Zoosystematics and Evolution 4»PErisPFr . 95 ( 2 ) 2019 ISSN 1435-1935 Zoosyst. Evol. 95 (2) 2019, 309-598 BERLIN Zoosystematics and Evolution A Bulletin of Zoology since 1898 Instructions for authors Scope Zoosystematics and Evolution is an international, peer-reviewed life science journal devoted to the diversity, systematics and evolution of all animal groups, except for insects. It publishes original research and review articles at all taxonomic levels. Its scope encompasses primary information from collection-relat¬ ed research, taxonomic descriptions and discoveries, revisions, annotated type catalogues, relevant aspects of the history of science, and contributions on new methods and principles of taxonomy and systematics. Articles whose main topic is ecolo¬ gy, functional anatomy, physiology, or ethology are only accept¬ able when of clear systematic or evolutionary relevance and perspective. 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Zoosystematics and Evolution A Bulletin of Zoology since 1898 Editor-in-Chief Thomas von Rintelen Museum fur Naturkunde, Leibniz-lnstitut fur Evolutions- und Biodiversitatsforschung Berlin, Germany phone: +49 (0)30-889140-8428 e-mail: thomas.vonrintelen@mfn.berlin Managing Editor Lyubomir Penev Pensoft Publishers, Sofia, Bulgaria phone: +359-2-8704281 fax: +359-2-8704282 e-mail: penev@pensoft.net Editorial Secretary Boryana Ovcharova Pensoft Publishers, Sofia, Bulgaria phone: +359-2-8704281 fax: +359-2-8704282 e-mail: journals@pensoft.net Editorial Board Vertebrata; Systematics Peter Bartsch - Museum fur Naturkunde, Berlin Amphibia Rate Brown - University of Kansas, Lawrence Decapoda; Taxonomy Sammy De Grave - Oxford University, Oxford Mollusca; Biogeography, Evolutionary Biology Matthias Glaubrecht - Oenter of Natural History, University of Hamburg, Hamburg Arachnida, Arthropoda; Taxonomy, Biodiversity & Oonservation Danilo Harms - Oenter of Natural History, University of Hamburg, Hamburg Pisces; Molecular Biology, Molecular Systematics; Population Genetics; Molecular Genetics Nicolas Hubert - Institut de Recherche pour le Developpement, Montpellier Arthropoda; Biodiversity & Oonservation, Molecular Biology, Taxonomy Martin Husemann - Oenter of Natural History, University of Hamburg, Hamburg Publisher Gastropoda; Freshwater, Terrestrial Martin Husemann - Australian Museum, Sydney Amphibia, Reptilia; Gonservation Biology, General Ecology, Taxonomy Johannes Penner - University of Freiburg, Freiburg Nematomorpha; Systematics, Marine, Taxonomy Andreas Schmidt-Rhaesa - Oenter of Natural History, University of Hamburg, Hamburg Invertebrata; Systematics Pavel Stoev - National Museum of Natural History and Pensoft Publishers, Sofia Orustacea; Freshwater Kristina von Rintelen - Museum fur Naturkunde, Berlin Mollusca Thomas von Rintelen - Museum fur Naturkunde, Berlin Zoosystematics and Evolution 2019. Volume 95. 2 Issues ISSN: 1435-1935 (print), 1860-0743 (online) Abbreviated keys title: Zoosyst. Evol. In Focus The cover picture shows Eulimacrostoma microsculpturata gen. nov. and sp. nov. See paper of de Souza LS, Pimenta AD Eulimacrostoma gen. nov., a new genus of Eulimidae (Gastropoda, Caenogastropoda) with description of a new species and reevaluation of other western Atlantic species Cover design Pensoft ^FENson : Zoosystematics and Evolution A Bulletin of Zoology since 1898 Content of volume 95 ( 2 ) 2019 Syaukani S, Thompson GJ, Yamasaki T, Othman AS, Muarrif S, Sarong MA, Djufri D, Eguchi K Taxonomy of the genus Longipeditermes Holmgren (Termitidae, Nasutitermitinae) from the Greater Sundas, Southeast Asia 309 Machado M, Guzati C, Viecelli R, Molina-Gomez D.Teixeira RA A taxonomic review of the crab spider genus Sidymella (Araneae, Thomisidae) in the Neotropics 319 Guimaraes EC, de Brito PS, Feitosa LM, Costa LFC, Ottoni FP A new cryptic species of Hyphessobrycon Durbin, 1908 (Characiformes, Characidae) from the Eastern Amazon, revealed by integrative taxonomy 345 Cunha CM, Rosenberg G Type specimens of Aplysiida (Gastropoda, Heterobranchia) in the Academy of Natural Sciences of Philadelphia, with taxonomic remarks 361 Boonmekam D, Krailas D, Gimnich F, Neiber MT, Glaubrecht M A glimpse in the dark? A first phylogenetic approach in a widespread freshwater snail from tropical Asia and northern Australia (Cerithioidea, Thiaridae) 373 Komai T, Chan T-Y, De Grave S Establishment of a new shrimp family Chlorotocellidae for four genera previously assigned to Pandalidae (Decapoda, Caridea, Pandaloidea) 391 de Souza LS, Pimenta AD Eulimacrostoma gen. nov., a new genus of Eulimidae (Gastropoda, Gaenogastropoda) with description of a new species and reevaluation of other western Atlantic species 403 Liu M, Li Z, Wei M Three new species of Macrophya Dahibom (Hymenoptera, Tenthredinidae) with a key to species of the Macrophya imitator group in Ghina 417 Cantalice KM, Martinez-Melo A, Romero-Mayen VA The paleoichthyofauna housed in the Coleccion Nacional de Paleontologia of Universidad Nacional Autonoma de Mexico 429 Salvador RB, Cavallari DC Taxonomic revision of the genus Hyperauiax Pilsbry, 1897 (Gastropoda, Stylommatophora, Odontostomidae) 453 Abstract & Indexing Information Biological Abstracts® (Thompson ISI) BIOSIS Previews® (Thompson ISI) Cambridge Scientific Abstracts (CSA/CIG) Web of Science® (Thompson ISI) Zoological Record™ (Thompson ISI) Zoosystematics and Evolution A Bulletin of Zoology since 1898 Content of volume 95 ( 2 ) 2019 de Souza Castanheira P, Baptista RLC, Dos Passos Pizzetti D, Teixeira RA Contributions to the taxonomy of the long-jav^/ed orb-v\/eaving spider genus Tetragnatha (Araneae, Tetragnathidae) in the Neotropical region, v\/ith comments on the morphology of the chelicerae 465 de Brito PS, Guimaraes EC, Carvalho-Costa LF, Ottoni FP A nevj species of Aphyocharax Gunther, 1868 (Characiformes, Characidae) from the Maracagume river basin, eastern Amazon 507 Valentas-Romera BL, Simone LRL, Mikkelsen PM, Bieler R Anatomical redescription of Cyrenoida floridana (Bivalvia, Cyrenoididae) from the Western Atlantic and its position in the Cyrenoidea 517 Yolanda R, Sawamoto S, Lheknim V A new species in the genus Heteromysoides (Crustacea, Mysida, Mysidae) from Songkhia Lagoon, southern Thailand 535 Sluys R The evolutionary terrestrialization of planarian flatworms (Platyhelminthes, Tricladida, Geoplanidae): a review and research programme 543 Albano PG, SchnedI S-M, Janssen R, Eschner A An illustrated catalogue of Rudolf Sturany’s type specimens in the Naturhistorisches Museum Wien, Austria (NHMW): Red Sea bivalves 557 Zoosyst. Evol. 95 (2) 2019, 309-318 | DOI 10.3897/zse.95.31636 4>yEnsPFr. BERLIN Taxonomy of the genus Longipeditermes Holmgren (Termitidae, Nasutitermitinae) from the Greater Sundas, Southeast Asia Syaukani Syaukani\ Graham J. Thompson^, Takeshi Yamasaki^, Ahmad Sohman Othman"^, Samsul Muarrif^, Muhammad Ali Sarong®, Djufri Djufri®, Katsuyuki Eguchi® 1 Biology Department, Faculty of Mathematics and Natural Science, Universitas Syiah Kuala, Darussalam 23111, Banda Aceh, Indonesia 2 Department of Biology, Western University, 1151 Richmond Street North, London N6A 5B7, Ontario, Canada 3 Systematic Zoology Laboratory, Department of Biological Science, Graduate School of Science, Tokyo Metropolitan University 1-1 Minami Osawa, Hachioji-shi, 192-0397 Tokyo, Japan 4 Molecular Ecology Laboratory, School of Biological Science, Universiti Sains Malaysia, 11800 Penang, Malaysia 5 Graduate School of Mathematics and Applied Sciences, Universitas Syiah Kuala, Darussalam 23111, Banda Aceh, Indonesia 6 Biology Department, Faculty of Education and Teacher Training, Universitas Syiah Kuala, Darussalam 23111, Banda Aceh, Indonesia http://zoobank org/D123A 73F-B056-46D3-AE9C- 74EAFD34F792 Corresponding author; Syaukani Syaukani (syaukani@unsyiah.ac.id) Academic editor: M/c/zae/OM ♦ Received 14 November 2018 ♦ Accepted 11 April 2019 ♦ Published 29 May 2019 Abstract More than 200 colonies of the genus Longipeditermes were collected in our field surveys across the Sundaland region of Southeast Asia from 1998 to 2014. Two species, L. kistneri Akhtar & Ahmad and L. logipes Holmgren, are recognized and redescribed with color photographs of the workers and major soldiers. We use variation in characters of soldier caste (head capsules, antennae, and pronotum) and worker caste (antennae and mandibles) to distinguish these two species. Longipeditermes kistneri seems to prefer high-altitude forests (above 1,000 m) and has so far been found exclusively in Java and Sumatra, while L. logipes seems to prefer lowland and swamp forests and is widespread in the Greater Sundas. Key Words Morphological characters, Nasutitermitinae, Open-air processional columns termites, species description Introduction A termite colony typically consists of a large number of workers and defensive soldiers together with a single king and a queen. This arrangement can, however, vary with seasonal cohorts of reproductive nymphs and dis¬ persive alates (Watson and Gay 1991; Pearce 1997). The castes within termite societies are morphologically and behaviorally specialized to perform particular tasks (Lee and Wood 1971; Grimaldi and Engel 2005) to the point where workers and soldiers are so specialized for non-re- productive helping and defensive tasks that they are con¬ sidered to be functionally sterile (Roisin 2000). The sex- uals, by contrast, are highly specialized for reproduction. As the largest subfamily among the higher termites (Termitidae), Nasutitermitinae consists of more than 550 species (Emerson 1955), which belong to more than 63 genera (Collins 1989). This subfamily probably originated in the Neotropical region during the Creta¬ ceous period (Emersonl955) and is characterized by a highly specialized defensive caste (Collins 1989) that has a modified rostrum or “nasus” (Deligne et al. 1981; Prestwich 1984). This subfamily was erected by Hare (1937), who emphasized the conspicuous prolongation on the front of the head, which is often accompanied by degeneration of mandibles and the concomitant devel¬ opment of a frontal gland. The development of the nasus is continuously variable among species within the Na- Copyright Syaukani Syaukani etai. This is an open access articie distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the originai author and source are credited. 310 Syaukani Syaukani et al.: Taxonomy of the genus Longipeditermes sutitermitinae from mild prolongation, as in Syntermes, to pronounced prolongation as in Subulitermes, Convex- itermes, and Nasutitermes. Longipeditermes, in the Nasutitermitinae, is a small genus with only two species so far recorded. These are L. longipes and L. kistneri from the Sundaland area (Gath- orne: Hardy 2001; Hoare and Jones 1998). Longipediter¬ mes is, however, well known for its remarkable foraging habit. Workers and soldiers form open-air processional columns on the ground, shrubs, and tree trunks typical¬ ly inside forests (Gray and Dhanarajan 1974; Hoare and Jones 1998; Miura and Matsumoto 1998; Takematsu et al. 2013; Syaukani et al. 2016). The foraging activities of both species seem to be greatest during the day (Hoare and Jones 1998), as evidenced by workers carrying packed balls of organic matter between their mandibles and returning to the nest. In the course of our long-term inventory and taxonom¬ ic research on termites in Southeast Asia, more than 200 colonies of Longipeditermnes have been sampled from different habitats and altitudes across the Greater Sunda. In this paper, the two species of the genus are redescribed and illustrated using newly obtained nest series. Informa¬ tion on their life history is provided. Materials and methods We examined 216 colonies of Longipeditermes from vari¬ ous habitats and altitudes across the Greater Sunda (Ta¬ ble 1; Fig. 1 ). The specimens examined are deposited at the Museum Zoologicum Bogoriense (MZB), Cibinong, Indonesia; the Natural History Museum, London (UK); and the Universitas Syiah Kuala (Biology Department), Darussalam, Banda Aceh (Indonesia). The head, body (in profile), pronotum, and antenna of the soldier caste (preserved in 70% ethanol) were photo¬ graphed using a digital microscope (KEYENCE HFVH- 8000, Osaka). From these images, multifocused montag¬ es were produced using Helicon Focus v. 6.2.2 (Helicon Soft Etd, Kharkov, Ukraine). Morphological terms and measurement characters follow those of Roonwal and Chhotani (1989), Sands (1998), Tho (1992), and Syau¬ kani et al. (2011). We measured the characters (in mil¬ limeters): head capsule length including nasus (HEN), head capsule length excluding nasus (HE), nasus length (NE), nasus index = NE/HE, maximum head width at anterior part (HWA), maximum head width at posterior part (HWP), maximum height of head capsule excluding postmentum (HH), pronotum length (PE), and pronotum width (PW). Results Taxonomic accounts Genus Longipeditermes Holmgren, 1913 Soldier. Bimodal in its size distribution. Head capsule pale brown to blackish; antenna much paler than head capsule in coloration, with the basal segments (first and second) generally darker than the remaining segments; pronotum paler than or as pale as head capsule; abdom¬ inal tergites pale brown to dark sepia brown; coxae yel- p o o p Q I/) .Acefi Pf wince' ovine 0 # Kayan Mertarang# Betung Keiinun* iaais» East Kalimantan Province* ilimantan Province* BanloUIn f'arara'rfcn* BeKil Suharto* GunuA^’alung* Bitklt Tangkrling* BengKrilu Prewce* Lampung Province* BuKil Banisfln 'BuJcffTinl Gunurvi Pargantlaran* 100’0'0"E 105“0'0'E 1t5’0'0"E 120°0'0'E o a «• o W ,p Q Figure 1. Distribution of Longipeditermes longipes (yellow dot) and L. kistneri (red dot) on the Greater Sundas, Southeast Asia. zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 309-318 311 lowish to pale brown; femora yellow to brown; tibiae pale yellow to yellow. In dorsal view head capsule excluding rostrum pear-shaped to somewhat triangular, weakly constricted behind antennal sockets; its posterior mar¬ gin weakly to strongly convex; dorsal outline in profile weakly to strongly concave; rostrum excluding the apex somewhat cylindrical rather than conical. Antenna long, with 14 segments; segment at least twice as long as 4*. Mandible relatively long, with sharp apical processes. Legs very long. Worker. Monomorphic but showing size variation. Pale brown to blackish. Antenna with 15 segments, with the basal segments darker than the following ones. Left mandible with 3'^ marginal tooth weakly to moderately protruding from cutting edge, and 4* partially hidden be¬ hind molar prominence. Remarks. In Longipeditermes, the soldier caste is dimorphic. The caste is subdivided into major and mi¬ nor soldiers that differ markedly in size (Thapa 1981; Tho 1992; Gathorne-Hardy 2001; Miura and Matsumoto 1998) and slightly in coloration; the former being darker than the latter. The major soldier has more characters use¬ ful for species recognition and identification. Two varied species, L. longipes and L. kistneri, were recognized in our collection (Table 1), while no additional species were found by our morphology-based examination. The two species are relatively easily distinguished from each oth¬ er by a combination of characters given in Table 2. These characters are re-described below in detail. Longpeditermes longipes (Haviland, 1898) Termes longipes Haviland 1898: 439-440. Type locality: Perak, Malay Peninsula. Syntypes examined. Eiitermes longipes Wasmann 1902: 131. Eutermes {Longipeditermes) longipes: Holmgren 1902: 215-217; John 1925:406. Longipeditermes longipes: Snyder 1949: 317; Ahmad 1958: 126; Tho 1992: 180-181: Miura and Matsumoto 1998: 179-189 (ecology); Hoare and Jones 1998: 1357-1366. Longipeditermes mandibidatiis Thapa 1981: 349-352. Sabah, Borneo. Synonymized by Hoare and Jones 1998: 1357-1366. Materials examined. Sumatra. SYK1998 & 1999-L- 1115, 1117, 1120, 1121, 1124, 1130, 1133, 1136, 1138, 1139,1141, 1143, 1147, 3025: soldiers and workers from Table 1. Summary of collection sites. Site Vegetation type Altitude (m) The Leuser Ecosystem, Sumatra Aceh Province (outside the Leuser Ecosystem), Sumatra BatangGadis National Park, Sumatra North Sumatra Province (except the Leuser Ecosystem) Kerinci Seblat National Park, Sumatra West Sumatra Province (outside Kerinci Seblat N. P), Sumatra Jambi Province ( outside Kerinci Seblat N.P), Sumatra Bukit Barisan Selatan National Park, Sumatra Bengkulu Province ( outside Bukit Barisan SelatanNational Park), Sumatra Lampung Province ( outside Bukit Barisan SelatanNational Park), Sumatra Endau Ronpin National Park, Malay Peninsula Teluk Bahang National Park, Malay Peninsula Bukit Tangkiling Nature Preserve, Borneo Pararawen Nature Preserve, Borneo Barito Ulu, Borneo Gunung Palung National Park, Borneo Betung Kerihun National Park, Borneo West Kalimantan Province ( outside Betung Kerihun NP) Borneo East Kalimantan Province ( outside Gunung Palung NP) Borneo Bukit Suharto, Borneo North Kalimantan Province ( outside Kayan Mentarang NP), Borneo Gunung Halimun National Park. West Java Pangandaran Natural Reserve, West Java Bukit Lengkong, West Java Lowland & subalpine forests Protected & unprotected 50-1400 Lowland forest 20-600 Lowland forest 800-1200 Lowland & subalpine forests 20-1100 Lowland & subalpine forests 300-1350 Lowland forest 50-400 Lowland forest 50-600 Lowland forest 50-600 Lowland forest 100-700 Lowland forest 10-300 Lowland forest 150-600 Lowland forest 5-200 Lowland forest 25-170 Lowland forest 50-350 Lowland & subalpine forests 900-1200 Lowland forest 50-300 Lowland forest 400-850 Lowland forest 50-650 Lowland forest 150-500 Lowland forest 40-300 Lowland & subalpine forests 500-1200 Lowland & subalpine forest 800-1350 Lowland 10-40 Lowland & subalpine forest 800-1250 Table 2. Summary of morphological characters for the Longipeditermes based on major soldiers and workers. Species Soldier Worker Coloration of head capsule Rostrum Anterior margin of pronotum Antennae Left mandible L. longipes Sepia brown to blackish Apical 2/3 lighter & basal 1/3 darker Nearly straight 4*^ segment longer than 5‘^ , 3^"'marginal tooth moderately protruding from cutting edge L. kistneri Pale brown to dark brown Apical 2/3 darker and basal 1/3 lighter Strongly indented in the middle 4*^ to 6'^ segments almost equal in length 3^^^ marginal tooth weakly protruding from cutting edge zse.pensoft.net 312 Syaukani Syaukani et al.: Taxonomy of the genus Longipeditermes Figures 2-9. (SYK2006-KSNP-0011). Longipeditermes longipes (major soldier & major worker). Soldier (2-4, 7, 8) and worker (5, 6, 9). Habitus in profile (2), head in dorsal view (3), head in profile (4), left (5) and right (6) mandibles, pronotum (7), antennae (8, 9). Scale bars; 0.6 mm (2), 0.3 mm (3, 4), 0.1 mm (5- 9), 1.7 mm (6). undisturbed forest, 300-500 m altitude, Ketambe, South¬ east Aceh. SYK1998-L-3005, 3010: soldiers and workers from disturbed forest, 450 m altitude, Lokop, East Aceh, Aceh. SYK-L-1148, 3006: soldiers and workers from disturbed forest, 80 m altitude, Soraya, Singkil, Aceh. SYK1998 & 2000-L-1126, 1127, 1131,1199, 1145, 3007, 3008: soldiers and workers from undisturbed forest, 150- 350 m altitude, Bukit Lawang, Langkat, North Sumatra. SYK1998-L-3009, 3022, 3024: soldiers and workers from disturbed forest, 200 m altitude, MRT Logging Concession, South Aceh. SYK1999 & 2001-L-1116, 1123, 1125, 1128, 1134, 1135, 1137, 1144, 1146: soldiers and workers from disturbed forest, 50 m altitude, Sekundur, Langkat, North Sumatra. SYK1999-L-1112,1118,1129,1132,1140,1142: soldiers and workers from undisturbed forest, 200^00 m altitude, Bengkung, Southeast Aceh. SYK2006-AL-0104: Soldiers and workers from disturbed forest, 50 m altitude, Maestong, Batang Hari, Jambi. SYK2006-KSNP-0011, 0019, 0080, 0091, 0093, 0095, 0096, 0097, 0104, 0206: soldiers and workers from the undisturbed forest, 300 m in altitude, Sungai Manau, Merangin, Jambi. SYK2006- AL-0100, 0101, 0102, 0103: soldiers and workers from disturbed forest, 50 m altitude, Maestong, Batang Hari, Jambi. SYK2007-LP-0019: soldiers and workers from undisturbed forest, 1350 m altitude, Sumber Jaya, Kota Bumi, Lampung. Java. SYK2001-HL-067, 072: soldiers and workers from protected forest, 1,450 m altitude, Hali- mun NP, West Java. SYK2006-PD-0011, 048: soldiers and workers from protected forest, 10 m altitude, Pan- gandaran Nature Reserve. West Java. Malay Peninsula. zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 309-318 313 Figures 10-17. (SYK2006-KSNP-0011). L. longipes (minor soldier & minor worker). Soldier (10-12, 15) and worker (13, 14, 17). Habitus in profile (10), head in dorsal view (11), head in profile (12), left (13) and right (14) mandibles, pronotum (15), antennae (16, 17). Scale bars: 0.6 mm (10), 0.3 mm (11, 12), 0.1 mm (13- 17), 1,7 mm (15). SYK2009-ER-084, 085, 086, 087, 088: soldiers and work¬ ers from protected forest, 50 m altitude, Endau Rompin NR SYK2011-TB-011, 012, 013, 014: soldiers and work¬ ers from protected forest, 10 m altitude, Teluk Bahang NR Borneo. SYK2014-BT-0054: soldiers and workers from protected secondary forest, 35 m altitude, Ralangkaraya, Central Kalimantan. SYK-2014-R-0012; soldiers and workers from undisturbed forest, 220 m altitude. North Barito, Central Kalimantan. SYK-2014-R-0024: soldiers and workers from undisturbed forest, 270 m altitude. North Barito, Central Kalimantan. Soldier (Figs 2-A, 7, 8, 10-12, 15, 16). Bimodal con¬ tinuum in size. Major soldier (Figs 2-A, 7, 8). Head cap¬ sule sepia brown to blackish, in dorsal view somewhat triangular, very weakly constricted behind antennal sock¬ ets, with posterior margin roundly convex; dorsal outline (including nasus) in lateral view moderately concave; ros¬ trum feebly bicolored, with apical 2/3 lighter and basal 1/3 darker. Mandible with short to long apical processes in dorsal view. Antenna with 14 segments, much paler than head capsule; antennal segment darker than the follow¬ ing segments, which are uniformly colored; 3'^'* segment approximately 1.5 times as long as 4*; 5* slightly short¬ er than 4*; 6*-14* gradually shortening toward the apex. Rronotum in dorsal view paler than head capsule; anterior margin nearly straight; posterior margin weakly indented in the middle. Coxae pale brown to brown; femora pale brown to yellow; tibiae pale yellow. Abdominal tergites pale brown to dark sepia brown. Measurements and index (20 major soldiers from 10 colonies): HEN 2.30-2.71 mm; zse.pensoft.net 314 Syaukani Syaukani et al.: Taxonomy of the genus Longipeditermes HL 1.57-1.62 mm; NL 0.72-0.77 mm; NL/HL 0.47-0.51; HWA0.81-0.98 mm; HWP 1.87-1.92 mm; HH 1.22-1.32 mm; PL 0.37-0.40 mm; PW 0.75-0.78 mm. Worker (Figs 5, 6, 9, 13, 14, 17). Monomorphic but showing size variation. Worker (large-sized) (Figs 5, 6, 9). Blackish brown to blackish. Left mandible with apical tooth shorter than marginal tooth; 3'^‘^ marginal moderately protruding from cutting edge; 4* completely hidden, scarcely visible behind molar prominence. Right mandible with posterior edge of 2"^ marginal tooth nearly straight or weakly concave; the inner layer of molar plate very moderately convex; notch moderately developed. Antenna with 15 segments, whitish yellow; and 2"'* segments darker than the following segments; 3'^‘^ clearly longer than 4*; 4* longer than 5*, 6*-15* gradually short¬ ening toward the apex. Distribution. Sumatra, Peninsular Malaysia, Java (new record), and Borneo. Longipeditermes kistneri Akhtar & Ahmad, 1985 Longipeditermes kistneri Ahktar and Ahmad 1985: 215-217. Bogor, West Java. Material examined. SYK1998 & 1999-L-1098, 2001, 3011: soldiers and workers from the undisturbed for¬ est, 1100-1400 m altitude, Kemiri Mountain, Southeast Aceh, Aceh, Sumatra. Figures 18-25. (SYK1999-L-3011). L. kistneri (major soldier and major worker). Soldier (18-20, 23, 25) and worker (21, 22, 25). Habitus in profile (18), head in dorsal view (19), head in profile (20), left (21) and right (22) mandibles, pronotum (23), antennae (24, 25). Scale bars: 0.6 mm (18), 0.3 mm (19, 20), 0.1 mm (21, 22, 24, 25), 1.7 mm (23). zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 309-318 315 Figures 26-33. (SYK1999-L-3011). L. kistneri (minor soldier and minor worker). Soldier (26- 28, 32) and worker (29, 30, 33). Habitus in profile (26), head in dorsal view (27), head in profile (28), left (29) and right (30) mandibles, pronotum (31), antennae (32, 33). Scale bars; 0.6 mm (26), 0.3 mm (27, 28), 0.1 mm (29, 30, 32, 33), 1.7 mm (31). Soldier (Figs 18-20, 23, 24, 26-28, 31, 32). Bimodal continuum in size. Major soldier (Figs 18-20, 23, 24). Head capsule pale to dark brown, in dorsal view some¬ what triangular, very weakly constricted behind antennal sockets, with posterior margin roundly convex; dorsal outline (including nasus) in profile moderately concave; rostrum feebly bicolored, with apical 2/3 darker and basal 1/3 paler. Mandible with short to long apical processes. Antenna with 14 segments, much paler than head capsule; V'- and 2"‘^ antennal segments darker than the following segments, which are uniformly colored; 3'^'* approximately 1.5 times as long as 4*; 5* slightly shorter than 4*; 6*-14* gradually shortening toward the apex. Pronotum in dorsal view as pale as or paler than head capsule; anterior margin indented in the middle; posterior margin weakly indented in the middle. Coxae and femora yellowish to pale brown; tibiae pale yellow to yellow. Abdominal tergites brown to dark sepia brown. Measurements and index (20 major sol¬ diers from 10 colonies): HLN 2.22-2.43 mm; HL 1.62- 1.68 mm; NL 1.02-1.12 mm; NL/HL 0.66-0.73 ; HWA 0.86-0.90 mm; HWP 1.63-1.71 mm; HH 0.98-1.04 mm; PL 0.40-0.44 mm; PW 0.77-0.82 mm. Worker (Figs 21,22,25,29,30,33). Monomorphic but showing size variation. Worker (large-sized) (Figs 21,22, 25). Body pale brown to brown. Left mandible with apical tooth shorter than marginal tooth; 3'^^ marginal weakly protruding from cutting edge; 4* not completely hidden behind molar prominence. Right mandible with posterior edge of 2"^ marginal tooth nearly straight; the inner layer of the molar plate very weakly concave; notch moderately zse.pensoft.net 316 Syaukani Syaukani et al.: Taxonomy of the genus Longipeditermes to strongly developed. Antenna with 15 segments, with and 2"^ segments slightly darker than the following seg¬ ments; clearly longer than 4*; 4*-6* nearly equal in length; 7*-15* gradually shortening toward the apex. Distribution. Sumatra and Java. Discussion Longipeditermes species are distinguished from those of the other groups of Southeast Asian nasutitermitine ter¬ mites by forming open-air processional foraging columns (e.g., Hospitalitermes and Lacessititermes) and by having major and minor soldiers that both have long legs and a long rostrum. No information is available for the ratio of the largest and smallest soldiers in typical colonies, but minor soldiers appear to be more numerous (Figs 34, 35). The soldier does not exhibit a clear dimorphism, but a bimodal continuum in size. Longipeditermes kistneri was only collected from Su¬ matra during our intensive surveys in the Greater Sundas, although the type locality is Java. Nine colonies were found in protected forests above 1,000 m in altitude. Four nests were located among buttresses and roots of big trees (under the ground covered with decayed leaf litter and small decayed branches). This nesting habit makes it dif¬ ficult to estimate the sizes of their colonies. Both soldiers and workers forage on the surface of the ground. The size of foraging columns is smaller in L. kistneri than in L. lon- gipes. Their pale- to dark-brown body color and smaller columns make it difficult to find colonies of L. kistneri. Longipeditermes longipes seems to prefer lowland and swamp forests and is widespread in the Greater Sundas. Nests of L. longipes are usually constructed among but¬ tresses of trees (Fig. 36) and in dead or decayed stumps, with decayed wood debris and leaves (Fig. 37). Several subterranean nests of this species were found in Lambir Hills National Park (Sarawak, Borneo) by Miura and Mat- Figures 34, 35. Foraging column of Longipeditermes longipes on the forest floor in Southeast Asia. Workers collecting and trans¬ porting food source from a decayed branch (34) and leaf litter (35), soldiers protecting the worker from predators such as Compono- tus gigas (Latreille). Figure 36. Nest entrances (red arrows) of subterranean nest of Longipeditermes longipes found in SE Asia tropical forests. sumoto (1998). Underground nests are sometimes made from consisting of soil and litter. This species forages on the ground during the day. However, Hoare and Jones (1998) reported foraging activity at night, during which no food balls were being carried. We also found foraging columns at night; the columns consisted of limited num¬ bers of workers carrying food balls to the nests. In typical processional foraging columns, numerous soldiers guard workers. Foraging ecology and diet of L. longipes have been poorly studied, but we know that this species feeds on leaf litter (Miura and Matsumoto 1998) and in the Pa- soh Forest Reserve on the Malay Peninsula it prefers to consume older leaf litter rather than newly fallen leaves (Matsumoto and Abe 1979). We observed foraging col¬ umns consuming a mixture of bark, twigs, and decayed wood in forests of Sumatra and the Malay Peninsula. zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 309-318 317 Figure 37. Above-ground nest of L. longipes constructed in and around a dead tree stump {Shorea sp.). The cavity was fulfill with a mixture of soil, debris and litter. Conclusion In general, from the current study, we conclude that only two species of Longipeditermes {L. longipes Holmgren and L. kistneri Akhtar & Ahmad) have so far been col¬ lected in the Greater Sundas. Long legs and size dimorph¬ ism of soldier caste can help to separate this genus from other nasute termites in Southeast Asia. The condition of rostrum, head capsule, pronotum, and antennae deter¬ mine the soldier caste, while the condition of antennae and left mandibles of worker caste serve to distinguish these two species. Longipeditermes kistneri seems to pre¬ fer high-altitude forests and has been found exclusively from Java and Sumatra, while L. longipes seems to pre¬ fer lowland and swamp forests and is widespread in the Greater Sundas. Acknowledgements We thank the Termite Research Group (Universitas Sy- iah Kuala, Indonesia), Sugesti, M. Rapi, Tarmizi, M. Isa, Usman, and Mat Plin (Leuser Development Program, Indonesia), Erwin Widodo (Conservation International), Epal (WWF, Indonesia), Dolly Priatna and Adnun Salam- pesi (Zoological Society of Eondon), F.X Susilo, I Gede Swibawa, and Rahmat Pranoto (Eampung University), Idris Ghani and Fazli Rahim (UKM, Malaysia), Rosichon Ubaidillah and Wara Asfiya (MZB, Bogor), Teguh Prib- adi (PGRI Palangkaraya University, Indonesia), and M. Shahril (USM, Malaysia) for assistance in the field and laboratory. We are grateful to Seiki Yamane (Kagoshima University Museum, Japan) for support, Paul Eggleton and David Jones (Natural History Museum, UK) and staff at the Museum Zoologicum Bogoriense (MZB, In¬ donesia) for allowing the first author to examine the type material. We thank to staff in the Forestry Department in Indonesia for their help and assistance during field sur¬ veys. This work was financially supported by the Feuser Development Program (FDP 1998-2001), Nagao Envi¬ ronmental Foundation (NEF 2006), the Ministry of Re¬ search, Technology and Higher Education RG to Syau- kani (International Research Collaboration and Scientific Publication 2012-2018), Syiah Kuala University (Pro¬ fessor Candidate Grant 2015, H-index Publication Grant 2016, International Conference Support 2016 (DITJEN RISBANG, the Ministry of Research, Technology and Higher Education, and the Zoology Eaboratory Grant 2018). Eguchi’s research activities were partly supported by Asahi Glass Foundation (Eeader: Katsuyuki Eguchi; FY2017-FY2020). References Akhtar MS, Ahmad M (1985) A new nasute termite from Java (Isop- tera: Termitidae: Nasutitermitinae). Pakistan Journal of Zoology 17: 215-217. Collins NM (1984) The termites (Isoptera) of the Gunung Mulu Na¬ tional Park, with a key to genera known from Sarawak. Sarawak Museum Journal 30: 65-87. Collins NM (1989) Termites. In: Lieth H, Werger MJA (Eds) Tropical Rain Forest Ecosystems. Biogeographical and Ecological Studies. Elsevier, Amsterdam, 455-471. https://doi.org/10.1016/B978-0- 444-42755-7.50032-8 Deligne J, Quennedy A, Blum MS (1981) The enemies and defense mechanisms of termites. In: Hermann HR (Ed.) Social Insects (Second Edition). Academic Press, New York, 1-76. https://doi. org/10.1016/B978-0-12-342202-6.50008-3 Emerson AE (1955) Geographical origins and dispersions of termite genera. Fieldiana, Zoology 37: 465-521. https://doi.org/10.5962/ bhl.title.2783 Gray B, Dhanarajan G (1974) Processional trails of the black ter¬ mite Longipeditermes Longipes (Haviland) (Isoptera: Termi- tisae). Insectes Sociaux 21: 151-155. https://doi.org/10.1007/ BF02222939 Gathorne-Hardy F (2001) A review of the South-East Asian Na¬ sutitermitinae (Isoptera: Termitidae), with descriptions of one new genus and a new species and including a key to the gen¬ era. Journal of Natural History 35: 1486-1506. https://doi. org/10.1080/002229301317067647 Grimaldi D, Engel MS (2005) Evolution of the Insects. Cambridge Uni¬ versity Press, New York, 755 pp. Haviland GD (1898) Observations on termites; with description on new species. Journal of the Einnean Society, Zoology 26: 358-442. https://doi.org/10.1111/j. 1096-3642.1898.tb00405.x zse.pensoft.net 318 Syaukani Syaukani et al.: Taxonomy of the genus Longipeditermes Hare L (1937) Termite phylogeny as evidenced by soldier mandible development. Annals of the Entomological Society of America 37; 459-486. https://doi.org/10.1093/aesa/30.3.459 Hoare A, Jones DT (1998) Note on the foraging behaviour and tax¬ onomy of the Southeast Asian termite Longipeditermes longipes (Termitidae; Nasutitermitinar). Journal of Natural History 32; 1357- 1366. https://doi.Org/10.1080/00222939800770681 Holmgren N (1912) Termitenstudien. 3. Systematik der Termiten. Die Familien Mastotermitidae. KungligaSvenskaVetensskapakademiens Handlingar 48; 1-166. Holmgren N (1913) Termitenstudien. 4. Versuch einer systematichen Monographic der Termiten der orientalischen Region. Kunglig- aSvenskaVetensskapakademiens Handlingar 50: 1-276. John O (1925) Termiten von Ceylon, der Malayaischen Halbinsel, Su¬ matra, Java und den Aru- Inseln. Treubia 6: 360-419. Lee KE, Wood TG (1971) Termites and soils. Academic Press, London and New York, 1-25. https://doi.org/10.2307/3758095 Matsumoto T, Abe T (1979) The role of termites in an equatorial rain forest ecosystem of West Malaysia. Oecologia 38: 261-274. https:// doi.org/10.1007/BF00345187 MiuraT, Matsumoto T (1998) Open-air litter foraging in the nasute termite Longipeditermes longipes (Isoptera; Termitidae). Journal of Insect Behavior 11: 179-189. https://doi.Org/10.1023/A:1021039722402 Pearce MJ (1997) Termites; Biology and Pest Management. CAB Inter¬ national, Willingford, 172 pp. Roisin Y (2000) Diversity and evolution of caste patterns. In; Abe T, Bignell DE, Higashi M (Eds) Termites: Evolution, Sociality, Sym¬ biosis, Ecology. Kluwer Academic, Dordrecht, 95-120. https;//doi. org/10.1007/978-94-017-3223-9_5 Roonwal ML, Chhotani OB (1989) The Fauna of India and the Adjacent Countries. Zoological Survey of India, Calcuta, 672 pp. Sands WA (1998) The Identification of Worker Caste of Termite from Soil of Africa and the Middle East. CAB International, Wallingford, 500 pp. Snyder TE (1949) Catalog of the Termites (Isoptera) of the World. Smithsonian Miscellaneous Collection 112; 1-374. https://reposito- ry. si. edu/handle/10088/22862 Syaukani S, Thompson GJ, Yamane S (2011) Hospitalitermes krishnai, a new nasute Termite (Nasutitermitinae, Termitidae, Isoptera), from southern Sumatra, Indonesia. ZooKeys 148: 161-169. https;//doi. org/10.3897/zookeys. 148.1768 Syaukani S, Thompson GJ, Zettel H, Pribadi T (2016) A new species of open-air processional column termite, Hospitalitermes nigrianten- nalis sp. n. (Termitidae), from Borneo. Zookeys 554: 27-36. https;// doi.org/10.3897/zookeys.554.6306 Takematsu Y, Kambara K, Yamaguchi T, Mitsumaki K (2013) Spatial segregation of four coexisting processional termites (Termitidae: Nasutitermitinae) in tropical rainforest. Entomological Science 16; 355-359. https://doi.org/10.111 l/ens. 12015 Thapa RS (1981) Termites of Sabah. Sabah Forest Record, Sandakan 12: 1-374. Tho YP (1992) Termites of Peninsular Malaysia. Malayan Forest Re¬ cords, Kuala Lumpur 36: 1-224. Wasmann E (1902) Termiten, termitophilen und myrmekophilen. Gessamelt auf Ceylon von Dr. Horn. Zoologische Jahrbuecher Abteilung fuer Systematik Oekologie und Geographic der Tiere 17: 99-164. https://www.zobodat.at/pdf/Zoologische-Jahrbuech- er-Syst_17_0099-0164.pdf Watson JAL, Gay FJ (1991) Isoptera (Termites). In: Naumann D, Came PB, Lawrence JF, Nielsen ES, Spradbery JP, Taylor RW, Whitten MJ, Littlejohn MJ (Eds) The Insects of Australia - A Textbook for Students and Research Workers. Vol. 1. Melbourne University Press, Australia, 330-347. zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 319-344 i DOI 10.3897/zse.95.34958 4>yEnsPFr. BERLIN A taxonomic review of the crab spider genus Sidymella (Araneae, Thomisidae) in the Neotropies Miguel Machado^ Catherine Guzati\ Rafaela Vieeelli\ Diana Molina-G6mez\ Renato Augusto Teixeira^ 1 Laboratdrio de Aracnologia, Faculdade de Biociencias, Pontificia Universidade Catdlica do Rio Grande do Sul (PUCKS), Avenida Ipiranga 6681, Porto Alegre, RS, Brazil http://zoobank.org/lAC7C423-2E9A-42B0-AD01-902985656BE6 Corresponding author: Miguel Machado (machaclom.arachno@gmail) Academic editor: Z)a«/7o♦ Received 28 March 2019 ♦ Accepted 2 May 2019 ♦ Published 29 May 2019 Abstract Four Neotropical species of Sidymella Strand, 1942, S. furcillata Keyserling, 1880, S. longispina (Mello-Leitao, 1943), S. lucida (Keyserling, 1880), and S. kolpogaster (Lise, 1973) are redescribed from both sexes. The holotype of S. nigripes (Mello-Leitao, 1947) is lost and this taxon is considered a species inquierenda. Sidymella obscura (Mello-Leitao, 1929), S. parallela (Mello-Leitao, 1929), and S. spinifera (Mello-Leitao, 1929) are all nomina dubia. Two new species are described: Sidymella excavata sp. nov. (males and females) and S. marmorata sp. nov. (female). Key Words crab spiders, morphology, new records, Stephanopinae, Stephanopis Introduction Crab spiders (Araneae, Thomisidae) are distributed worldwide but the highest diversity is found in tropical regions (WSC 2019). The group has been studied in many recent phylogenetic works, but its relationships are still being discussed (Benjamin et al. 2008; Benjamin 2011; Ramirez 2014; Wheeler et al. 2017) and broader rela¬ tionships among basal thomisids such as the subfamily Stephanopinae remain weakly supported and unstable (Ramirez 2014). The presence of cheliceral teeth, which was previously considered as a synapomorphy for this group (Ono 1988), was recovered as a plesiomorphy by Benjamin (2011), and this subfamily remains as the most controversial and the least studied group in Thomisidae; it has many genera in need of revision and a considerable number of species yet to be described (Benjamin 2011). Based on the work of Mello-Leitao (1929), subsequent efforts were made to update the taxonomy of some Neo¬ tropical stephanopines (Lise 1973, 1981; Bonaldo and Lise 2001; Machado et al. 2015, 2017; Silva-Moreira and Machado 2016; Prado et al. 2018). However, many gen¬ era are still known only from the original descriptions and poor diagnoses, and the accurate identification of many species is practically impossible. The genus Sidymella Strand, 1942 is a prime example of such difficulties. The genus has a disjunct Gondwanan distribution, with 11 described species occuring in Austra¬ lia and New Zealand whilst 10 are found in the Neotropies (WSC 2019). Sidymella is currently defined by a convex prosoma, both anterior and posterior eye rows recurved, anterior tibiae and metatarsi (I and II) with stout and spin- iform macrosetae, and opisthosoma posteriorly bifurcated (Strand 1942; Mello-Leitao 1929; Lise 1973). Although the Neotropical Sidymella have been revised by Lise (1973), this author focused on somatic characters to de¬ scribe and diagnose the species, neglecting both external and internal structures of female genitalia and the position, shape, and size of palpal apophyses of males. Therefore, the present paper provides a taxonomic review of the Neo- Copyright Miguel Machado etal. This is an open access articie distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the originai author and source are credited. 320 Miguel Machado et al.: Taxonomic review of Neotropical Sidymella tropical Sidymella, where the males of S. furcillata and S. longispina are described for the first time, two new species added to the genus, and new synonyms established. Methods The examined specimens belong to the Museo Argentine de Ciencias Naturales “Bernardino Rivadavia”, Buenos Aires (MACN, Martin Ramirez), Museu de Ciencias e Tecnologia da Pontificia Universidade Catolica do Rio Grande do Sul, Porto Alegre (MCTP, A. A. Lise), Museu de Ciencias Naturais da Fundagao Zoobotanica do Rio Grande do Sul, Porto Alegre (MCN, R. Ott), and the Mu¬ seum of Comparative Zoology of Harvard, Cambridge (MCZ, G. Giribet and L. Liebensperger). The terminology for somatic and copulatory structures follows Machado et al. (2018). Photos of copulatory struc¬ tures were taken with a Multipurpose Zoom Microscope Leica M205A with a digital camera, and scanning electron microscopy were made with a Philips XL 30 Field Emis¬ sion ESEM from the Centro de Microscopia e Microanalis- es (CEMM) of the Pontificia Universidade Catolica do Rio Grande do Sul (PUCRS). All measurements were taken in millimeters. Morphology abbreviations: AEE—anterior lateral eyes, AME—anterior median eyes, MOQ—median ocular quadrangle, PEE—posterior lateral eyes, PME— posterior median eyes, RTA— retrolateral tibial apophysis. Results Sidymella Strand, 1942 Sidyma Simon, 1895; 1056. Type species: Stephanopis lucida Keyser- ling, 1880; by original designation and monotypy; Borland 1913: 95, pi. 9, figs 51-56. Preoccupied by Walker (1856) in Lepidoptera. Sidymella Strand, 1942: 399 (generic replacement name). Diagnosis. Sidymella is similar to Coenypha Simon, 1895 and some species of Stephanopis (e.g. S. antenna- ta, S. ditissima, and S. nodosa) and these species share a male palp with a long, thin and curled embolus, well-de¬ veloped pars pendula, and a retrolateral tibial apophysis with a short basal branch (Figs lA, B, IIC-F, 13D); the epigynes have a septum formed by the posterior folds of the epigynal plate and long and coiled copulatory ducts (Figs 1C, D, lOC-F). However, Sidymella species can be recognized and distinguished from these genera by hav¬ ing a bifid opisthosoma (Fig. IE, F), presence of spini- form macrosetae on the mesial surface of femora I and above the AEE (Fig. lOA, B), the epigyne have a single pair of walnut-shaped spermathecae compartmentalized in several smaller chambers, with accessory glandular heads (Figs 4D, F, 6D, F, 8D, F, lOD, F, 12D, F, 13D, F) while the male palp has a tibial trichobothrium and the RTA have a nodose surface (Fig. lA, B) unlike other Stephanopinae which present a grooved RTA (with paral¬ lel creases on its surface). Description. Small spiders (total length 3.00-3.96 in males, 4.68-7.93 in females) with slight sexual size di¬ morphism, presenting evidence of predominant green colouration in vivo (Fig. IE, F). Prosoma longer than wide, covered with short and conical setae. Both ante¬ rior and posterior eyes disposed in two recurved rows; AEE almost two times larger than the other eyes; ocular macrosetae may be present only above the AEE or on the MOQ area; sternum heart-shaped with concave anteri¬ or border, and clypeus with a pair of macrosetae; labi¬ um trapezoidal and endites with rounded edges, longer than wide. Opisthosoma with two projections of different sizes, shapes and angles in relation to the opisthosomal axis, varying from rounded and short ones, disposed hor¬ izontally/posteriorly, or long, acute and vertically orient¬ ed (Figs 2A-F, 3A-F). Eeg formula: 1-2-4-3; anterior tibiae and metatarsi (I and II) ventrally armed with stout macrosetae. Epigynum with membranous and hyaline copulatory ducts, long spermathecae with many cham¬ bers or with a single median constriction (Fig. 1OD); male palp with RTA rounded or truncated, discoid tegulum and ribbon-like embolus and (Fig. lOC-F). Composition. Six Neotropical species: Sidymella excav- ata sp. nov., S. furcillata (Keyserling, 1880), S. longispi¬ na (Mello-Eeitao, 1943), S. lucida (Keyserling, 1880), S. marmorata sp. nov., and S. kolpogaster (Else, 1973); 11 additional species from the Australasian region that are not within the scope of the present study. Distribution. Ecuador, Colombia, Peru, Brazil, Argenti¬ na, and Uruguay (Fig. 15). Sidymella excavata Machado & Guzati, sp. nov. http://zoobank.org/4D59E033-C71C-4588-ADBC-6563B92ACDAE Figures 2D, 3D, 4, 5 Type material. Holotype $, ECUADOR: Cerro Troya, Carchi, 0°43'59.7"N, 77°4r00.3"W, E. Pena leg. (MCZ 133396). Paratypes: S, ECUADOR: Pichincha, 0°15'00.0"S, 78°35'00.0"W, 19.iv.l988, W. Maddison leg. (MCZ 133397); $, COLOMBIA: Cundinamarca, Mosquera (Mondonedo), 4°4r0.06"N, 74°15'0.25"W, 20.X.2000, E. Florez & J. Pinzon leg. (ICN 3404). Material examined. COLOMBIA: 2j, Magdalena (San¬ ta Marta), 11°12'54"N, 74°06'01"W, 18.iv.l977, W. Gal- vis & A.J. Moreno leg. (ICN-Ar 9140); 1$ Ij, Quindio (Estacion Bremen), 4°40'0"N, 75°39'0"W, 14-20.iv.l998, P. Ariza leg. (MPUJ 45511); 1$, IJ, same data as preced¬ ing (MPUJ 11490); 1$, Cundinamarca, Mosquera (via a Ea Mesa), 4°40T2.0"N, 74°16'00.7"W, 20.X.2000, E. Flo¬ rez & J. Pinzon leg. (ICN-Ar 3404); 2$, Mosquera (De- sierto de Zabrinsky), 4°44'30"N, 74° 14'8"W, 23.ii.2002, E. Benavides, C. Nino, A. Castaneda & G. Mora leg. (ICN-Ar 2157); 2$, 5j, 06.iv.2002, same locality and col¬ lectors (ICN-Ar 2146); 4$, 7j, same locality, April 2006, zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 319-344 321 Figure 1. a-d Diagnostic sexual features of Sidymella a Detail of the nodose RTA surface on male palp b Tibial trichobothria on male palp (indicated by an arrow) c Glandular head on spermatheca d Walnut-shaped spermatheca e, f Photos of live specimens of Sidymella lucida. Photo credits; e Diego Galarraga Sugoniaev; f Damian Hagopian. zse.pensoft.net 322 Miguel Machado et al.: Taxonomic review of Neotropical Sidymella Figure 2. Habitussen of females, lateral a Sidymella longispina b Sidymella furcillata c Sidymella kolpogaster d Sidymella excavata sp. nov. e Sidymella marmorata sp. nov. f Sidymella lucida. zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 319-344 323 Figure 3. Habitussen of males, lateral a Sidymella longispina b Sidymella furcillata c Sidymella kolpogaster d Sidymella excavata sp. nov. e Sidymella lucida. zse.pensoft.net 324 Miguel Machado et al.: Taxonomic review of Neotropical Sidymella Figure 4. Female of Sidymella excavata sp. nov. a habitus, dorsal b prosoma, anterior c, e epigynum, ventral d, f epigynum, dorsal. zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 319-344 325 Figure 5. Male of Sidymella excavata sp. nov. a habitus, dorsal b prosoma, anterior c, e left palp, ventral view d, f left palp, retro- lateral view. zse.pensoft.net 326 Miguel Machado et al.: Taxonomic review of Neotropical Sidymella J. Martinez leg. (ICN-Ar 10500); 1$, 23.ii.2002, same lo¬ cality, C. Nino, G. Mora, A. Castaneda & J. Martinez leg. (CAUD-216-ARA 139); 1$, 27.vii.2002, same locality, J. Martinez, C. Nino & G. Mora leg. (CAUD-216-ARA 366); 1$, iv.2002, same locality, C. Nino, G. Mora, A. Castaneda & J. Martinez leg. (CAUD-216-ARA 1944); 1$, 08.ix.2016, same locality, D. Molina leg. (ICN-Ar 10449); 1$, same data as preceding (ICN-Ar 10498). 1$, 2j , Boyaca, Villa de Levya, 5°40'21'’N, 73°27'42"W, 09.vi.2001, L. Benavides leg. (ICN-Ar 1248). 1(5', Valle del Cauca, Saladito, 3°34'15.78"N, 76°36'9.27"W, W. Eber- hard leg. (MCZ 133395). ECUADOR: 1(5', Quito, Pichin- cha, 0°15'00"S, 78°35'0'’W, 19.iv.l988, W. Maddison leg. (MCZ 133397). 1(5', 2j, Azuay, Cerro Tinajillas, 3°10'60"S, 79°02'00''W, 18-21.iii.1965, L. Pena leg. (MCZ 133403). PERU: 1(5', Chavin, Ancash, 9°25'56"S, 77°32'07"W, 14.i.l972 (MACN-Ar 19068). ARGENTINA: 3(5', 4 j, Catamarca, Las Chacritas, 27°4ri9.25"S, 65°55'24.97"W, 20.i.l987, P. Goloboff leg. (MACN-Ar 19098). Etymology. The specific name refers to the shape of the anterior border of the opisthosoma with a remarkable me¬ dian re-entrance/excavation. Diagnosis. Females of S. excavata sp. nov. resemble those of S. marmorata sp. nov. by the large body size, short opisthosomal projections and long copulatory ducts; however, they can be distinguished by the presence of long needle-shaped macrosetae on the ocular area (Fig. 4B), and by having five mesial macrosetae on femora I and II (Fig. 4A). Males are similar to those of S. luci- da, but can be recognized by cephalic setae and a basal branch of the RTA which is truncated instead of conical (Fig. 5C-F). Description. Female: Prosoma, sternum and legs totally orange; legs I and II covered by numerous spiniform se¬ tae; opisthosoma predominantly light-yellow with a dark median stain on the dorsum that splits posteriorly towards to the tips of the projections (Fig. 4A). Epigynum with a wide septum formed by the median junction of the pos¬ terior folds of the epigynal plate (Fig. 4C, E); copulatory ducts hyaline, long and coiled, leading to a pair of wal¬ nut-shaped spermathecae (Fig. 4D, F). Measurements: eyes sizes and interdistances: AME 0.03, ALE 0.09, PME 0.05, PLE 0.03, AME-AME 0.15, AME-ALE 0.09, PME-PME 0.17, PME-PLE 0.17, MOQ length 0.41, MOQ width 0.17; leg formula: 1-2-4-3: leg I - femur 3.40/ patella 1.36/ tibiae 3.04/ metatarsus 2.04/ tarsus 0.88/ total 10.72; II - 2.68/ 1.12/ 2.08/ 1.72/ 0.68/ 8.28; III - 1.40/ 0.72/ 0.88/ 0.80/ 0.60/ 4 . 44 ; IV - 1.76/ 0.76/ 1.12/ 0.96/ 0.60/ 5.20. Total body length 5.48; prosoma 2.48 length, 2.04 wide; opisthoso¬ ma length 3.00; clypeus 0.35 height; sternum 1.12 length, 1.04 width; endites 0.56 length, 0.20 width; labium 0.32 length, 0.36 width. Male: Prosoma yellow with a darker median stain; legs I and II yellow with brownish spots on the femo¬ ral setiferous tubercles and at the distal portion of each leg segment; legs III and IV light-yellow. Opisthosoma as in female (Fig. 5A). Palpi with a well-developed pars pendula and embolus emerging at three o’clock, curling at the tip (Fig. 5C-F); basal branch of RTA stout and di¬ rected retro laterally while the RTA is short, rounded and oriented vertically (Fig. 5D, F). Measurements: eyes diameters and eyes interdistances: AME 0.05, ALE 0.07, PME 0.05, PLE 0.05, AME-AME 0.13, AME-ALE 0.05, PME-PME 0.11, PME-PLE 0.13, MOQ length 0.17, MOQ width 0.11; leg formula: l-2^-3: leg I - femur 2.04/ patella 0.72/ tibiae 1.92/ metatarsus 1.68/ tarsus 0.76/ total 7.12; II - 1.44/ 0.60/ 1.20/ 1.12/ 0.60/ 4.96; III - 0.72/ 0.60/ 0.56/ 0.40/ 0.40/ 2.68; IV - 0.96/ 0.36/ 0.60/ 0.52/ 0.40/ 2.84. Total body length 3.12; prosoma 1.56 length, 1.24 wide; opisthoso¬ ma length 1.56; clypeus 0.19 height; sternum 0.68 length, 0.62 width; endites 0.35 length, 0.13 width; labium 0.17 length, 0.25 width. Distribution. COLOMBIA: Magdalena, Quindio, Cun- dinamarca, Boyaca, and Valle del Cauca; ECUADOR: Pichincha and Azuay; PERU: Chavin; ARGENTINA: Catamarca (Fig. 15). Sidymella furcillata (Mello-Leitao, 1944) Figures 2B, 3B, 6, 7 Sidyma spinifera Mello-Leitao, 1943: 209, f. 36 preoccupied by Mello-Leitao 1929). Sidyma multispinulosa Mello-Leitao, 1944: 4 (replacement name). Lise 1973: 13, figs 24-33 (holotype $ from Santa Catarina, Brazil, P. Buck leg., MNRJ 41934, examined). (New synonymy) Stephanopis furcillata Keyserling, 1880: 179, pi. 4, fig. 98 (holotype $ from Santa Cruz, Rio Grande do Sul, Brazil, Hensel leg., ZMB 2406, examined). (New combination) Material examined. ARGENTINA: Ij, Misiones, San Vicente, 26°55T2"S, 54°31T2"W, 12.i.2005, L. Lopardo et al. leg. (ICN-Ar 27642); Ij, same locality, viii.1954, Schiapelli de Carlo leg. (ICN-Ar 19099); 2j, same locality, 1954, Schiapelli de Carlo leg. (ICN-Ar 19096). BRAZIL: 1(5', Bahia: Maracas, 13°28T5'’S, 40°26T6"W, 14.hi.2012, E.S. Araujo & A.S Medeiros leg. (UFMG 15165); 1(5, Sao Paulo, Botucatu (Ru- biao Jumior), 22°53'53"S, 48°29'23"W, 15.ii.l966, VC. Jesus leg. (MNRJ 10391); 1$, Parana: Colombo, 25°17'31"S, 49°13'26"W, 02.xii.l990 (MCN 20684); 1(5, Tres barras do Parana, 25°25'08"S, 53°10'51"W, 20-26.ii. 1993 (MCN 23043); 1$, Curitiba (Parque Bi- rigui), 25°25'31.58"S, 49°18'38.69"W, 01.xii.l990, A.B. Bonaldo leg. (MCN 20626); 1$, Curitiba, 25°25'47"S, 49°16T9"W, 15.iv.2005, J. Ricetti leg. (MCTP 37237); 1$, Santa Catarina: Itapiranga, 27°10'08"S, 53°42'43"W, Pio Buck leg. (MNRJ 41934); 2$, 3j, Rancho Queimado, 27°40'22"S, 49°01T9"W, 13-15.1.1995 (MCN 26464); 1(5, Rio Grande do Sul, Tenente Portela, 27°22'25.69"S, 53°47'32.43"W, ll.ix.l976, S. Scherer leg. (MCN 4862); zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 319-344 327 1 Nonoai (Parque Estadual de Nonoai), 27°27'21.25"S, 53°6'46.98"W, 14.i.l985, A.A. Lise leg. (MCN 13077); 1$, Arroio do Meio, 29°23'47.15'’S, 51°56'54.26"W, 09.i. 1985, same collector (MCN 13071); Ij, Iral, 27°ir38"S, 53°15'03"W, 20 September 1975, same col¬ lector (MCN 8084); 1(5', Derrubadas (Parque Estadual do Turvo), 27°13'57'’S, 53°5r04''W, 16.i.l985, same collec¬ tor (MCN 13057); 1$, same locality, 01 February 1996, A.B. Bonaldo, A. Kury & R. Pinto-da-Rocha leg. (MCN 27100); Ij, 27-31.viii.2003, R. Ott leg. (MCN 37827); Ij, Caxias do Sul, 29°07'17'’S, 51°0r07''W, 18-21.xi.l993 (MCTP4176); 1$, Farroupilha, 29°13'30"S, 51°20'52"W, 29.ix.1978, H. Bischoff leg. (MCN 8268); 1$, 2 j, same data as preceding (MCN 8270); 1(5, Estrela Velha (Bar- ragem de Itauba), 29°15'19''S, 53°13'36'’W, 07.iii. 2001, R. Ott leg. (MCN 33700); 1$, Sao Francisco de Paula (Barragem Passo do Inferno), 29°16'29"S, 50°44'15"W, 26.ix.2000, M.A.L. Marques leg. (MCN 33220); 1$, Sao Francisco de Paula, 29°27'00"S, 50°34'59"W, lO.xii. 2007, L. Moura & R. Moraes leg. (MCN 43748); 1$, Parobe (Rio dos Sinos), 29°37'29"S, 50°49'56"W, ll.v. 2008, E.N.L. Rodrigues leg. (MCN 47943), 1 Maquine, 29°39'41"S, 50°12'47'’W, 08-09.iii.1998, L.A. Moura leg. (MCN 29060); 1 $, Campo Bom, 29°40'44"S, 51°3'10"W, 19-20.V.1986, C.J. Becker leg. (MCN 15080); 2$, Santa Maria, 29°40'59"S, 53°48'00"W, 17.iv.l996, C.B. Kot- zian & L. Indrusiak leg. (MCTP 40093); Ij, Montenegro, 29°41'20"S, 51°27'39"W, 06.viii.l977, H. Bischoff leg. (MCN 6822); Ij, same locality, 03.xi. 1977, A.A. Lise leg. (MCN 7141); 1(5,1$, 3j, same locality, 15.xii.l977, same collector (MCN 7510); 1$, same locality, 20.xii.l977, same collector (MCN 7614); Ij, same locality, 03.xi.l977, same collector (MCN 8089); 2j, same locality, 29.ix. 1977, same collector (MCN 9058); 5j, same data as preceding (MCN 9059); 1 $, Sao Leopoldo, 29°45'36''S, 51°8'49'’W, 12.vi.l992 (MCN 24755); 2$, 3j, Montenegro (Pesque- iro), 29°46'16"S, 51°23'57"W, 12.v. 1977, A.A. Lise leg. (MCN 5323); 1$, Santo Antonio da Patrulha, 29°49'4"S, 50°3ri2"W, 18.vii.2000, same collector (MCN 33116); 2$, Triunfo (Parque Copesul de Prote^ao Ambiental), 29°5r57''S, 51°2r54"W, 25.1.1990, A.B. Bonaldo leg. (MCN 19344); 1$, same locality, 13.1.1994, A.F. Fran- ceschini leg. (MCN 24771); Ij, same locality, 25.xi.1994, L.A. Moura leg. (MCN 26116); 1(5', 2$, same locality, 08.1.1997, E.H. Backup leg. (MCN 28176); 2$, 14- 15.1.1997, L.A. Moura leg. (MCN 28263); 3$, same lo¬ cality, 16-17.hi. 1998, L.A. Moura leg. (MCN 29238); 1$, same locality, 05.ii.2003, R. Ott leg. (MCN 35125); 2$, same locality, 29-30.iv.2003, R. Ott leg. (MCN 35740); 2$, same locality, 06.1.2005, R. Ott leg. (MCN 38352); 1$, same locality, 01.hi.2005, R. Ott leg. (MCN 39924); 1$, same locality, 28.v. 2007, E.N.L. Rodrigues leg. (MCN 43383); 1(5, 1$, same locality, 14.ii.2008, R. Moraes leg. (MCN 44009); 1$, same locality, 03.iv.2008, A. Barcellos leg. (MCN 44127); 1$, same locality, 26.ii.2010, M.C. Pairet Jr. leg. (MCN 47032); 1$, same locality, 26.ii.2010, M.A.L. Marques leg. (MCN 47049); 3$, Canoas, 29°55'12"S, 51°10'48"W, 13.xii.l990 (MCN 20142); 1(5', 1$, 2j, same locality, 24.i.l991, H. Gali¬ leo & E.H. Backup leg. (MCN 20431); Ij, same local¬ ity, 25.xii.1983, A.D. Brescovit leg. (MCN 11904); 3$, Triunfo, 29°56'34"S, 51°43'4"W, 20.x. 1977, H. Bischoff leg. (MCN 6900); 3$, same locality, 28.xi.1977, same collector (MCN 7314); 1 Triunfo (Parque Estadual Del¬ ta do Jacui), 29°57T6"S, 51°12'55"W, 23.ii.1999, A.B. Bonaldo leg. (MCN 30488); Ij, Porto Alegre, 30°0r58"S, 51°13'48"W, 04.ix.l977, A.A. Lise leg. (MCN 6469); 1$, same locality, 17.xii. 1983, A.D. Brescovit leg. (MCN 11882); 2(5', 1$, same locality, 18.i.l992, A.D. Bresco¬ vit leg. (MCN 21958); 1$, same locality, 2010, biology students leg. (MCTP 41329); Ij, Porto Alegre (Morro Santana), 30°02'34"S, 51°08'39"W, 22.X.1981, A.A. Lise leg. (MCN 11421); Ij, Viamao (Estagao Experimen¬ tal Fitotecnica Aguas Belas), 30°02'51"S, 51°00'53"W, 24.1.1977, A.A. Lise leg. (MCN 5140); 1(5', same local¬ ity, 06.1.1977, same collector (MCN 5763); 1$, Viamao (Escola Marista), 30°04'41"S, 51°03'02"W, 1994, A. Braul leg. (MCTP 4726); 3j, Porto Alegre (Ponta Gros- sa), 30°10'29"S, 51°ir50"W, 03.ix.l975, A.A. Lise leg. (MCN 3029); Ij, same locality, 07.V.1976, same collector (MCN 8086); 3j, Viamao (Morro do Coco), 30°16T1"S, 51°03T5"W, 04.x. 1975, A.A. Lise leg. (MCN 8085); 1$, same locality, 25.vii.1985, same collector (MCN 13378); Ij, Viamao, 30°4'51"S, 51°r22"W, 12.1.1996, same col¬ lector (MCTP 8365); 1$, same locality, 23.1.1996, same collector (MCTP 9294); 3j, same locality, 02.xh. 1994, A.A. Lise & A. Braul leg. (MCTP 5876); 5j, same lo¬ cality, 12.vii. 1994, A.A. Lise leg. (MCTP 5244), 1$, Ij, Guaiba, 30°6'50"S, 51°19'30''W, 09.1.1996, A.A. Lise leg. (MCTP 8247); 1(5', Cristal (Rio Camaqua), 31°00T2"S, 52°04'02"W, 03.ii.2008, E.N.L. Rodrigues leg. (MCN 48665). Diagnosis. Females of S.furcillata are similar to those of S. longispina by having macrosetae above the ALE and long opisthosomal projections that are directed vertical¬ ly (Fig. 2B); however, in S.furcillata the projections are rounded at the apex with a smaller terminal tubercle (Fig. 2B). They can be also distinguished by having a group of clavate setae on the median region of the prosoma (Fig. 6B), a triad of mesial macrosetae on femora I, presence of seven pairs of ventral macrosetae on tibiae I, a dark transversal line on the dorsum of opisthosoma (Fig. 6A), and by the long and coiled copulatory ducts (Fig. 6D, F). Males can be distinguished by having tubercles at the apexes of the opisthosomal projections (Fig. 3B), wide pars pendula (Fig. 7C, E), retrolateral tegular process, and the bifid basal branch of the RTA (Fig. 7D, F). Description. Female: Prosoma dark-yellow to light- brown, covered by hyaline setae; legs of the same colour as pro soma, with some sparse darker circular taints ran¬ domly distributed; dorsal surface of coxae I and II with wide guanine spots (Fig. 6A). Opisthosoma dark-yellow with two transversal dark lines, being the posterior line disposed along the projections (Fig. 6A). zse.pensoft.net 328 Miguel Machado et al.: Taxonomic review of Neotropical Sidymella Figure 6. Female of Sidymella furcillata a habitus, dorsal b prosoma, anterior c, e epigynum, ventral d, f epigynum, dorsal. zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 319-344 329 Figure 7. Male of Sidymella furcillata a habitus, dorsal b prosoma, anterior c, e left palp, ventral view d, f left palp, retrolateral view (white arrow indicates the basal branch of the RTA). zse.pensoft.net 330 Miguel Machado et al.: Taxonomic review of Neotropical Sidymella Measurements: eyes diameters and eyes interdistances: AME 0.05, ALE 0.09, PME 0.05, PEE 0.05, AME-AME 0.13, AME-ALE 0.11, PME-PME 0.15, PME-PLE 0.15, MOQ length 0.33, MOQ width 0.15; leg formula: 1-2^- 3: leg I - femur 3.36/ patella 1.40/ tibiae 3.12/ metatarsus 2.24/ tarsus 0.92/ total 11.04; II - 3.17/ 1.32/ 2.40/ 1.96/ 0.76/ 9.61; III - E16/ 0.68/ 0.92/ 0.60/ 0.52/ 3.88; IV- 1.36/ 0.72/ 0.84/ 0.52/ 0.48/ 3.92. Total body length 5.05; prosoma 2.04 length, 2.08 wide; opisthosoma length 3.01 (considering the projections); clypeus 0.29 height; ster¬ num 1.03 length, 0.94 width; endites 0.47 length, 0.29 width; labium 0.27 length, 0.27 width. Male: Prosoma dark-yellow, legs I and II predomi¬ nantly light-brown with dark spots; tibiae I yellowish on the median region; legs III and IV yellowish (Fig. 7A). Opisthosoma yellowish with a pair of dorsolateral and a median posterior black spot (Fig. 7A). Embolus long, emerging from tegulum at four o’clock, with wide pars pendula and curled at the tip; RTA rounded and with cen¬ tral excavation (Fig. 7C-F). Measurements: eyes diameters and eyes interdistances: AME 0.03, ALE 0.07, PME 0.05, PEE 0.05, AME-AME 0.11, AME-ALE 0.09, PME-PME 0.17, PME-PLE 0.13, MOQ length 0.31, MOQ width 0.11; leg formula: 1-2^- 3: leg I - femur 3.08/ patella 1.32/ tibiae 2.88/ metatarsus 2.32/ tarsus 0.88/ total 10.48; II - 2.48/ 1.00/ 2.04/ 1.96/ 0.80/ 8.28; III - 0.76/ 0.52/ 0.76/ 0.56/ 0.48/ 3.08; IV - 1.24/ 0.44/ 0.76/ 0.48/ 0.48/ 3.40. Total body length 3.96; prosoma 1.96 length, 1.88 wide; opisthosoma length 2.00 (considering the projections); clypeus 0.29 height; ster¬ num 0.90 length, 0.82 width; endites 0.43 length, 0.25 width; labium 0.23 length, 0.27 width. Distribution. ARGENTINA: Missiones; BRAZIL: Ba¬ hia, Sao Paulo, Parana, Santa Catarina, and Rio Grande do Sul (Fig. 16). Sidymella kolpogaster Lise, 1973 Figures 2C, 3C, 8, 9 Sidyma kolpogaster Lise, 1973: 5, figs 5-9 ($); Lise 1981: 130, figs 1-12 (( 3 ,?). Sidymella kolpogaster. Brignoli, 1983: 605. Type material. Holotype BRAZIL: Parana, Rio Ne¬ gro, Franciscanos leg. (MNRJ 58065, examined; lost in the fire of the Museu Nacional do Rio de Janeiro). Neo¬ type BRAZIL: Parana, Curitiba (Parque Barigui), 25°25T2'’S, 49°18'21"W, A.B. Bonaldo leg., 01.xii.l990 (MCN 20622, designated here). Other material examined. BRAZIL: 1$, Parana, Rio Negro, 26°05' 11" S, 49°46'47"W (MNRJ 58065); 1 S, San¬ ta Catarina, Rancho Queimado, 27°40'22"S, 49°01T9"W, 09-13.x. 1995, A.A. Lise, A. Braul & M. Silveira leg. (MCTP 7009); 1$, Rio Grande do Sul, Estrela Velha, 29°15T9"S, 53°13'36"W, 21.X.1998, A.B. Bonaldo leg. (MCN 30791); Ij, Arroio do Tigre (Itauba), 29°16'47"S, 53°06'34"W, 18.iv.l978, A.A. Lise leg. (MCN 8047); Ij, Arroio do Tigre, 29°19'58"S, 53°05'34"W, 23.iv.1978 (MCN 8150); 1$, Sao Francisco de Paula, 29°24'52"S, 50°15'24"W, 20.ix.l998, A.A. Lise leg. (MCTP 14446); Ij, same locality, 21-24.iii. 1995, same collector (MCTP 12010); Ij, Montenegro, 29°4F20"S, 51°27'39"W, 20.xii. 1977, A.A. Lise leg. (MCN7613); 16', 1$, Glorinha (Rincao Sao Joao), 29°49'00"S, 50°50'00"W, 14.vii.2000, A.B. Bonaldo leg. (MCN 33053); 16', Triunfo (Parque Copesul de Prote9aoAmbiental),29°51'57"S,51°21'54"W, 01.ix.2008, E.N.L. Rodrigues leg. (MCN 45434); 1$, Triunfo (Parque Braskem), 29°51'57"S, 51°21'54"W, 04.xii.2009, E.N.L. Rodrigues leg. (MCN 46847); Ij, Porto Alegre (Morro Santana), 30°02'34"S, 51°08'39"W, 17.V.1980, A.A. Lise leg. (MCN 9065); Ij, Viamao (Es- ta9ao Experimental Fitotecnica Aguas Belas), 30°02'51"S, 51°00'53''W, 12.viii.l994, A.A. Lise leg. (MCTP 5243); 1$, same locality, 23.ix. 1994, same collector (MCTP 5498); 16', same locality, 06.v. 1994, A.A. Lise & A. Braul leg. (MCTP 4683); 1$, same locality, 17.i. 1977, A.A. Lise leg. (MCN 4995); 1$, Viamao, 30°04'51"S, 51°0F22'’W, 14.V.2005, R. Jalisco leg. (MCTP 19737). Diagnosis. Females of S. kolpogaster are similar to those of S. lucida by the general shape of the spermathecae, short copulatory ducts, and femora I with two mesial macrosetae. However, females of S. kolpogaster can be easily distin¬ guished from congeners by curved tibiae I and II, fiattened prosoma, dark body colouration, varying from dark-brown to black and contrasting with their vivid yellow legs III and IV (Fig. 8A). The males can also be recognized by their body colour pattern, which is the same as in females, and by their truncated RTA fused with basal branch (Fig. 9D, F). Description. Female: Prosoma dark-brown, lighter on the ocular area and clypeus, covered by hyaline setae and dor- soventrally compressed (Fig. 8B). legs I and II entirely dark- brown, except for the reduced and yellowish tarsi; metatarsi I and II curved; legs III and IV entirely light-yellow (Fig. 8A). Opisthosoma dark-brown with posterior projections stout and conical. Epigynal plate wide and with short sep¬ tum; copulatory ducts short and hyaline (Fig. 8C-F). Measurements: eyes diameters and eyes interdistances: AME 0.07, ALE 0.11, PME 0.05, PEE 0.05, AME-AME 0.17, AME-ALE 0.09, PME-PME 0.23, PME-PLE 0.19, MOQ length 0.21, MOQ width 0.15; leg formula: l-2^-3: leg I - femur 4.95/ patella 1.80/ tibiae 4.15/ metatarsus 2.95/ tarsus 0.95/ total 14.80; II - 4.45/ 1.75/ 3.20/ 2.65/ 0.85/ 12.90; III - 1.65/ 0.75/ 1.15/ 1.05/ 0.55/ 5.15; IV-2.05/ 0.75/ 1.25/ 1.15/ 0.55/ 5.75. Total body length 7.65; proso¬ ma 2.80 length, 2.85 wide; opisthosoma length 4.85; cly¬ peus 0.19 height; sternum 1.21 length, 1.20 width; endites 0.68 length, 0.37 width; labium 0.41 length, 0.50 width. Male: Prosoma and legs colouration as in female (Fig. 9A, B). Palpi with fiattened cymbium and dorsal trichobohtria on tibiae; tegulum discoid and embolus thin, slightly curved at the tip and emerging from tegulum at four o’clock (Fig. 9C-F). zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 319-344 331 Figure 8. Female of Sidymella kolpogaster a habitus, dorsal b prosoma, anterior c, e epigynum, ventral d, f epigynum, dorsal. zse.pensoft.net 332 Miguel Machado et al.: Taxonomic review of Neotropical Sidymella Figure 9. Male of Sidymella kolpogaster a habitus, dorsal b prosoma, anterior c, e left palp, ventral view d, f left palp, retro- lateral view. zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 319-344 333 Measurements: eyes diameters and eyes interdistances: AME 0.03, ALE 0.05, PME 0.03, PEE 0.05, AME-AME 0.11, AME-ALE 0.05, PME-PME 0.09, PME-PLE 0.09, MOQ length 0.21, MOQ width 0.11; leg formula: l-2^-3: leg I - femur 2.04/ patella 0.68/ tibiae 1.68/ metatarsus 1.56/ tarsus 0.60/ total 6.56; II - 1.64/ 0.36/ 1.44/ 1.20/ 0.52/ 5.16; III - 0.72/ 0.36/ 0.52/ 0.52/ 0.32/ 2.44; IV - 0.92/ 0.32/ 0.76/ 0.48/ 0.36/ 2.84. Total body length 3.12; prosoma 1.40 length, 1.32 wide; opisthoso- ma length 1.72; clypeus 0.07 height; sternum 0.64 length, 0.68 width; endites 0.27 length, 0.32 width; labium 0.13 length, 0.25 width. Distribution. BRAZIL: Parana, Rio Grande do Sul, San¬ ta Catarina (Fig. 17). Sidymella longispina (Mello-Leitao, 1943) Figures 2A, 3A, 10, 11 Sidyma longispina Mello-Leitao, 1943: 208, fig. 35 ($); Lise 1973: 10, figs 19-23 ($). Type material. Holotype BRAZIL: Rio Grande do Sul, B. Rambo leg. (MNRJ 41911, examined; lost in the fire of the Museu Nacional do Rio de Janeiro). Neotype female, BRAZIL: Rio Grande do Sul, Sao Francisco de Paula, 29°24'52''S, 50°15'24"W, A.A. Lise et af, 24-26. iv.2006 (MCTP 35899, designated here). Other material examined. BRAZIL: 1(5', Minas Gerais, Vale Verde (Parque Nacional do Caparao), 20°25'05"S, 41°50'48"W, 24-30.xi.2014, B.T. Faleiro leg. (UFMG 18264). 1(5', Ij, Parana, Foz do Igua^u (Refugio Bi- ologico de Bela Vista), 25°26'56"S, 54°33'23"W, 09-11. xi.l991, A.B. Bonaldo leg. (MCN 21809). 1$, Santa Catarina, Rancho Queimado, 27°40'22"S, 49°01T9"W, 08-12.X.1994, A.B. Bonaldo & L.A. Moura leg. (MCTP 5952). 3j, Rio Grande do Sul, Tenente Portela, 27°22' 15" S, 53°45'28"W, 17.i.l985, A.A. Lise leg. (MCN 13068); 2j, Irai, 27°11'38"S, 53°15'03"W, 18.xi. 1975, A.A. Lise leg. (MCN 3142); lOj, same locality, 21.xi.l975, same collector (MCN 8082); Ij, Derrubadas (Parque Estadu- al do Turvo), 27°13'57"S, 53°5r04"W, 04-06.ii.l980, A.A. Lise leg. (MCN 8984); 3$, same locality, 04- 07.V.2004, R. Ott leg. (MCN 38864); 1$, same locality, 19-22.X.2004. R. Ott leg. (MCN 38878); If, Caxias do Sul (Vila Oliva), 29°12'56"S, 50°53'22''W, 14.ix.l976, H. Bischoff leg. (MCN 4485); 1(5', 10.iv.l992, L.A. Moura leg. (MCN 22124); Ij, Canela, 29°2r57"S, 50°48'57"W, 24.viii.1975, A.A. Lise leg. (MCN 3026); 2$, Sao Fran¬ cisco de Paula, 29°27'00'’S, 50°34'59"W, 14.V.1993, A. Braul leg. (MCTP 3213); 3j, same locality, 05.i.1985, A.A. Lise leg. (MCN 13107); 1$, Maquine, 29°40'47"S, 50°11'20"W, 22.vi.2008, E.N.L. Rodrigues leg. (MCN 54202); 1$, Santa Maria, 29°40'59"S, 53°48'00"W, 18.i.l999, C.B. Kotzian & L. Indrusiak leg. (MCTP 40101); 1(5, 1$, same locality, 30.ix. 1992, biology stu¬ dents leg. (MCTP 41330); 1$, Gravatai (Mato Alto), 29°57'20"S, 50°57'46"W, 13.iii.l985, A.D. Brescovit leg. (MCN 13094); 2$, Cachoeira do Sul, 30°0'0"S, 53°0'0"W, B. Rambo leg. (MNRJ 41911); 1$, Panta- no Grande, 30°11'27"S, 52°22'26"W, 05.iv.2008, G. Depra leg. (MCTP 40084); 1$, Guaiba, 30°6'50'’S, 51°19'30"W, 28.iv.1995, A. Braul leg. (MCTP 7542); 1$, Cristal (Rio Camaqua), 31°00T2"S, 52°04'02"W, 19.xii.2007, E.N.L. Rodrigues leg. (MCN 48618); 1(5', same locality, 14.ii.2008, same collector (MCN 48832); 1$, same locality, 21.iv.2008, same collector (MCN 49031); Ij, Sao Borja, 28°10T6.10"S, 55°26'49.78"W, 06.xii. 1975, A.A. Lise leg. (MCN 8088). ARGENTINA: 1$, Misiones, Puerto Iguazu (Parque Nacional Iguazu), 25°4r01"S, 54°27T4"W, 22-30.viii.1986, M. Ramirez leg. (MACN-Ar 19069); 2j, same locality, 19-20.i.2005, C. Grismado et al. leg. (MACN-Ar 27637); 1$, same locality, vi.l985, M. Ramirez leg. (MACN-Ar 19067); Ij, Iguazu, 25°50'53.71"S, 54°20'48.17"W, x.1954, Schiapelli de Carlo leg. (MACN-Ar 19070); Ij, Monte- carlo, 26°33'58"S, 54°45'25"W, i.l966, same collector (MACN-Ar 19072); 3(5', 2j, Santa Maria, 27°53'39'’S, 55°2r20"W, 1952, C. Viana leg. (MCTP 3537). Diagnosis. Females of S. longispina are similar to those of S. furcillata by their long and vertically oriented opist- hosomal projections (Figs 2A, lOA), and the presence of stout macrosetae above the ALE (Fig. lOB); however, their opisthosomal projections have pointy conical apex¬ es instead of being rounded with a small apical protuber¬ ance as in S. furcillata. The females of S. longispina also have shorter copulatory ducts (Fig. lOD, F). Males of S. longispina resemble those of S. furcillata but their palpi bear a truncated RTA with a conical and well-developed basal branch (larger than the RTA itself) (Fig. IID, F); the male palp has a long, acute tegular process and a dor- so-basal projection (Fig. 1 ID, F). Both males and females have just one mesial macroseta on femora I (Fig 11 A). Description. Female: Prosoma yellowish-orange, darker on the cephalic area and presenting a pair of needle-shaped macrosetae on conical projections above the ALE (Fig lOB). Legs yellowish-orange; femora I with a single pro¬ lateral macrosetae; both the anterior tibiae and metatarsi (I and II) ventrally armed with five pairs of ventral macrose¬ tae. Opisthosoma light-yellow with posterior projections long, pointed and vertically oriented (Figs 2A, lOA). Measurements: eyes diameters and eyes interdistances: AME 0.05, ALE 0.09, PME 0.05, PLE 0.05, AME-AME 0.15, AME-ALE 0.11, PME-PME 0.15, PME-PLE 0.17, MOQ length 0.35, MOQ width 0.15; leg formula: 1-2^- 3: leg I - femur 4.45/ patella 1.60/ tibiae 4.20/ metatarsus 3.05/tarsus 0.85/total 14.15; 11-3.35/ 1.30/2.95/ 1.95/ 0.75/ 10.30; III - 1.25/ 0.65/ 1.05/ 0.55/ 0.55/ 4.05; IV - 1.60/ 0.70/ 0.95/ 0.55/ 0.45/ 4.25. Total body length 7.93; prosoma 2.56 length, 2.12 wide; opisthosoma length 5.37 (considering the projections); clypeus 0.23 height; ster¬ num 1.11 length, 1.00 width; endites 0.60 length, 0.27 width; labium 0.31 length, 0.37 width. zse.pensoft.net 334 Miguel Machado et al.: Taxonomic review of Neotropical Sidymella Figure 10. Female of Sidymella longispina a habitus, dorsal b prosoma, anterior c, e epigynum, ventral d, f epigynum, dorsal. zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 319-344 335 Figure 11. Male of Sidymella longispina a habitus, dorsal b prosoma, anterior c, e left palp, ventral view d, f left palp, retrolateral view (white arrow indicates the tegular process). 1 mm 0,5 mm zse.pensoft.net 336 Miguel Machado et al.: Taxonomic review of Neotropical Sidymella Male: Body colouration pattern and other characteris¬ tics as in female (Fig. 11 A, B). Measurements: eyes diameters and eyes interdistances: AME 0.03, ALE 0.05, PME 0.03, PEE 0.03, AME-AME 0.09, AME-ALE 0.05, PME-PME 0.11, PME-PLE 0.11, MOQ length 0.21, MOQ width 0.13; leg formula: 1-2^- 3: leg I - femur 2.44/ patella 1.12/ tibiae 3.12/ metatarsus 2.28/ tarsus 0.92/ total 9.88; II - 2.08/ 0.76/ 1.64/ 1.48/ 0.72/ 6.68; III - 0.60/ 0.48/ 0.60/ 0.40/ 0.40/ 2.48; IV - 1.04/ 0.44/ 0.60/ 0.40/ 0.40/ 2.88. Total body length 3.04; prosoma 1.40 length, 1.72 wide; opisthosoma length 1.64 (considering the projections); clypeus 0.17 height; ster¬ num 0.76 length, 0.66 width; endites 0.33 length, 0.17 width; labium 0.17 length, 0.25 width. Distribution. BRAZIL: Minas Gerais, Parana, Santa Ca¬ tarina, and Rio Grande do Sul; ARGENTINA: Misiones (Fig. 17). Sidymella lucida (Keyserling, 1880) Figures 2F, 3E, 12, 13 Stephanopis lucida Keyserling, 1880: 190, pi. 4, fig. 105 ($). Sidyma lucida. Simon 1895: 1056. Berland 1913: 95, pi. 9, figs 51-56 ((?¥)- Sidyma cancellata. Mello-Leitao 1934: 207, fig. 34 ((3). Sidymella lucida. Strand 1942: 399. Machado et al. 2017: 454, figs S11A,F, SMB ( 3 '). Sidyma lucida Lise, 1973: 6, figs 10-18 (d'?)- Type material. Holotype S, BRAZIL: Rio Grande do Sul, B. Rambo (MNRJ 41748, examined). Other material examined. ECUADOR: 1(5', Tunguhua, Banos, 01°23'51"S, 78°25'27''W, 10.iv.l939 (MCZ 133401); 1(5', 1$, 2j, same locality, iv.l939 (MCZ 133402); COLOMBIA: 1$, Valle del Cauca, Cali, 3°20'50"N, 76°34'34"W, x.1972 (MCZ), BRAZIL: 1$, Minas Gerais, Tiradentes (Serra de Sao Jose), 21 °06'36"S, 44°10'23"W, 28.x. 1989, Becker, Johann & Roppa leg. (MNRJ 2694); 1 Rio de Janeiro, Santa Maria Madale- na, 21°57T8"S, 42°0'28"W, 15.V.2002 (MCN 34671); 1$, Sao Paulo, Vale do Ribeira, 24°37'28.68"S, 47°23'33.99"W, vi.2002, E.H. Wienskoski leg. (MNRJ 11513); 1$, Ij, Parana, Sao Jose dos Pinhais, 25°32'06"S, 49°12'21"W, 09.xii.2015, A.C. Domahovski leg. (MCTP 39092); 1(5, 2$, Santa Catarina, Rancho Queimado, 27°40'22"S, 49°01T9"W, 13-15.i.l995, L.A. Moura leg. (MCN 26465); 1$, Alta Feliz, 29°23'31"S, 51°18'43"W, vii.1988, A.A. Lise leg. (MCTP 526); 1 Rio Grande do Sul, Derrubadas (Parque Estadual do Turvo), 27°13'57"S, 53°5r04"W, 27-31.X.2003, R. Ott leg. (MCN 37826); 1$, same locality, 19-22.X.2004, same collector (MCN 38879); 1$, Ij, Sao Valentim, 27°33'28"S, 52°3F26"W, 16.x. 1976 (MCN 4704); 1$, Santa Rosa, 27°52T5"S, 54°28'34"W, 02.i.l984, A.D. Brescovit leg. (MCN 11989); 2j, Esmeralda, 28°03T4'’S, 51°11'24"W, 24.V. 1975, A.D. Brescovit leg. (MCN 2881); 1$, Esmer¬ alda (Estagao Ecologica de Esmeralda), 28°03'8.35"S, 51°1F36.92"W, 12.xii.l978, C.J. Becker leg. (MCN 8467);6$,3J,Garruchos,28°10'16.21'’S,55°26'49.38"W, 08.xii.l975, A.A. Lise leg. (MCN 8083); 1$, same local¬ ity, 08.xi.l979, H. Bischoff leg. (MCN 8965); 4j, Vacaria, 28°30'43"S, 50°56'02"W, 23.V.1981 (MCN 9758); 1(5', Augusto Pestana, 28°3F01"S, 53°59'31"W, 06.ix.2009, L.V Silva-Leomar & B. Medeiros (MCTP 30588) 1$, same locality, 12.ix.2008, L.V. Silvia-Leomar et al. leg. (MCTP 27094); 1$, Sao Borja (Reserva Biologica de Sao Donato), 28°39'39"S, 56°00T4"W, 23.1.2012, Mach¬ ado, M. leg. (MCTP 34729); 2(5', 3$, Salto do Jacui (Hor- ta da CELL), 29° 05'21.03'’S, 53°12'41.24"W, 19.x. 1998, A.B. Bonaldo leg. (MCN 30761); 1 Caxias do Sul (Fa¬ zenda Souza), 29°07T7''S, 51°01'07"W, ll-12.xi.l995, lab staff leg. (MCTP 7322); 1(5', Mugum, 29°09'54"S, 51°52'04"W, 02.iii.l984 (MCN 12090); 1$, Caxias do Sul (AguaAzul), 29°1F51"S, 50°59'27"W, 15.ix.l976, E.H. Buckup (MCN 4498); 1(5', 4j, Caxias do Sul (Vila Oliva), 29°12'56"S, 50°53'22"W, 05.iv.l975, H. Bischoff leg. (MCN 2872); 1$, Estrela Velha (Barragem de Itau- ba), 29°15T9"S, 53°13'36"W, 20.x. 1998, L.A. Moura leg. (MCN 30763); 1$, same locality, 28.x. 1999, A. Silva (MCN 31959); 1(5', Sao Francisco de Paula, 29°16'29"S, 50°44'15"W, 19.xi.l990, E.H. Buckup leg. (MCN28841); 1$, same locality, 16.xii.l999, A.F. Franceschini leg. (MCN 32047); Ij, Canela, 29°21'57'’S, 50°48'57"W, 07.x. 1967, A.A. Lise leg. (MCN 649); 3j, same locality, 05.ii.l970, same collector (MCN 651); 2$, same locality, 20.i.l972, same collector (MCN 1025); 1$, same locali¬ ty, 31.xii.l973, same collector (MCN 2029); 1$, same locality, 2Fix. 1974, same collector (MCN 2249); 3$, same locality, 26.xii.1974, same collector (MCN 2492); Ij, same locality, 08.xi.l975, same collector (MCN 5968); Ij, same locality, 05.i. 1973, same collector (MCN 9056) ; Ij, same locality, ll.i.l966, same collector (MCN 9057) ; 1 $, Sao Francisco de Paula (FLONA), 29°25'47" S, 50°23'35"W, 19.xii.2010, R.A. Teixeira leg. (MCTP 33303); 1$, same locality, lO.x.2012, same collector (MCTP 41327); 3j, Sao Francisco de Paula, 29°27'00"S, 50°34'59"W, 05.i.l985, A.A. Lise leg. (MCN 12723); Ij, Tres Coroas, 29°30'55"S, 50°46'46"W, 15.xii. 1976, A.A. Lise leg. (MCN 4924); 1$, Sao Martinho da Serra, 29°32'16"S, 53°5F18"W, 19.x. 1993, L. Indrusiak & R.A. Boelter leg. (MCTP 40116); 1$, Itaara, 29°36'36"S, 53°45'54"W, 22.xi.2006, L. Indrusiak & R.A. Boelter leg. (MCTP 21356); 2j, same locality, 23.vi.2007, A.A. Lise et al. leg. (MCTP 21357); Ij, same locality, 14.vii.2007, L. Indrusiak leg. (MCTP 21358); 1 $, Agudo, 29°38'31''S, 53°15'10"W, 21.X.1988, L. Indrusiak leg. (MCN 18751); 1$, Campo Bom, 29°40'44"S, 51°3T0"W, 19.x. 1987, L. Indrusiak leg. (MCTP 135); 1 Santa Maria, 29°40'59"S, 53°48'00"W, 03.vii.2000, L. Indrusiak leg. (MCTP 40092); 2$, same locality, 29.vi.1998, same collector (MCTP 40094); 1 same locality, 24.iii.1992, same col¬ lector (MCTP 40122); 1(5, same locality, 28.vii.1995, C.B. Kotzian & L. Indrusiak leg. (MCTP 40100); 1$, same locality, 14.X.2004, C.B. Kotzian leg. (MCTP zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 319-344 337 40091); 1$, same locality, 03.iv.2013, L. Indmsiak leg. (MCTP 39419); 16', 1$, same locality, 24.V.2007, A. Aued & E. Felzmamm leg. (MCTP 40103); 1$, 30. vii. 2000, L. Indmsiak leg. (MCTP 40090); 1$, same lo¬ cality, 15.vii.l998, C.B. Kotzian & L. Indmsiak leg. (MCTP 40102); 1$, same locality, 22.V.1996, same col¬ lector (MCTP 40097); 1$, same locality, 20.x. 1995, same collector (MCTP 40098); 3(6, 1$, same locality, 19.vi.l998, C. Kotzian & M. Monteiro leg. (MCTP 41324); 1 same locality, 17.xii.l992, L. Indmsiak & M. Monteiro leg. (MCTP 41323); 1$, same locality, 21. xii.1993, L. Indmsiak leg. (MCTP 41322); 1$, same lo¬ cality, 17.iii.l998, C. Kotzian & L. Indmsiak leg. (MCTP 41326); 1(6, 3$, same locality, 21.viii.l994, biology stu¬ dents leg. (MCTP 41325); 1(6, 1$, same locality, 26. viii. 1992, E. Eang & E. deques leg. (MCTP 40096); 3(6, 1$, same locality, 30.ix.l998, E. Indmsiak & M. Mon¬ teiro leg. (MCTP 40095); 1(6, same locality, 15.x. 1998, C. Kotzian & E. Indmsiak leg. (MCTP 40104); Ij, same locality, 10.x. 1979, D. Eink leg. (MCN 9548); 3$, same locality, 02.xi.l985, A.D. Brescovit leg. (MCN 14565); 2(6, same locality, 31.viii. 1994, R.G. Buss leg. (MCTP 10254); 5j, Montenegro, 29°41'20"S, 51°27'39'’W, 01. ix. l977 (MCN 6362); Ij, same locality, 06.X.1977, T. Arigony leg. (MCN 6808); Ij, same locality, 29.ix.1977, A.A. Else leg. (MCN 8180); 1(6, Santa Cruz do Sul, 29°43'04"S, 52°25'33"W, 14.viii.l994, A.A. Else leg. (MCN 33985); 1(6, Novos Cabrais, 29°46'59"S, 52°58T8"W, 01.xi.2008, R.G. Buss leg. (MCTP 27971); 1(6, same locality, ll.ix.2008, same collector (MCTP 28090); Ij, Alegrete, 29°47'26"S, 55°47'43"W, 28. viii. 1984, M.A.E. Marques leg. (MCN 9717); 1(6, Mo- rungava, 29°51T1''S, 50°54'31"W, 02.ii. 1992, A.D. Bres¬ covit leg. (MCN 23177); 1(6, 3$, Cachoeira do Sul (Porteira 7), 30°01T8.65"S, 52°55'3.70''W, 31.X.1992, R.G. Buss leg. (MCTP3485); 1$, Ij, same locality, 26. vii. 1992, same collector (MCTP 3488); 1 $, Porto Alegre, 30°01'58"S, 51°13'48"W, 01.xi.2009, I. Martins leg. (MCTP26258); 1(6,2$, Ij, Cachoeira do Sul,30°01'59"S, 52°54'00"W, 10.x. 1992, R.G. Buss leg. (MCTP 3491); 1(6,1 $, same locality, 26.ix. 1992, same collector (MCTP 3493); 1(6, 1$, same locality, 27.ix.1992, same collector (MCTP 3487); Ij, Porto Alegre (Morro Santana), 30°02'34"S, 51°08'39"W, 01.ix.l984, A.A. Else leg. (MCN 12546); Ij, Viamao (Estagao Fitotecnica de Viamao), 30°02'51"S, 51°00'53"W, 12.viii.l994, A.A. Else leg. (MCTP 5245); Ij, same locality, 30.iii.l976 same collector (MCN 5860); 1(6, Viamao, 30°04'51"S, 51°01'22"W, 22.xi.1995, A.A. Else leg. (MCTP 12246); 1(6, Guaiba, 30°06'50"S, 51°19'30"W, 04.X.1995, (MCTP 12334); 1$, Sao Sepe, 30°09'50"S, 53°34'18"W, iv.l999, E.C. Costa leg. (MCTP 40099); 1(6, same locality, 03.X.2002, C.B. Kotzian leg. (MCTP 41321); 1$, Cachoeira do Sul (Fazenda das Pedras), 30°12'50"S, 52°50'21"W, 27.x. 1992, R.G. Buss leg. (MCTP 3490); 1$, same locality, 09.ix. 1992, same collector (MCTP 3486); 1(6, same locality, 30.xi.l993, same collector (MCTP 4251); 1$, Cachoeira do Sul (Capanezinho), 30°18'35"S, 52°58'50"W, 29.viii.1992, R.G. Buss leg. (MCTP 3492); 1(6, 2$, same locality, 17.X.1992, same collector (MCTP 3489); 2$, Viamao (Morro Fortaleza), 30°20'45"S, 51°01'35"W, 22.xi.1992, A. Braul leg. (MCTP 2656); 2j, Quarai, 30°23'03"S, 56°26'56"W, 24- 28.V.1991, A. Braul leg. (MCTP 461); 4j, Quarai (Estan- cia Sao Roberto), 30°25T4"S, 55°51'53"W, 07.ii.l978, J.W. Thome leg. (MCN 7779). ARGENTINA: 7(6, 3$, Jujuy, Parque Nacional Calilegua, 23°38'20"S, 64°34T7"W, 23-24.ix.l995 (MACN-Ar 19229); 1(6, Mi- siones, El Pinalito, 25°58'59"S, 53°53'59"W, xi.l954, Schiapelli di Carlo leg. (MACN-Ar 19095); 1$, Tucuman, Raco, 26°38T5"S, 65°22'43"W (MACN-Ar 19102); Ij, Arroyo Yabebiry (Ruta Nacional 12), 27°17'05"S, 55°32'01'’W, vii.1980, P. Goloboff leg. (MACN-Ar 19100); 1$, Santa Maria, 27°53'39"S, 55°21'20"W, vii. 1954, M.J. Viana leg. (MACN-Ar 19097); 1 $, same locality, x.1953, M.J. Viana & Schiape- li de Carlo leg. (MACN-Ar 3804); 3(6, same locality, xii.1952, J. Viana leg. (MACN-Ar 3535); Ij, same locali¬ ty, xi.l952, J. Viana leg. (MACN-Ar 19071); Ij, same locality, 1954, M. Belgrano & Schiapelli de Carlo leg. (MACN-Ar 19096); 1 $, same locality, xii.1947, J. Viana leg. (MACN-Ar 2455); 2$, same locality, xi.l952, J. Vi¬ ana leg. (MACN-Ar 3534); 1(6, Entre Rios, Rosario del Tala, 32°18'00"S, 59°08'00"W, 20.xi.l988, M. Ramirez leg. (MACN-Ar 19091). URUGUAY: 1$, Eavalleja, Cerro Arequina, 34°17'09"S, 55°16'05"W, 03.xii.l997, A.A. Else leg. (MCTP 12677$ Diagnosis. Females of S. lucida resemble those of S. ex- cavata sp. nov. and S. marmorata sp. nov. by their short and rounded opisthosomal projections (Figs 2F, 12A). They can be distinguished by shorter copulatory ducts and spermathecae with just a median twisted constriction instead of many chambers (Fig. 12D, F). Males are sim¬ ilar to those of S. excavata sp. nov. by the colour pattern of the opisthosoma and the shape of its posterior projec¬ tions. However, males of S. lucida have a roundish RTA with a discrete basal branch (Fig. 13D, F), narrower pars pendula, and a shorter embolus emerging from tegulum at five o’clock (Fig. 13C, E). Description. Female: Prosoma yellowish, with cephalic ridges delimited by lines of small papules; needle-shaped setae concentrated on the median area of prosoma, be¬ ing the largest ones disposed right back of the PEE (Fig. 12B). Eegs yellowish, with femora I bearing three equal¬ sized needle-shaped macrosetae on their mesial surface (Fig. 12A); both the anterior tibiae and metatarsi (I and II) ventrally armed with five pairs of ventral macrosetae; tibiae I also bear a pair of smaller macrosetae along their mesial surface (Fig. 12A). Opisthosoma light-yellow with a median black stain on its anterior portion; posteri¬ or projections stout, obtuse and vertically oriented (Fig. 12A). Epigynal plate wide, depressed on the median field; posterior folds thick and converging in the middle to form a septum (Fig. 12C, E). zse.pensoft.net 338 Miguel Machado et al.: Taxonomic review of Neotropical Sidymella Figure 12. Female of Sidymella lucida a habitus, dorsal b prosoma, anterior c, e epigynum, ventral d, f epigynum, dorsal. zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 319-344 339 Figure 13. Male of Sidymella lucida a habitus, dorsal b prosoma, anterior c, e left palp, ventral view d, f left palp, retrolateral view (black arrow indicates the basal branch of the RTA). zse.pensoft.net 340 Miguel Machado et al.: Taxonomic review of Neotropical Sidymella Measurements: eyes diameters and eyes interdistances: AME 0.05, ALE 0.07, PME 0.05, PEE 0.05, AME-AME 0.11, AME-ALE 0.05, PME-PME 0.09, PME-PLE 0.17, MOQ length 0.29, MOQ width 0.23; leg formula: 1-2^- 3: leg I - femur 3.10/ patella 1.28/ tibiae 3.16/ metatarsus 2.32/ tarsus 0.84/ total 10.70; II - 2.00/ 0.96/ 1.64/ 1.48/ 0.56/ 6.64; III - 0.80/ 0.52/ 0.68/ 0.44/ 0.40/ 2.84; IV - 1.04/ 0.60/ 0.68/ 0.52/ 0.36/ 3.20. Total body length 4.68; prosoma 1.96 length, 1.76 wide; opisthosoma length 2.72 (considering the projections); clypeus 0.27 height; ster¬ num 1.08 length, 0.84 width; endites 0.44 length, 0.28 width; labium 0.28 length, 0.32 width. Male: Prosoma and legs as in female; opisthosoma predominantly yellow with lateral whitish lines, posterior projections dark (Fig. 13A, B). Palp flattened dorsoven- trally and elongated; cymbium distally narrowed; tegu- lum rounded with scaled surface (Fig. 13C-F). Measurements: eyes diameters and eyes interdis¬ tances: AME 0.05, ALE 0.07, PME 0.05, PEE 0.05, AME-AME 0.11, AME-ALE 0.05, PME-PME 0.09, PME-PLE 0.17, MOQ length 0.29, MOQ width 0.23; leg formula: 1-2-4-3: leg I - femur 2.56/ patella 0.88/ tibiae 2.48/ metatarsus 2.12/ tarsus 0.72/ total 8.76; II - 1.68/ 0.60/ 1.36/ 1.28/ 0.56/ 5.48; III - 0.64/ 0.32/ 0.56/ 0.28/ 0.36/ 2.16; IV- 0.84/ 0.44/ 0.48/ 0.24/ 0.36/ 2.36. Total body length 3.00; prosoma 1.40 length, 1.20 wide; opisthosoma length 1.60 (considering the projections); clypeus 0.15 height; sternum 0.58 length, 0.76 width; endites 0.27 length, 0.13 width; labium 0.17 length, 0.15 width. Distribution. ARGENTINA: Entre Rios, Jujuy, Mis- siones, and Tucuman. BRAZIL: Parana, Rio de Janeiro, Rio Grande do Sul, and Santa Catarina. COLOMBIA: Cali and Nova Granada. ECUADOR: Loja and Tungura- hua. URUGUAY: Lavalleja (Fig. 16). Sidymella marmorata Machado & Guzati, sp. nov. http;//zoobank.org/0EB278EC-9D36-4D68-8D52-9CE838B66501 Figures 2E, 14 Type material. Holotype ECUADOR: Quito (Road to Santo Domingo), 0°19'3.83"S, °59T9.95"W, L. Pena leg. (MCZ 133399). Paratypes. ECUADOR: 1 $, Qui¬ to (Road to Santo Domingo), 0°19'3.83"S, °59'19.95"W, 24.ii.1965, L. Pena leg. (MCTP 42654); COLOM¬ BIA: 1 Cundinamarca (Chia, Alto de Yerbabuena), 4°5r53.13"N, 74°3'3.30'’W, 09.X.2005, K. Lara & X. Marquinez leg. (ICN 7578) Other material examined. COLOMBIA: 1 Cundina¬ marca, Zipacon, 04°45'44"N, 74°22'46"W, 01.i.2011, D. Luna & C. Romero leg. (ICN-Ar 6379); 1 Ij, Boyaca, 05°41'14''N, 73°26'9"W, 01.viii.2003 (MPUJ_ENT); 1$, Narino (La Planada), 01°11'16"N, 78°3'19"W, 29.ii.1991, C. Valderrama leg. (LPN 226); 1$, Valle del Cauca (Chicoral), 03°39'24"N, 76°41'18"W, iii.2005. J. Cabra leg. (MUSENUV 28231); 1$, Quindio (Bue- navista), 04°21'59"N, 75°44'37"W, 13.ii.2010 (CIUQ 9663); 1$, Cundinamarca (Universidade Javeriana), 04°37'44"N, 74°03'51"W, 03.xi.l995 (lAvH 1157); 1$, Quindio (Bengala), 04°40'0"N, 75°39'0"W, 05.V.2003, A. Ardila leg. (MAUQ 1691); 3$, 2j, Cundinamar¬ ca (Alto de Yerbabuena), 04°51'55"N, 74°0r27'’W, ix.2005, K. Lara & X. Marquinez leg. (ex. ICN-Ar 7578); 1$, Cundinamarca (Suesca), 05°06'12"N, 73°47'56"W, 06.X.2013 (ICN-Ar 7637); 1$, Caldas (Sa- mana), 05°36'0"N, 75°2'0''W, 23.xi.1995, V. Rueda & H. Pineros leg. (lAvH 180518); 1 Boyaca, (Villa de Ley¬ va), 05°40'21"N, 73°27'42"W, 09.vi.2001, L. Benavides & J. Pinzon leg. (ICN-Ar 1249). Etymology. The epithet means marbled or overlaid with marble and refers to the reticulated colour pattern of the opisthosoma. Diagnosis. Females of S. marmorata sp. nov. resemble those of S. excavata sp. nov. by the large body size and the general shape of opisthosoma with an anterior concavity and short posterior projections (Figs 2E, 14A). However, females of S. marmorata sp. nov. bear numerous spini- form macrosetae on the mesial surface of femora I and II and five, instead of four, pairs of ventral macrosetae on tibiae I and II (Fig. 14A). Description. Female: Prosoma and legs I and II en¬ tirely orange while the posterior legs (III and IV) are yellowish. Opisthosoma predominantly yellow but with brownish irregular stains distributed randomly, giving a “marbled” aspect to the spider’s dorsum (Fig. 14A); opisthosoma projections short and stout and anal region projected backwards (Figs 2E, 14A). Epigynal plate elevated, with a wide septum and lateral folds concen¬ trically developed towards to the copulatory openings (Fig. 14C, E) copulatory ducts long, hyaline and coiled, leading to a pair of walnut-shaped spermathecae sub¬ divided in chambers and with a tubular glandular-head (Fig. 14D, F). Measurements: eyes diameters and eyes interdistances: AME 0.07, ALE 0.11, PME 0.07, PEE 0.07, AME-AME 0.17, AME-ALE 0.13, PME-PME 0.19, PME-PLE 0.19, MOQ length 0.37, MOQ width 0.17; leg formula: l-2^-3: leg I - femur 3.20/ patella 1.20/ tibiae 2.85/ metatarsus 1.80/ tarsus 0.88/ total 9.93; II - 2.40/ 1.03/ 2.12/ 1.60/ 0.77/ 7.92; III - 1.40/ 0.76/ 0.92/ 0.84/ 0.44/ 4.36; IV - 1.72/ 0.76/ 1.08/ 0.92/ 0.52/ 5.00. Total body length 5.44; prosoma 2.36 length, 2.32 wide; opisthoso¬ ma length 3.08; clypeus 0.33 height; sternum 1.03 length, 1.07 width; endites 0.58 length, 0.23 width; labium 0.33 length, 0.41 width. Male: Unknown. Distribution. ECUADOR: Quito. COLOMBIA: Cundi¬ namarca, Boyaca, Narino, Valle del Cauca, and Quindio (Fig. 15). zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 319-344 341 Figure 14. Female of Sidymella marmorata sp. nov. a habitus, dorsal b prosoma, anterior c, e epigynum, ventral d, f epigy- num, dorsal. zse.pensoft.net 342 Miguel Machado et al.: Taxonomic review of Neotropical Sidymella Figure 15. Distribution records of Sidymella excavata sp. nov. and Sidymella marmorata sp. nov. Figure 16. Distribution records of Sidymella furcillata and Sidymella lucida. zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 319-344 343 Figure 17. Distribution records of Sidymella kolpogaster and Sidymella longispina. Additional taxonomic changes Sidymella nigripes Mello-Leitao, 1947: 276 ((5'). Lise 1973: 16, figs 34-38 ($). Note. The holotype female was collected in Curitiba by Padre Moure in April 1942 and deposited in the In¬ stitute de Protegao do Patrimonio Natural da Secretaria da Agricultura do Estado do Parana under the number 2497. However, this institution has closed and its arthro¬ pod collection was transferred to unknown institutions. The repository of this species is unknown, even after we searched for it in all major Brazilian institutions with arachnological collections, and we assume that the holo¬ type of S. nigripes is lost. Therefore, we propose that the species should be considered a species inquirenda. Sidymella obscura Mello-Leitao, 1929: 65. Holotype juvenile collected in Serro da Caraga, Minas Gerais, Brazil (MNHN 8263, examined). Nomen dubium. Sidymella parallela Mello-Leitao, 1929: 64. Holotype juvenile collected in Salobro, Bahia, Brazil (MNHN 31114, examined). Nomen dubium. Sidymella spinifera Mello-Leitao, 1929: 66. Lise 1973: 21, figs 47-53. Syntypes, 1 immature S and 1 imma¬ ture $ collected in Serro da Caraga, Minas Gerais, Brazil (MNHN 8202, examined). Note. In the original work, Mello-Leitao (1929) men¬ tioned both specimens as “type”, but they are labelled as syntypes. Both specimens are juveniles that cannot be di¬ agnostic of any species. Nomen dubium. Acknowledgments We thank all curators who provided the material or access to the collections to examine the specimens listed above. We are also thankful to the staff of the Centro de microsco- pia e microanalises (CEMM) of the Pontificia Universi- dade Catolica do Rio Grande do Sul (PUCRS) for technical support, to Diego Galarraga Sugoniaev and Damian Ha- gopian for the images of live specimens, and to Dr Arthur Anker for the examination and photos of the holotypes of S. obscura, S. parallela, and S. spinifera. We thank the ref¬ erees and Steven Chu for comments and suggestions that helped to improve this manuscript. This study was financed in part by the Coordenagao de Aperfeigoamento de Pessoal de Nivel Superior, Brasil (CAPES) (Finance Code 001). References Benjamin SP, Dimitrov D, Gillespie RG, Hoimiga G (2008) Family ties: molecular phylogeny of crab spiders (Araneae: Thomisidae). Cladis- tics 24; 708-722. https;//doi.org/10.1111/j.l096-0031.2008.00202.x zse.pensoft.net 344 Miguel Machado et al.: Taxonomic review of Neotropical Sidymella Benjamin SP (2011) Phylogenetics and comparative morphology of crabs piders (Araneae; Dionycha, Thomisidae). Zootaxa 3080; 1-108. https://doi.org/10.11646/zootaxa.3080.1.1 Bonaldo AB, Lise AA (2001) A review of the Neotropical spider genus Stephanopoides (Araneae, Thomisidae, Stephanopinae). Biocien- cias 9; 63-80. Keyserling E (1880) Die Spinnen Amerikas, I. Laterigradae. Bauer & Raspe (E. Kuster), Ntirnberg, 283 pp. https;//doi.org/10.5962/bhl. title.64832 Lise AA (1973) Contribuipao ao conhecimento do genero Sidyma no Brasil, com descripao de uma nova especie (Araneae-Thomisidae). Iheringia43: 3-47. Lise AA (1981) Tomisideos Neotropicais V: Revisao do genero Ono- colus Simon, 1895 (Araneae, Thomisidae, Stephanopinae). Iheringia 57: 3-97. Machado M, Teixeira RA, Lise AA (2015) Taxonomic notes on the crab spider genus Tobias Simon, 1895 (Araneae,Thomisidae, Stepha¬ nopinae). Zootaxa 4034(3); 565-576. https://doi.org/10.11646/zoo- taxa.4034.3.8 Machado M, Teixeira RA, Lise, AA (2017) Cladistic analysis supports the monophyly of the Neotropical crab spider genus Epicadus and its senior synonymy over Tobias (Araneae; Thomisidae). 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Eibrairie Encyclopedique de Roret, Paris, 1084 pp. https://doi.org/10.5962/bhl.title.47654 Strand E (1942) Miscellanea nomenclatorica zoologica et palaeontolog- ica. X. Folia Zoologica et Hydrobiologica 11; 386-402. Wheeler WC, Coddington JA, Crowley EM, Dimitrov D, Goloboff PA, Griswold CE, Hormiga G, Prendini E, Ramirez MJ, Sierwald P, Almei- da-Silva E, Alvarez-Padilla F, Amedo MA, Benavides ER, Benjamin SP, Bond JE, Grismado CJ, Hasanf E, Hedin M, Izquierdo MA, Ea- barque EM, Eedford J, Eopardo E, Maddison WP, Miller JA, Piacentini LN, Platnick NI, Polotow D, Silva-Davila D, Scharff N, Szuts T, Ubick D, Vink CJ, Wood ITM, Zhang J (2017) The spider tree of life: phylog¬ eny of Araneae based on target-gene analyses from an extensive taxon sampling. Cladistics 33: 574-616. https://doi.org/10.llll/cla.12182 World Spider Catalog (2019) World Spider Catalog. Natural His¬ tory Museum Bern. Version 20.0. http://wsc.nmbe.ch [accessed March 2019] zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 345-360 | DOI 10.3897/zse.95.34069 ^yEHSPFT A new cryptic species of Hyphessobrycon Durbin, 1908 (Characiformes, Characidae) from the Eastern Amazon, revealed by integrative taxonomy Erick Cristofore Guimaraes^’^’^, Ptoella Silva de Brito^’^’^, Leonardo Manir Feitosa^, Luis Fernando Carvalho Costa^, Felipe Polivanov Ottoni^’^’^’® 1 Universidade Federal do Maranhao, Programa de Pds-Graduagao em Biodiversidade e Biotecnologia da Amazonia Legal. Av. dos Portiigiieses 1966, Cidade Universitdria do Bacanga, CEP 65080-805, Sdo Lids, MA, Brazil 2 Universidade Federal de Pernambuco, Programa de P6s-Graduagdo em Biologia Animal. Av. Professor Moraes Rego 1235, Cidade Universitdria, CEP 50670-901, Recife, PE, Brazil 3 Universidade Federal do Maranhao, Departamento de Biologia, Laboratorio de Genetica e Biologia Molecular, Av. dos Portugueses 1966, Cidade Universitdria do Bacanga, CEP 65080-805, Sdo Lids, MA, Brazil 4 Universidade Federal do Maranhao, Laboratorio de Sistemdtica e Ecologia de Organismos Aqudticos, Centro de Ciencias Agrdrias e Ambientais, Campus Universitdrio, CCAA, BR-222, KM 04, S/N, Boa Vista, CEP 65500-000, Chapadinha, MA, Brazil 5 Universidade Federal do Maranhao, Programa de Pds-Graduagao em Biodiversidade e Conservagdo. Av. dos Portugueses 1966, Cidade Universitdria do Bacanga, CEP 65080-805, Sdo Lids, MA, Brazil 6 Universidade Federal do Maranhdo, Programa de Pds-graduagdo em Ciencias Ambientais, Centro de Ciencias Agrdrias e Ambientais, Campus Universitdrio, CCAA, BR-222, KM 04, S/N, Boa Vista, CEP 65500-000, Chapadinha, MA, Brazil http://zoobank.org/E45AD907-EFD2-46B9-B056-31250BC8BFEE Corresponding author; Erick Cristofore Guimardes (erick.ictio@yahoo.com.br) Academic editor; Nicolas Hubert ♦ Received 25 February 2019 ♦ Accepted 7 May 2019 ♦ Published 12 June 2019 Abstract Hyphessobrycon caru sp. nov. is described based on five different and independent methods of species delimitation, making the hy¬ pothesis of this new species supported by an integrative taxonomy perspective. This new species has a restricted distribution, occur¬ ring just in the upper Pindare river drainage, Mearim river basin, Brazil. It is a member of the rosy tetra clade, which is characterized mainly by the presence of a dark brown or black blotch on dorsal fin and absence of a midlateral stripe on the body. Hyphessobrycon caru sp. nov. is distinguished from the members of this clade mainly by the shape of its humeral spot, possessing few irregular in¬ conspicuous vertically arranged chromatophores in the humeral region, or sometimes a very thin and inconspicuous humeral spot, and other characters related to teeth count, and color pattern. The phylogenetic position of the new species within the rosy tetra clade was based on molecular phylogenetic analysis using sequences of the mitochondrial gene cytochrome oxidase subunit 1. In addition, a new clade (here termed Hyphessobrycon micropterus clade) within the rosy tetra clade is proposed based on molecular data, com¬ prising H. caru sp. nov., H. micropterus, H. piorskii, and H. simulatus, and with H. caru sp. nov. and H. piorskii recovered as sister species. Our results suggest cryptic speciation in the rosy tetra clade and, more specifically, in the H. micropterus clade. We recom¬ mend the use of integrative taxonomy for future taxonomic revisions and species descriptions when dealing with species complexes and groups containing possible cryptic species. Key Words bPTP, DNA barcoding, rosy tetra clade, species complex, Stethaprioninae Copyright Erick Cristofore Guimaraes etai. This is an open access articie distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the originai author and source are credited. 346 Erick Cristofore Guimaraes et al.: A new cryptic species of Hyphessobrycon... Introduction Hyphessobrycon Durbin, 1908 is a species-rich charac- id genus comprising about 160 valid species (Fricke et al. 2019). It is widely distributed along the river basins of the Neotropical region, from southern Mexico to the La Plata River basin in northeastern Argentina (Car¬ valho and Malabarba 2015; Garcia-Alzate et al. 2017; Guimaraes et al. 2018). The genus was first proposed as a subgenus of Hemigrammus Gill, 1858 by Durbin in Eigenmann (1908), differing from the latter only by the absence of scales covering the caudal-fin. Hyphessobry¬ con was reviewed by Eigenmann (1918, 1921) in a work which still constitutes the most comprehensive revision¬ ary studies on the genus. The large number of species in¬ cluded within Hyphessobrycon and the poor knowledge of the alpha and beta-taxonomy of species and species groups are among the major challenges for a more com¬ prehensive taxonomic study and phylogenetic analyses of the genus. It is widely known that Hyphessobrycon does not constitute a monophyletic group (Weitzman and Palmer 1997a; Mirande 2010, 2018; Oliveira etal. 2011; Carvalho and Malabarba 2015; Carvalho et al. 2017; Moreira and Eima 2017; Betancur-R. et al. 2018; Guim¬ araes et al. 2018). Nevertheless, groups of species have been proposed based primarily on similarities of color pattern and other external features (e.g. Weitzman and Palmer 1997a; Garcia-Alzate et al. 2008; Moreira and Eima 2017). Some of them are probably merely artificial operational assemblages to aid species identification, whereas others represent potential monophyletic groups, delimited by exclusive character states (e.g. Castro-Paz et al. 2014; Carvalho and Malabarba 2015; Guimaraes et al. 2018). Several genetic studies focusing on characoid fishes, such as Astyanax Baird & Girard, 1854 (e.g. Omelas-Gar- cia et al. 2008), Caenotropus Gunther, 1864 (e.g. Melo et al. 2014), Chilodus Muller & Troschel, 1844 (e.g. Melo et al. 2014), Curimatopsis Steindachner, 1876 (e.g. Melo et al. 2016a), Gymnocorymbus Eigenmann, 1908 (e.g. Be- nine et al. 2015), Hyphessobrycon (e.g. Castro-Paz et al. 2014, Guimaraes et al. 2018), Piabina Reinhardt, 1867 (e.g. Pereira et al. 2011), Prochilodus Agassiz, 1829 (e.g. Melo et al. 2016b), Nannostomus Gunther, 1872 (e.g. Ben- zaquem et al. 2015) and Tetragonopterus Bleeker, 1863 (e.g Melo et al. 2016c) have evidenced that some species may exhibit large discontinuities in their geographic dis¬ tribution patterns, with high genetic divergences, but little morphological variability among geographically isolated lineages. These results suggest that these groups may represent species complexes or cryptic species, that is, they might even including morphologically quite similar or undistinguishable species that are hidden and errone¬ ously classified (Brown et al. 1995; Bickford et al. 2006; Adams et al. 2014; Souza et al. 2018). Studies relying solely on morphology may be inadequate in recognizing species within groups including cryptic species (Guim¬ araes et al. 2018). Integrative studies, using more than one criteria, such as character-based, tree-based, genetic distance and coalescent-based approaches, especially in¬ cluding molecular data, are useful and powerful for the recognition of hidden and/or possible new species in such species complexes (Sytsma and Schaal 1985; Bickford et al. 2006; Goldstein and Desalle 2010; Padial et al. 2010; Adams et al. 2014; Costa-Silva et al. 2015; Souza et al. 2018; Ottoni et al. 2019). In this context of integrative taxonomy, the present study aims to investigate the diversity within the rosy tetra clade sensu Weitzman and Palmer (1997a). This clade comprises around 30 species, including some spe¬ cies of Hyphessobrycon and other allied species, that are appreciated as aquarium fishes due to their attrac¬ tive color patterns (e.g. Weitzman and Palmer 1997a, 1997b, 1997c, 1997d; Zarske 2008; Hein 2009; Guim¬ araes et al. 2018). This group has had its composition and name changed over the last decades, and a detailed taxonomic history is presented by Weitzman and Palmer (1997a). Two pre¬ vious papers (e.g. Castro-Paz et al. 2014; Guimaraes et al. 2018) applied molecular approaches to investigate the diversity of rosy tetra clade, and they suggested that its taxonomic resolution should be better investigated as it could include cryptic species or valid species which may have been synonymized. A new species of Hy¬ phessobrycon and member of the rosy tetra clade is de¬ scribed from the upper Pindare river drainage, Mearim river basin, a coastal river basin of the Eastern Amazon region, Brazil, based on both morphology and molecu¬ lar data. Furthermore, a new clade, within the rosy tetra clade, is proposed based on the phylogenetic tree topol¬ ogy presented. Materials and methods Taxa sampling, specimens collection, and preservation Individuals collected for this study were euthanized with a buffered solution of MS-222 at a concentration of 250 mg E“' for a period of 10 min or more until oper¬ cular movements completely ceased. Specimens select¬ ed for morphological analysis were fixed in formalin and left for 10 days, after which they were preserved in 70% ethanol. Molecular data were obtained from specimens that were euthanized, fixed, and preserved in absolute ethanol. Specimens for morphological analysis are listed in type and comparative material lists. Specimens for molecular approaches are listed in Table 1. We also re¬ trieved sequences from other species of Hyphessobry¬ con and allied species for a comparative analysis from the Barcode of Fife Database (BOED) and the National Center for Biotechnology Information (NCBI) databas¬ es (Table 1). zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 345-360 347 Table 1. List of species, specimens and their respective catalogue numbers, Region/state/country, and BOLD Systems and GenBank sequence accession numbers. Sequences available in the current study are in Bold. N° Species Catalogue number Region/state/country Accession no. 1 Hyphessobrycon erythrostigma INPA 37681-HERYl Tabatinga/Amazonas/Brazil HYP076-13 2 Hyphessobrycon erythrostigma INPA 37681-HERYlO Tabatinga/Amazonas/Brazil HYP077-13 3 Hyphessobrycon erythrostigma INPA 37681-HERY2 Tabatinga/Amazonas/Brazil HYP078-13 4 Hyphessobrycon erythrostigma INPA 37681-HERY3 Tabatinga/Amazonas/Brazil HYP079-13 5 Hyphessobrycon pyrrhonotus INPA 37672-TROlO Santa Isabel do Rio Negro/Amazonas/Brazil HYP040-13 6 Hyphessobrycon pyrrhonotus INPA 37672-TROll Santa Isabel do Rio Negro/Amazonas/Brazil HYP04M3 7 Hyphessobrycon pyrrhonotus - Barcelos/Amazonas/Brazil HYP157-13 8 Hyphessobrycon pyrrhonotus - Barcelos/Amazonas/Brazil HYP158-13 9 Hyphessobrycon socoiofi INPA_39530-6152 Barcelos/Amazonas/Brazil HYP131-13 10 Hyphessobrycon socoiofi INPA_39530-6155 Barcelos/Amazonas/Brazil HYP134-13 11 Hyphessobrycon socoiofi INPA_39530-6178 Barcelos/Amazonas/Brazil HYP135-13 12 Hyphessobrycon socoiofi INPA 39530-BCR8 Barcelos/Amazonas/Brazil HYP148-13 13 Hyphessobrycon copeiandi INPA_37683-TU1 Tabatinga/Amazonas/Brazil HYP094-13 14 Hyphessobrycon copeiandi INPA_37683-TU2 Tabatinga/Amazonas/Brazil HYP095-13 15 Hyphessobrycon copeiandi INPA_37683-TU3 Tabatinga/Amazonas/Brazil HYP096-13 16 Hyphessobrycon eques INPA_37678-IC2 Santarem/Para/Brazil HYP070-13 17 Hyphessobrycon eques INPA_37679-PE1 Macapa/Amapa/Brazil HYP071-13 18 Hyphessobrycon eques INPA_37680-AL1 Parintins/Amazonas/Brazil HYP072-13 19 Hyphessobrycon eques OL-0544 Bonito/Mato Grosso do Sul/Brazil DSMIS077-09 20 Hyphessobrycon epicharis INPA_37665-JUF1 Sao Gabriel da Gachoeira/Amazonas/Brazil HYP002-13 21 Hyphessobrycon epicharis INPA_37665-JUF3 Sao Gabriel da Cachoeira/Amazonas/Brazil HYP004-13 22 Hyphessobrycon epicharis INPA_37665-JUF4 Sao Gabriel da Cachoeira/Amazonas/Brazil HYP005-13 23 Hyphessobrycon epicharis INPA_37665-JUF8 Sao Gabriel da Cachoeira/Amazonas/Brazil HYP006-13 24 Hyphessobrycon compressus CINV-NEC7411 Flores Magon/Campeche/Mexico FYPM054-10 25 Hyphessobrycon compressus ECOCH Hatie ville/Belize/Belize MXV765-15 26 Hyphessobrycon compressus ECOCH Hatie ville/Belize/Belize MXV766-15 27 Hyphessobrycon compressus ECOCH Hatie ville/Belize/Belize MXV767-15 28 Hyphessobrycon bentosi INPA_37684-5939 Barcelos/Amazonas/Brazil HYP097-13 29 Hyphessobrycon bentosi INPA_37684-5940 Barcelos/Amazonas/Brazil HYP098-13 30 Hyphessobrycon bentosi INPA_39527-BA1 - HYP116-13 31 Hyphessobrycon bentosi INPA_39527-BA2 - HYP117-13 32 Hyphessobrycon bentosi CICCAA02349 Santarem/Para/Brazil MK240339 33 Hyphessobrycon bentosi CICCAA02350 Santarem/Para/Brazil MK240340 34 Hyphessobrycon bentosi CICCAA02351 Santarem/Para/Brazil MK240341 35 Hyphessobrycon simuiatus MHNG 2743.087 Pisiemoengo/Commev^ijne/Suriname GBOL761-15 36 Hyphessobrycon simuiatus MHNG 2743.087 Pisiemoengo/Commev^ijne/Suriname GBOL762-15 37 Hyphessobrycon simuiatus - Sinnamary/Cayenne/French Guiana GBOL1771-17 38 Hyphessobrycon simuiatus MHNG 2735.007 Sinnamary/Gayenne/French Guiana GBOL3296-18 39 Hyphessobrycon simuiatus MHNG 2757.080 Kourou/Cayenne/French Guiana GBOL3298-18 40 Hyphessobrycon simuiatus MHNG 2759.026 Kav^/Gayenne/French Guiana GBOL3300-18 41 Hyphessobrycon simuiatus MHNG 2759.026 Kav^/Cayenne/French Guiana GBOL3301-18 42 Hyphessobrycon simuiatus MHNG 2759.035 Regina/ Cayenne/French Guiana GBOL3302-18 43 Hyphessobrycon cf. swegiesi INPA_37668-JAR3 Sao Gabriel da Cachoeira/Amazonas/Brazil HYP026-13 44 Hyphessobrycon cf. swegiesi INPA_37668-JAR4 Sao Gabriel da Cachoeira/Amazonas/Brazil HYP027-13 45 Hyphessobrycon cf. swegiesi INPA_37668-JAR5 Sao Gabriel da Gachoeira/Amazonas/Brazil HYP028-13 46 Hyphessobrycon cf. swegiesi INPA_37668-JAR7 Sao Gabriel da Cachoeira/Amazonas/Brazil HYP030-13 47 Hyphessobrycon micropterus - Varzea da Palma/Minas Gerais/Brazil BSB287-10 48 Hyphessobrycon micropterus - Varzea da Palma/Minas Gerais/Brazil BSB288-10 49 Hyphessobrycon micropterus - Varzea da Palma/Minas Gerais/Brazil BSB289-10 50 Hyphessobrycon micropterus - Varzea da Palma/Minas Gerais/Brazil BSB290-10 51 Hyphessobrycon piorskii CIGGAA00725-1 Chapadinha/Maranhao/Brazil MF765796 52 Hyphessobrycon piorskii GICCAA00726-1 Chapadinha/Maranhao/Brazil MF765797 53 Hyphessobrycon piorskii CICGAA01650-1 Barrel rinhas/Maranhao/Brazil MG791915 54 Hyphessobrycon piorskii CICCAA01651-1 Barrel rinhas/Maranhao/Brazil MG791914 55 Hyphessobrycon piorskii Cl CGAA02164-1 Godo/Maranhao/Brazil MK240337 56 Hyphessobrycon piorskii CICCAA02164-4 Codo/Maranhao/Brazil MK240338 57 Hyphessobrycon caru CICCAA00748-1 Buriticupu/Maranhao/Brazil MH338230 58 Hyphessobrycon caru CICCAA00749-1 Buriticupu/Maranhao/Brazil MH338231 59 Hyphessobrycon caru CICCAA02300-1 Buriticupu/Maranhao/Brazil MH338232 60 Hyphessobrycon caru CICCAA02301-1 Buriticupu/Maranhao/Brazil MH338233 61 Pristeiia maxiiiaris - - KU568982.1 62 Pristeiia maxiiiaris - - KU568981.1 63 Pristeiia maxiiiaris - Marlborough/Pomeroon-Supenaam/Guyana TZGAA025-06 64 Pristeiia maxiiiaris - Santa Cruz/Barima-Waini/Guyana TZGAAl 78-06 65 Moenkhausia hemigrammoides INPA38532-PR1 Guyana HYPlOl-13 66 Moenkhausia hemigrammoides INPA_38532-PR2 Guyana HYP102-13 67 Moenkhausia hemigrammoides INPA_38532-PR3 Guyana HYP103-13 68 Hyphessobrycon panamensis STRI-05303 Cocle/Panama BSFFA760-07 69 Hyphessobrycon fiammeus LBPV-40464 Biritiba-Mirim/Sao Paulo/Brazil FUPR988-09 zse.pensoft.net 348 Erick Cristofore Guimaraes et al.: A new cryptic species of Hyphessobrycon... Morphological analysis Measurements and counts were made according to Fink and Weitzman (1974), with exception of the scale rows below lateral line, which were counted to the insertion of pelvic-fin. Vertical scale rows between the dorsal-fin ori¬ gin and lateral line do not include the scale of the median predorsal series situated just anterior to the first dorsal-fin ray. Counts of supraneurals, vertebrae, procurrent cau¬ dal-fin rays, unbranched dorsal and anal-fin rays, bran- chiostegal rays, gill-rakers, premaxillary, maxillary, and dentary teeth were taken only from cleared and stained paratypes (C&S), prepared according to Taylor and Van Dyke (1985). The four modified vertebrae that constitute the Weberian apparatus were not included in the vertebrae counts and the fused PUl + U1 was considered as a single element. Osteological nomenclature follows Weitzman (1962). Institutional abbreviations follow Fricke and Es- chmeyer (2019), with addition of LIOP.UFAM Colegao Ictiologica do Laboratorio de Ictiologia e Ordenamento Pesqueiro do Vale do Rio Madeira da Universidade Fed¬ eral do Amazonas. DNA extraction, amplification, and sequencing DNA was extracted from fin clips using Wizard Genomic DNA Purification kit (Promega) according to the man¬ ufacturer’s protocol. Fragments of the cytochrome c oxidase subunit 1 gene (hereafter COI) from mitochon¬ drial DNA were amplified, using the universal primers designed by Ward et al. (2005) for fish. Polymerase chain reactions (PCR) comprised a total volume of 15 pi con¬ taining lx Polymerase buffer, 1.5 mM MgCl 2 , 200 pM dNTP, 0.2 uM of each primer, lU of Taq Polymerase (In- vitrogen), 100 qg of DNA template, and ultrapure water. The PCR cycles were as follows: 2 min at 94 °C, fol¬ lowed by 35 cycles of 94 °C for 30s, 54 °C for 30s, and 72 °C for 1 min, and 10 min at 72 °C. Amplicons were purified using Illustra GFX PCR DNA and Gel Purifica¬ tion Kit (GE Healthcare Systems) and sequenced using the forward primer by an outsourced sequencing service at the University of Sao Paulo, using BigDye Terminator kit 3.1 Cycle Sequencing kit in ABI 3730 DNA Analyser (Applied Biosystems). Data partition, evolution models, and alignment The dataset included the following gene: COI (680 Base pairs, BP). Sequences were aligned using ClustalW (Chenna et al. 2003). The DNA sequences were trans¬ lated into amino acids residues to test for the absence of premature stop codons or indels using the program MEGA 7 (Kumar et al. 2016). In the alignment, gaps were coded with a dash (-) and missing data with a ques¬ tion mark (?), but during analyses, both were treated as missing data. Measure Substitution Saturation tests were performed in DAMBE5 (Xia 2013) according to the al¬ gorithm proposed by Xia et al. (2003). The best-fit evo¬ lutionary model (GTR+G) was calculated, using the cor¬ rected Akaike Information Criterion (AICc) determined by the JModelTest 2.1.7 (Darriba et al. 2012). Species concept, species delimitation, and diagnoses The unified species concept is herein adopted by ex¬ pressing the conceptual definition shared by all tra¬ ditional species concepts, “species are (segments of) separately evolving metapopulation lineages”, disen¬ tangling operational criterion elements to delimit taxa from species concepts (de Queiroz 2005, 2007). Ac¬ cording to this concept, species are treated as hypothet¬ ical units and could be tested by the application of dis¬ tinct criteria (species delimitation methods) (de Queiroz 2005, 2007). It allows for any criteria to separately pro¬ vide evidence about species limits and identities, inde¬ pendently from other criteria (de Queiroz 2005, 2007). However, evidence corroborated from multiple opera¬ tional criteria is considered to produce stronger support for hypotheses of lineage separation (de Queiroz 2007; Goldstein and Desalle 2010), a practice called “integra¬ tive taxonomy” (Dayrat 2005; Goldstein and Desalle 2010; Padial et al. 2010). Five distinct and independent operational criteria for species delimitation, based on morphological and molec¬ ular data, were implemented here: Population Aggrega¬ tion Analysis (Davis and Nixon 1992) (hereafter PAA); DNA barcoding, as proposed by Hebert et al. (2003a, 2003b, 2004 a, 2004b) (hereafter DBG); a tree-based method as proposed by Wiens and Penkrot (2002) (here¬ after WP, following Sites and Marshall 2003); a charac¬ ter-based DNA barcoding as proposed by Desalle et al. (2005) (hereafter CBB); and a coalescent species delimi¬ tation method termed the Bayesian implementation of the Poisson tree processes (hereafter bPTP, following Zhang et al. 2013). All species delimitation methods here adopt¬ ed, except PAA, were performed on cytochrome c oxi¬ dase subunit 1 (COI) sequences, as it is a mitochondrial gene with fast evolutionary rate, suitable for single locus species delimitation approaches (Avise 2000). Population aggregation analysis (PAA) The PAA (Davis and Nixon 1992) is a character-based method, in which species are delimited by unique combi¬ nation of morphological character states occurring in one or more populations (Costa et al. 2014). The morpholog¬ ical data was based on both examined material and liter¬ ature (e.g. Steindachner 1882; Meek 1904; Eigenmann, 1908; Durbin 1909; Eigenmann 1915; Ahl 1937; Fowler 1943; Gery 1960, 1961, 1964, 1977; Gery and UJ 1987; Burgess 1993; Planquette et al. 1996; Weitzman and zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 345-360 349 Palmer 1997a, 1997b, 1997c, 1997d; Zarske 2008; Hein 2009; Lima et al. 2013; Zarske 2014; Carvalho and Mala- barba2015; Carvalho et al. 2017; Guimaraes et al. 2018). Traditional DNA barcoding (DEC) and Phylogenetic analysis We used the Kimura-2-parameters model (K2P) (Kimura 1980) to estimate the pairwise genetic distances between species in MEGA 7 software (Kumar et al. 2016). We used DnaSP V. 6 (Rozas et al. 2003) to estimate the number of variable sites and haplotypes. A Bayesian inference-based phylogenetic (BI) tree was estimated in MrBayes (Huelsen- beck and Ronquist 2001) plugin in Geneious 9.0.5 to recon¬ struct the evolutionary relationships among terminals using General Time Reversible (GTR+G) as evolutionary model. Bayesian tree inference was based in a chain length of 10 million, a bum-in length of 500,000 generations subsam¬ pling trees every 10,000 generations. We used a sequence of Hyphessobrycon flammem Myers, 1924 as outgroup. Wiens and Penkrot analysis (WP) WP is based on the direct inspection of haplotype trees generated from the phylogenetic analysis having as termi¬ nals at least two individuals (haplotypes) of each focal spe¬ cies. In this method, the term “exclusive” is used instead of monophyletic, as the term monophyly is considered inap¬ plicable below the species level (Wiens and Penkrot 2002). Clustered haplotypes with concordant geographic distribu¬ tion forming mutual and well supported clades (exclusive lineages) are considered strong evidence for species dis¬ crimination (absence of gene flow with other lineages). When haplotypes from the same locality fail to cluster to¬ gether, there is potential evidence for gene flow with other populations (Wiens and Penkrot 2002). Statistical support for clades is assessed by the posterior probability, consid¬ ered as significant at values about 0.95 or higher (Alfaro and Holder 2006). When only one haplotype (specimen) from one putative population was available, the species delimitation was based on the exclusivity of the sister clade of this single haplotype, supported by significant values, allowing us to perform the test in populations with only one haplotype (Wiens and Penkrot 2002). In addition, the method allows recognition of nonexclusive lineages as species since their sister clades are exclusive and support¬ ed by significant values (Wiens and Penkrot 2002). Character-based DNA barcoding (CBB) The CBB is similar to the population aggregation anal¬ ysis proposed by Davis and Nixon (1992), but directed to nucleotides as an alternative method for diagnosing taxa through DNA barcodes, as the original method is based on subjective cut-off distance measures to species designation (Hebert et al. 2003a, 2003b, 2004a, 2004b). This method delimits species based on a unique combi¬ nation of nucleotides within a site shared by individuals of the same population or group of populations. In addi¬ tion, species were diagnosed by nucleotide substitutions following Costa et al. (2014). Optimization of nucleotide substitutions among lineages of the Hyphessobrycon mi- cropterus clade were obtained from the Maximum Parsi¬ mony topology, using TNT 1.5 (Goloboflf and Catalano 2016). Maximum Parsimony analysis (MP) was obtained with the following parameters: traditional search, tree bi¬ section reconnection branch swapping (TBR), 1 random seed, setting random taxon-addition replicates to 1,000, multi-trees in effect, collapsing branches of zero length, characters equally weighted, and 10,000 trees saved per replication. MP tree branch support was given by boot¬ strap analysis (Felsenstein 1985), using a heuristic search with 1,000 replicates and the same settings used in the MP search, saving a maximum of 1,000 trees in each random taxon-addition replicate. The analysis was rooted on Hy¬ phessobrycon flammeus Myers, 1924. Each nucleotide substitution is represented by its relative numeric position determined through sequence alignment with the complete mitochondrial genome of Astyanax paranae Eigenmann 1914 (KX609386.1:5503-7062 - mitochondrion complete genome), followed by the specific nucleotide substitution in parentheses. The results of this analysis are presented in Suppl. material 1: Box 1 and molecular diagnosis section. Bayesian implementation of the poisson tree proeesses (bPTP) The bPTP is a coalescent phylogeny-based species delim¬ itation method aimed at delimiting species based on single locus molecular data (Zhang et al. 2013). An advantage of bPTP is that it does not need an ultrametric calibration like other coalescent approaches, avoiding errors and comput¬ er intensive processes (Zhang et al. 2013). The method relies on the number of substitutions between haplotypes and assumes that more molecular variability is expected between species than within a species (Zhang et al. 2013). In our analysis the dataset was reduced to include only unique haplotypes from the species of the H. micropterus clade. Outgroups were restricted to Hyphessobrycon ben- tosi Durbin, 1908 and Hyphessobrycon copelandi Durbin 1908. Sequences were aligned using ClustalW (Chenna et al. 2003). The best-fit evolutionary model (GTR+G) for the reduced dataset was calculated using the corrected Akaike Information Criterion (AICc) determined by the JModelTest 2.1.7 (Darriba et al. 2012). The input phylo¬ genetic tree was performed in MrBayes 3.2.6 (Ronquist et al. 2012), with the following parameters: independent runs of two Markov chain Monte Carlo (MCMC) runs of four chains each for 3 million generations and sampling frequency of 1,000. The bPTP analysis was performed in the Exelixis Eab’s web server http://species.h-its.org/ptp/, following the default parameters except for a 20% burn in. zse.pensoft.net 350 Erick Cristofore Guimaraes et al.: A new cryptic species of Hyphessobrycon... Results Hyphessobrycon earn sp. nov. http://zoobank.org/3BC35EBB-E138-4E24-A06E-DF985F015ED5 Figures 1, 2a; Table 2 Holotype. CICCAA 02286, 22.2 mm SL, Brazil, Mara- nhao state, Buriticupu municipality, Buritizinho river, Pindare river drainage, Mearim river basin, 04°22'52"S, 46°30'35'’W, 24 Jan. 2017, Guimaraes E. C., Brito P. S. Paratypes. All from Brazil, Maranhao state: CICCAA 00706, 37, 15.9-25.4 mm SL; CICCAA 0709, 12 C&S, 15.1-20.6 mm SL; LIOP.UFAM 1009, 1, 16.2 mm SL collected with holotype. CICCAA00707, 3, 17.2-22.1 mm SL, Buriticupu municipality, Buritizinho river. Pin- dare river drainage, Mearim river basin, 4°25'45"S, 46°29'41"W, 24 Jan. 2017, Guimaraes E. C., Brito P. S. CICCAA00708, 2, 19.9-21.6 mm SL, Buriticupu munic¬ ipality, Buritizinho river, Pindare river drainage, Mearim river basin, 04°19'45"S, 46°29'46"W, 24 Jan. 2017, Gui¬ maraes E. C., Brito P. S. UFRJ11745, 1, 22.4 mm SL, Buriticupu municipality, Buritizinho river, Pindare river drainage, Mearim river basin, 04°19'45"S, 46°29'46"W, 24 Jan. 2017, Guimaraes E. C., Brito P. S. Diagnosis (PAA). The new species Hyphessobrycon caru sp. nov. differs from most of its congeners, except members of the rosy tetra clade, by the presence of a dark brown or black blotch on dorsal-fin (vs absence) and ab¬ sence of a midlateral stripe on the body (vs presence). The new species differs from most of its congeners in the rosy tetra clade by possessing few irregular inconspicuous vertically arranged chromatophores in the humeral region, or sometimes a very thin and inconspicuous humeral spot (Fig. 2a) [vs inconspicuous vertically elongated humeral spot in H. hasemani Fowler, 1913,7/. piorskii Guimaraes, De Brito, Feitosa, Carvalho-Costa, Ottoni, 2018 (Fig. 2b); approxi¬ mately rounded humeral spot in H. erythrostigma (Fowler, 1943), H. jackrobertsi Zarske, 2014,77 minor Dmhm, 1909, 77 pando Hein, 2009, 77 paepkei Zarske, 2014, 77 pyrrho- notus Burgess, 1993, 77 roseus (Gery, 1960), 77 socolofi Weitzman, 1977, and 77 sweglesi (Gery, 1961) (Fig. 2c); hu¬ meral spot horizontally or posteriorly elongated in 77 epich- aris Weitzman & Palmer, 1997,77 khardinae Zarske, 2008, and 77 wemeri Gery & Uj, 1987 (Fig. 2d); conspicuous hu¬ meral spot at least on males in 77 copelandi Durbin, 1908, 77 (Steindachner, 1882), 77 haraldschultzi Travassos, 1960, 77 micropteriis (Eigenmann, 1915), 77 megalopter- us (Eigenmann, 1915), 77 simulatus (Gery, 1960) and 77 takasei Gery, 1964 (Fig. 2e); and absence of humeral spot in 77 compressus (Meek, 1904), 77 dorsalis Zarske, 2014, 77 georgettae Gery, 1961, 77 pulchripinnis Ah\, 1937, and 77 rosaceus Durbin, 1909 (Fig. 2f)]. Furthermore, the new species differs from 77 bentosi Durbin, 1908, 77 erythrostigma, 77 pyrrhonotus, 77 ro¬ saceus, and 77 socolofi by presenting only one tooth in the outer row of premaxillary, and this unique tooth just slightly displaced from inner row [vs two or more teeth, displaced from the inner row]; from 77 hasemani and 77 micropterus by the dorsal-fin spot located approximately at the middle of the fin’s depth, not reaching its tip (vs spot located approximately at the middle of the fin’s depth, reaching its tip in adults); from 77 hasemani by present¬ ing tri to unicuspid teeth in the inner row of premaxil¬ lary and dentary (vs tricuspid or pentacuspid teeth); from 77 piorskii by having the anal-fin profile usually nearly straight (vs anal-fin profile usually falcate). In addition, 77 caru sp. nov. is easily distinguished from Pristella maxillaris (Ulrey, 1894), Moenkhausia hemigrammoides Gery, 1965, and Hemigrammus unilineatus (Gill, 1858) by the absence of a black oblique stripe or band on the anterior portion of the anal-fin (Fig. 1) (vs presence). Description. Morphometric data of holotype and para¬ types are presented in Table 2. Body small (with maximum SL of 25.4 mm), compressed, moderately deep, greatest body depth slightly anterior to dorsal-fin base. Lateral body profile straight and downward directed from the end of dorsal-fin to adipose-fin, straight or slightly convex be¬ tween later point and origin of dorsal most procurrent cau¬ dal-fin ray. Dorsal profile of head convex from upper lip to vertical through eye; predorsal profile of body roughly straight, dorsal-fin base slightly convex, posteroventrally inclined; ventral profile of head convex from lower jaw to pelvic-fin origin. Ventral profile of body straight or slight¬ ly convex from pelvic-fin origin to anal-fin origin; straight and posterodorsally slanted along anal-fin base; and slightly concave on caudal peduncle. Jaws equal, mouth terminal, anteroventral end of dentary protruding. Maxilla reaching vertical to anterior margin of pupil. Table 2. Morphometric data {N = 45) of Hyphessobrycon caru sp. nov. SD: Standard deviation. Holotype Paratypes Mean SD Standard length 22.2 14.8-25.4 18.9 - Percentages of standard length Depth at dorsal-fin origin (body depth) 37.3 33.1-38.5 35.2 1.1 Snout to dorsal-fin origin 53.7 49.4-55.0 51.7 1.2 Snout to pectoral-fin origin 29.5 28.2-32.3 29.9 1.0 Snout to pelvic-fin origin 46.0 43.6-48.8 45.6 1.0 Snout to anal-fin origin 62.5 58.5-64.0 61.0 1.3 Caudal peduncle depth 12.3 8.5-12.3 10.3 0.8 Caudal peduncle length 11.7 9.5-12.7 11.2 0.8 Pectoral-fin length 23.2 16.5-23.7 19.6 1.9 Pelvic-fin length 20.6 14.1-20.5 17.4 1.4 Dorsal-fin base length 15.2 12.9-15.7 14.3 0.8 Dorsal-fin height 32.2 27.9-34.1 30.8 1.5 Anal-fin base length 32.4 26.4-32.7 29.6 1.3 Eye to dorsal-fin origin 37.5 34.4-38.8 37.3 0.9 Dorsal-fin origin to caudal- fin base 55.1 50.6-56.1 53.4 1.1 Head length Percentages of head length 29.8 27.4-31.1 29.3 1.0 Horizontal eye diameter 39.2 35.4-43.6 39.2 1.7 Snout length 24.4 17.3-24.3 21.5 1.8 Least interorbital \A/idth 29.1 22.4-30.7 27.2 1.8 Upper ja\A/ length 37.8 33.1-42.5 37.4 2.1 zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 345-360 351 Figure 1. Hyphessobrycon earn sp. nov., CICCAA 02286, holotype, 22.2 mm SL; Brazil: Maranhao state; Buritizinho River, Pin- dare river drainage, Mearim river basin. Figure 2. Humeral spot of; a Hyphessobrycon earn, holotype, CICCAA 02286 b H. piorskii, holotype, CICCAA 00695 c H. pyr- rhonotus, holotype, MZUSP 45714 d H. werneri, holotype, MZUSP 42365 e H. eques, CICCAA 00300 fH. compressus, paratype, MHNG 2181.076. zse.pensoft.net 352 Erick Cristofore Guimaraes et al.: A new cryptic species of Hyphessobrycon... 1.8mm 1mm Figure 3. Hyphessobrycon caru sp. nov., jaw suspensory, CICCAA 00697, paratype, 19.3 mm SL; premaxillary (a), maxillary (b), and dentary (c). Premaxillary teeth in two rows. Outer row with one unicuspid or tricuspid tooth, just slightly displaced from inner row; inner row with 6(5), 7(6), or 8(1) tricuspid teeth and one unicuspid tooth. Maxilla with 3(2) tricus¬ pid teeth and two unicuspid teeth, 4(3) tricuspid teeth and two unicuspid teeth or 5(7) tricuspid teeth. Dentary with five (10) or six (1) larger tricuspid teeth followed by one smaller tricuspid teeth 5(2), 6(2), 7(3), and 8(5) smaller unicuspid teeth (Fig. 3). Scales cycloid, three to eight radii strongly marked, cir- culi well-marked anteriorly, weakly marked posteriorly; lat¬ eral line incompletely pored, with 5(1), 6(2), 7(24), 8(14), or 9(4) perforated scales. Longitudinal scales series includ¬ ing lateral-line scales 31(1), 32(7), 33(14), 34(13), 35(3), or 36(7). Longitudinal scales rows between dorsal-fin origin and lateral line 5(3), 6(32), or 7(10). Horizontal scale rows between lateral line and pelvic-fin origin 4(43) or 5(2). Scales in median series between tip of supraoccipital spine and dorsal-fin origin 10(9), 11(12), 12(21), or 13(3). Cir- cumpeduncular scales 11(6), 12(35), 13(2), or 14(2). Dorsal-fin origin at midbody. Base of last dorsal-fin ray at vertical through first third of anal-fin. Dorsal-fin rays ii + 9(48), hi + 9(5), ii + 10(4). First dorsal-fin pte- rygiophore main body located behind neural spine of 4* vertebrae. Adipose-fin present. Anal-fin origin aligned with vertical line through middle of dorsal-fin, between 6* and 8* dorsal-fin rays base. Anteriormost anal-fin pterygiophore inserting posterior to haemal spine of 11* vertebrae. First anal-fin ray in vertical through the mid¬ dle of dorsal-fin (with about 7* or 8* ray base). Anal- fin hi + 22(10) or hi + 23(47); anal-fin origin aligned with vertical line through middle of dorsal-fin (between base of 6* and 8* dorsal-fin rays); Anal-fin profile nearly straight; Anal-fin rays with a sexually dimorphic pattern, which is absent in females, described below. Pectoral-fin rays 12(57) total rays. Tip of pectoral-fin surpassing pel¬ vic-fin base. Pelvic-fin rays 8(57) total rays, surpassing anal-fin origin. Pelvic-fin rays with a sexually dimorphic pattern, which are absent in females, described below. Caudal-fin forked, upper and lower lobes similar in size. Principal caudal-fin rays 11+10(50) or 10+9(7); dorsal procurrent rays 8(2), 9(8) or 11(2) and ventral procurrent rays 7(4) or 8(8). Branchiostegal rays 4(12). First gill arch with 1(11), 2(1) hypobranchial, 11(1), 12(10), or 13(1) ceratobranchi- al, 1(12) on cartilage between ceratobranchial and epibran- chial, and 5(1) or 6(11) epibranchial gill-rakers. Supraneu- rals 3(2), 4(9), or 5(1). Total vertebrae 28(2) or 29(10). zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 345-360 353 - 47 ° 36 ' . 45 ° 54 ' - 44 “ 12 ' - 42 ° 30 ' - 40 ° 48 ' - 39 “ 6 ' 0 ° 0 ' - 1“42 • 3'=24 - 5 ° 6 ' Figure 4. Geographical distribution of Hyphessobrycon caru sp. nov. and H. piorskii. Red star denotes Holotype and blue cir¬ cles denote paratypes of H. caru sp. nov., and white star denotes Holotype and the black circles denote the known distribution of H. piorskii. Colour in alcohol. Ground coloration light yellowish brown. Humeral region with few irregular inconspicuous vertically arranged chromatophores, sometimes very thin and inconspicuous humeral spot. Flank with chromato¬ phores homogeneously scattered, more concentrated on posterior region to humeral spot, posterior region of dor¬ sal-fin base origin and below mid-portion of trunk, between anal-fin origin and caudal peduncle. Ventral region lacking dark-brown chromatophores. Dark-brown chromatophores present on head and more concentrated on dorsal portion, becoming sparser on cheek and preopercular regions. Dorsal-fin ground coloration hyaline, with conspicuous black or dark-brown spot located on anterior portion of fin, reaching about 6* ray, approximately between one-half to two-thirds of fin depth. Anal and caudal-fins hyaline. Cau¬ dal-fin with a darker, usually dark brown, posterior margin and on its base. Adipose-fin hyaline to light brown, with dark-brown or black chromatophores more concentrated on its dorsal portion, depending on the specimen preserva¬ tion state. Pectoral and pelvic-fins hyaline; pelvic-fin with variable amounts of dark-brown pigmentation remaining depending on the specimen preservation state. Sexual dimorphism. Mature males with small bone hooks on anal and pelvic-fin rays. Bone hooks absent on females. Anal-fin presenting bone hooks from 4*, or 5* rays to the last ray. Number of hooks variable, increas¬ ing from the first to the last rays. Pelvic-fin presenting 2"^, 3rd, 4th^ Qj. 5th j-^yg Qj. j smaller hooks. Etymology. The specific epithet honors the term ''Caru''. Caru is the name of an area (about 70.000 ha) inhabited by Brazilian native tribes from the ethnicities Guaja and Guajajara. People from this area use the Tupi language and have sulFered consequences of European coloniza¬ tion and are under threat due to the pressure for explora¬ tion of the protected territory. Geographic distribution. Hyphessobrycon caru sp. nov. has a restricted geographic distribution, being known only from the upper Pindare river drainage, Mearim riv¬ er basin, in the state of Maranhao, northeastern Brazil (Fig. 4). This species was never collected in the lower portions of this river drainage during 8 years of field trips conducted by EG and PB, including about 15 expeditions. Molecular diagnosis (CBB). Hyphessobrycon caru sp. nov. belongs to the H micropterus clade possessing 20 synapo- morphic nucleotide substitutions: COI 73 (C^T), COI 88 (T^C), COI 217 (C^T), COI 274 (C^T), COI 298(C^T), COI 334 (C^G), COI 338 (T^C), COI 370 (A^G), COI 418 (A^G), COI 433 (C^T), COI 439 (C^A), COI zse.pensoft.net 354 Erick Cristofore Guimaraes et al.: A new cryptic species of Hyphessobrycon... FUPR988-09 / Hyphessobrycon flammeus L FYPM054-10/Hyphessobrycon compressus MXV767-15 / Hyphessobrycon compressus I I- MX\/765-15 / Hyphessobrycon compressus ^ MX\/766-15 / Hyphessobrycon compressus 6SFFA760-07 / Hyphessobrycon panamensis 0.04 0 a 0.64 GBOL761 -15/ Hyphessobrycon simulatus GBOL762-15 / Hyphessobrycon simulatus GBOL1771-17 / Hyphessobrycon simulatus GBOL3296-18 / Hyphessobrycon simulatus GBOL3298-18 / Hyphessobrycon simulatus GBOL3300-18 / Hyphessobrycon simulatus GBOL3301 -18 / Hyphessobrycon simulatus GBOL3302-18 / Hyphessobrycon simulatus BSB287-10 / Hyphessobrycon micropterus BSB289-10 / Hyphessobrycon micropterus BSB290-10 / Hyphessobrycon micropterus BSB291 -10 / Hyphessobrycon micropterus 0J2 j~ MG791914/ Hyphessobrycon piorskii MG791915/ Hyphessobrycon piorskii r MF675796 / Hyphessobrycon piorskii ^ MF675797 / Hyphessobrycon piorskii MK240337 / Hyphessobrycon piorskii MK240SS8 / Hyphessobrycon piorslai 0.52r MH338230 / Hyphessobrycon caru ~ *1 |-t MH33823'\ / Hyphessobrycon caru I L— MH338233 / Hyphessobrycon caru FIYP076-13 / Hyphessobrycon erythrostigma FIYP077-13 / Hyphessobrycon erythrostigma FIYP078-13 / Hyphessobrycon erythrostigma HYP079-13 / Hyphessobrycon erythrostigma HYP040-13 / Hyphessobrycon pyrrhonotus HYP041-13 / Hyphessobrycon pyrrhonotus FIYP157-13 / Hyphessobrycon pyrrhonotus HYP158-13 / Hyphessobrycon pyrrhonotus F1YP131-13/ Hyphessobrycon socolofi FIYP134-13/ Hyphessobrycon socoloh FIYP135-13/ Hyphessobrycon socolofi FIYP 148-13 / Hyphessobrycon socolofi MK240339 / Hyphessobrycon bentosi MK240340 / Hyphessobrycon bentosi MK240341 / Hyphessobrycon bentosi HYP097-13 / Hyphessobrycon bentosi FIYP098-13 / Hyphessobrycon bentosi FIYPI16-13/ Hyphessobrycon bentosi HYP117-13/ Hyphessobrycon bentosi o 0 ) Q. CD O S S' s C/5 MH338232 / Hyphessobrycon caru FIYP070-13 / Hyphessobrycon eques FIYP071-13/ Hyphessobrycon eques HYP072-13/ Hyphessobrycon eques DSMIS077-09 / Hyphessobrycon eques r FIYP094-13/ Hyphessobrycon copeiandi I HYP095-13 / Hyphessobrycon copeiandi I FIYP096-13 / Hyphessobrycon copeiandi HYP002-13 / Hyphessobrycon epicharis HYP004-13 / Hyphessobrycon epicharis FIYP005-13 / Hyphessobrycon epicharis HYP006-13 / Hyphessobrycon epicharis r HYP026-13 / Hyphessobrycon cf. sweglesi _0j9^ HYP027-13 / Hyphessobrycon cf. sweglesi I* FIYP029-13 / Hyphessobrycon cf sweglesi I HYP030-13 / Hyphessobrycon cf sweglesi KU568981.1 / Pristella maxillaris KU568982.1 / Pristella maxillaris TZGAA025-06 / Pristella maxillaris TZGAA178-06 / Pristella maxillaris HYP 101-13 / Moenkhausia hemigrammoides HYP 103-1 3 / tJloenkhausia hemigrammoides HYP102-13 / Moenkhausia hemigrammoides Bl 99 100 •- 95 C 67 84 D 99 92 BSB289-10 / Hyphessobrycon micropterus BSB290-10 / Hyphessobrycon micropterus BSB291-10 / Hyphessobrycon micropterus BSB287-10 / Hyphessobrycon micropterus GBOL3302-18 / Hyphessobrycon simulatus GBOL3296-18 / Hyphessobrycon simulatus GBOL3300-18 / Hyphessobrycon simulatus QBOL32Q8-^ 8 /Hyphessobrycon simulatus GBOL3301-18 / Hyphessobrycon simulatus -GBOL1771-17/ Hyphessobrycon simulatus -GBOL762-15 / Hyphessobrycon simulatus -GBOL 761-15 / Hyphessobrycon simulatus -MH388231 / Hyphessobrycon caru -MH338233 / Hyphessobrycon caru -MH338232 / Hyphessobrycon caru -MH338230 / Hyphessobrycon caru -MK240338 / Hyphessobrycon piorskii -MK240337 / Hyphessobrycon piorskii -MG791914 / Hyphessobrycon piorskii -MG791915 / Hyphessobrycon piorskii -MF675797 / Hyphessobrycon piorskii -MF675796 / Hyphessobrycon piorskii MP t CD Co Co o O' I O o Figure 5. Phylogenetic tree based on Bayesian Inference (BI). Numbers above branches are posterior probability values. Posterior probability value supporting the Hyphessobrycon micropterus clade is indicated in green (haplotypes marked with a green bar); posterior probability value supporting the H. caru sp. nov. lineage under WP method is indicated in red (haplotypes marked with a red bar); and the other species (lineages) under WP method, within this clade, are indicated in black, b Strict consensus phylogenetic tree based on Maximum Parsimony (MP), obtained from the 38 most parsimonious trees, in which 587 characters were constant, 20 variable but parsimony-uninformative, and 248 parsimony-informative (total length 833, consistency index 0.489, retention index 0.901). The image is focusing on the Hyphessobrycon micropterus clade. Numbers above branch are bootstrap values and letters below branches correspond to nucleotide substitutions, listed in Suppl. material 1: Box 1, corresponding to the CBB method. Green circle indicating Hyphessobrycon micropterus clade, red circle H. caru sp. nov., blue circles the other congeners within the clade, and black circle the clade H. caru sp. nov. + H. piorskii. zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 345-360 355 HYP094-13/ Hyphessobrycon copelandi - ■ ■ MK240339 / Hyphessobrycon bentosi [ • • ■ HYP116-13 / Hyphessobrycon bentosi • ■ ■ HYP 117-13 / Hyphessobrycon bentosi • • •HYP097-13 / Hyphessobrycon bentosi [ GBOL3298-18 / Hyphessobrycon simuiatus GBOL3300-18 / Hyphessobrycon simuiatus • ■ ■ GBOL3296-18 / Hyphessobrycon simuiatus [ GBOL762-15 / Hyphessobrycon simuiatus GBOL1771-17 / Hyphessobrycon simuiatus ■ ■ ■ GBOL761-15 / Hyphessobrycon simuiatus L- ■ • GBOL3302-18 / Hyphessobrycon simuiatus { ■ MF675796 / Hyphessobrycon piorskii r ‘ ■ MK240337 / Hyphessobrycon piorskii •'' MK240338 / Hyphessobrycon piorskii MG791914 / Hyphessobrycon piorskii ■MG791915 / Hyphessobrycon piorskii MH338231 / Hyphessobrycon caru MH338233 / Hyphessobrycon caru ^ • MH338230 / Hyphessobrycon caru ■ MH338232 / Hyphessobrycon caru [ BSB290-10 / Hyphessobrycon micropterus BSB291 -10 / Hyphessobrycon micropterus • ■ ■ BSB289-10 / Hyphessobrycon micropterus • ■ ’ BSB287-10 / Hyphessobrycon micropterus 0.10 Figure 6. Species delimitation tree generated by the Bayesian Poisson Tree Processes (bPTP) model, using a fragment of the mitochon¬ drial gene COL The blue lines indicate branching processes among species, while red lines indicate branching processes within species. 457 (A^G), COI 469 (T^C), COI 478 (A^T), COI 559 (A^G), COI 562 (T^A), COI 592(A^G), COI 631 (A^T), COI 655 (A^C), COI 673 (A^C). It shares nine synapomorphic nucleotide substitutions with H. piorskii, which separate them from H. simuiatus and H. micropter¬ us-. COI 181 (A^C), COI 208 (A^G), COI 245 (C^T), COI 325 (T^C), COI 349 (T^C), COI 436 (A^T), COI 472 (A^G), COI 538 (C^T), COI 556 (T^C). In addi¬ tion, it has six unique nucleotide substitutions within the H. micropterus clade: COI 148 (C^T), COI 154 (C^T), COI 175 (T^C), COI 364 (G^A), COI 487 (T^C), COI 517 (A^G) (Fig. 5; Suppl. material 1: Box 1). DBC. COI sequences support the existence of a new spe¬ cies of Hyphessobrycon inhabiting the Pindare river basin in Maranhao state. After trimming, the final alignment yielded 680 base pairs with 159 polymorphic sites and 26 haplotypes. Average genetic distances were 18.3%, with the highest values between H. epicharis and H. erythrostig- ma (23.4%), while the lowest value (0.7%) was between H. pyrrhonotus and H. erythrostigma (Table 3). Hyphessobry¬ con caru sp. nov. is divergent on average 17.0% from the other taxa, with a minimum distance of 3.6% to H piorskii and a maximum of 21.8% to Pristella maxillaris (Table 3). WP and CBB. Both phylogenetic analysis based on BI and MP supported a clade comprising H. caru sp. nov., H. micropterus, H piorskii, and H. simuiatus, hereafter termed Hyphessobrycon micropterus clade, with maxi¬ mum posterior probability value and 99% bootstrap value in BI and MP, respectively. Hyphessobrycon caru sp. nov. formed a single exclusive lineage with maximum poste¬ rior probability value (posterior probability =1) and 99% bootstrap value in BI and MP, respectively. zse.pensoft.net 356 Erick Cristofore Guimaraes et al.: A new cryptic species of Hyphessobrycon... Table 3. Kimura-2 parameters pairwise genetic distances among species. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 1 H. erythrostigma 2 H. pyrrhonotus 0.007 3 H. socolofi 0.037 0.035 4 H. simulatus 0.192 0.194 0.179 5 H. micropterus 0.205 0.202 0.198 0.041 6 H. piorskii 0.188 0.185 0.178 0.063 0.064 7 H. caru 0.206 0.210 0.203 0.062 0.057 0.036 8 H. eques 0.175 0.178 0.163 0.154 0.157 0.160 0.160 9 H. copelandi 0.192 0.189 0.186 0.170 0.176 0.158 0.168 0.102 10 H. epicharis 0.234 0.230 0.220 0.170 0.175 0.189 0.190 0.187 0.197 11 H. bentosi 0.106 0.103 0.112 0.197 0.194 0.205 0.204 0.195 0.220 0.219 12 H. cf. sweglesi 0.210 0.207 0.197 0.182 0.187 0.201 0.209 0.181 0.197 0.030 0.222 13 P. maxillaris 0.225 0.232 0.206 0.201 0.211 0.220 0.218 0.194 0.213 0.180 0.202 0.183 14 M. hemigrammoides 0.219 0.219 0.209 0.179 0.185 0.178 0.179 0.203 0.211 0.196 0.221 0.199 0.166 15 H. compressus 0.214 0.218 0.215 0.202 0.208 0.198 0.208 0.198 0.201 0.212 0.212 0.212 0.203 0.215 16 H. panamensis 0.210 0.213 0.209 0.202 0.199 0.179 0.187 0.215 0.229 0.221 0.213 0.218 0.204 0.208 0.145 17 H. flammeus 0.201 0.201 0.198 0.169 0.174 0.186 0.204 0.203 0.205 0.192 0.200 0.189 0.206 0.200 0.170 0.196 These species delimitation analysis (WP and CBB) have identical results, delimitating four species within the Hyphessobrycon micropterus clade: H. earn sp. nov., H. micropterus, H. piorskii, and H. simulatus (Fig. 5a, b). The nucleotide substitutions supporting these four lin¬ eages within the H. micropterus clade, and the nucleotide substitutions supporting this clade are presented in Fig¬ ure 5b and Suppl. material 1: Box 1. The combination of nucleotide substitutions diagnosing H. caru sp. nov. are presented in the molecular diagnosis section. bPTP. This species delimitation analysis also indicates four lineages (species) within the Hyphessobrycon micro¬ pterus clade: H. caru sp.n., H. micropterus, H piorskii, and H. simulatus (Fig. 6). This outcome was similar to the aforementioned results. The species included as out¬ groups {H. bentosi and H. copelandi) were also supported as independent lineages. Discussion Currently molecular techniques are frequently useful for solve species complexes and discover cryptic species (e.g. Bickford et al. 2006; Costa and Amorim 2011; Pereira et al. 2011; Adams etal. 2014; Costa-Silva 2015; Costa et al. 2012, 2014, 2017; Amorim 2018; Guimaraes et al. 2018; Ottoni et al. 2019) and could be an excellent complement for traditional taxonomy (Kekkonen and Hebert 2014). DNA barcoding has demonstrated to be very efficient for delimiting species of Hyphessobrycon, mainly in groups with little morphological variation (i.e., cryptic species) (see Castro-Paz et al. 2014; Guimaraes et al. 2018), pref¬ erably when applied together with other species delimi¬ tation methods, such as PAA, DBC, CBB, bPTP, and WP in an integrative taxonomy perspective (Guimaraes et al. 2018). The recognition of different genetic patterns and lineages in groups with very similar morphology has been a common pattern in the tree of eukaryotic life. This is observed particularly often in species-rich genera, such as in several Neotropical fishes (e.g. Pereira et al. 2011; Roxo et al. 2012; Castro-Paz et al. 2014; Melo et al. 2014, 2016a; Benzaquem et al. 2015; Benine et al. 2015; Ottoni et al. 2019). DNA techniques can help to uncover mor¬ phological hidden diversity (Bickford et al. 2006; Adams et al. 2014), delimiting a putative population or group of populations as an independent lineage (species), and, subsequently, through a more meticulous analysis of mor¬ phological features, morphological differences between cryptic species can be found. The large number of the described Hyphessobrycon species (about 160 spp.), with new species described ev¬ ery year, reveal an astonishing diversity within the genus. During the past 10 years, about 50 new species have been described (Fricke et al. 2019). However, historically Hy¬ phessobrycon species have been described only on the ba¬ sis of morphological features, including differences in the pigmentation patterns and teeth numbers and morpholo¬ gy, using few individuals per species (e.g. Steindachner 1882; Eigenmann 1915; Zarske 2008, 2014; Bragan^a et al. 2015). Recently, DNA barcoding in characoid fishes has been used to discriminate species, identify new ones, and reveal that it is not always possible to differentiate species based solely on their morphology (Ornelas-Gar- cia et al. 2008; Pereira et al. 2011; Castro-Paz et al. 2014; Melo et al. 2014, 2016a; Benine et al. 2015). Our results suggest a cryptic speciation in the rosy tetra clade, more specifically in a new clade here defined, the Hyphessobrycon micropterus clade, including H. caru sp. nov., H. micropterus, H piorskii, and H. simulatus, so far only known from the Pindare, Itapecuru, Munim, Pre- gui^as, and Sao Francisco river drainages of Brazil and the coastal river basins of French Guiana and Suriname (Guimaraes et al. 2018; Brito et al. 2019; Fricke et al. 2019; this study). The clade proposed here is supported by high node support values (maximum posterior prob¬ ability value and 99% of bootstrap value in BI and MP, respectively). In addition, this clade was corroborated by 20 synapomorphic nucleotide substitutions (Fig. 5; Sup¬ pl. material 1: Box 1). zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 345-360 357 Hyphessobrycon earn sp. nov. is herein described within the Hyphessobrycon micropterus clade based on five different and independent methods of species delimitation (PAA, DBC, WP, CBB and bPTP), charac¬ terized by different criteria and assumptions. Hyphesso¬ brycon caru sp. nov. is distinguished from all its conge¬ ners by a combination of unambiguous morphological character states [see Diagnosis (PAA)]. In our Bayesian phylogenetic analysis (Fig. 5A), haplotypes of H. caru sp. nov. formed a single exclusive clade with maximum posterior probability value (posterior probability = 1) (WP). Furthermore, the COI average genetic distance of H. caru sp. nov. when compared with the other taxa herein analyzed was 19.6% and its minimum COI ge¬ netic distance was 3.6% to H. piorskii (DBC). Consid¬ ering this value, the threshold of H. caru sp. nov. would be greater than that inferred by delimitations among Neotropical fish species (2% according to Pereira et al. 2011). Moreover, H. caru sp. nov. was also molecular- ly diagnosed by six synaphomorphic nucleotide sub¬ stitutions (Fig. 5b; Suppl. material 1: Box 1), as well as, by a combination of other nucleotide substitutions (see CBB - molecular diagnosis), and corroborated by a bPTP analysis. Thus, it makes the hypothesis of this new species stronger from an integrative taxonomy per¬ spective (see Dayrat 2005; de Queiroz 2007; Goldstein and Desalle 2010; Padial et al. 2010). Therefore, we recommend the use of integrative taxonomy for future taxonomic revisions and species descriptions when dealing with species complexes and groups containing possible cryptic species. Comparative material Hyphessobrycon amandae: UFRJ 1557, 5 spems, Jussara municipality, Goias state, Brazil. H. bentosi: MCZ 20842, 1 spem (Syntype), Obidos municipality, Para state, Brazil. H. bifasciatus: UFRJ 0068, 6 spems, Marataizes and Guarapari municipality, Espirito Santo state, Brazil. H. compressus. BMNH 1905.12.6.4-5, 2 spems (Paratypes), Oaxaca state. Mexico. H. copelan- di. CAS 42683, 1 spem (Syntype); MCZ 20771, 1 spem (Syntype), Tabatinga municipality, Amazonas state, Bra¬ zil. H. eques. CICCAA00715,4 spems (C&S); CICCAA 00710, 51 spems, Tombos municipality Carangola riv¬ er, Minas Gerais state, Brazil. H. erythrostigma. ANSP 70208, 1 spem (Holotype), Peru and Brazil. H. epich- aris. FMNHl00609, 1 spem (Paratype), Baria river, Am¬ azonas, Venezuela. H. haraldschultzi. CICCAA 00873, 20 spems, Ilha do Bananal municipality, Javaes river, Tocantins state, Brazil. H. hasemani. ANSP 39230, 1 spem (Holotype), Guajaramirim municipality, Madeira river, Rondonia state, Brazil. H. micropterus. FMN- HH 57916, 1 spem (Holotype), Sao Francisco river at Lagoa de Porto, Minas Gerais state, Brazil. H. piorskii: CICCAA 00695, 1 spem (Holotype); CICCAA 00430, 15 spems (Paratype); CICCAA 00431, 21 spems (Para¬ type); CICCAA 00696, 15 spems (Paratype); CICCAA 00697, 16 spems (C&S) (Paratype); CICCAA 00698, 6 spems, 1 spem (C&S) (Paratype); CICCAA 00750, 9 spems (Paratype); CICCAA01654, 1 spem (Paratype); CPUFMA 171664, 15 spems (Paratype); UFRJ 11553, 6 spems (Paratype), stream at the Anapurus municipality, Munim river, Maranhao state, Brazil. CICCAA 00089, 1 spem (C&S) (Paratype); CICCAA 00881, 1 spem (Para¬ type); CICCAA 01563, 1 spem (Paratype); stream at Mata de Itamacaoca, Chapadinha municipality, Munim river, Maranhao state, Brazil. CICCAA 01382, 5 spems (Paratype); CICCAA 02008, 12 (C&S) spems (Para¬ type), stream at Mata Fome, Barreirinhas municipality, Pregui^as river, Maranhao state, Brazil. H. pyrrhonotus. MZUSP 45714, 1 spem (Holotype), Erere river, Brazil. H. rosaceus. FMNH 52791, 1 spem (Holotype), Gluck Island, Essequibo River, Guyana. H werneri. MZUSP 42365, 1 spem (Holotype), Santa Maria do Para and Sao Miguel de Guama municipality, Guama river, Para state, Brazil. CICCAA 00751, 1 spem, Paragominas mu¬ nicipality, Candiru river, Para state, Brazil. H. socolofi. MZUSP 13181, 1 spem (Holotype), Barcelos municipal¬ ity, Negro river, Amazonas state, Brazil. Acknowledgements We thank Axel Makay Katz, Axel Zarske, Flavio Lima, Ingo Schindler, Oscar Lasso-Alcala and Ronald Fricke for providing useful literature; Vale S.A and Amplo Engenha- ria for the cession of part of the data analysed in this study; Raphael Covain and collaborators of the Project Gui-BOL Barcoding Guianese fishes for the cession of part of the data analysed in this study; Marcelo Rodrigues dos An- Jos and Wilson Costa for the loan and donation of mate¬ rial; Mark Sabaj Perez (CAS), James Maclaine (FMNH), Riedel Bettina (NMW) for providing photographs, x-ray images, and information on the type material of some spe¬ cies; FACEPE for providing the scholarship to LMF under the process IBPG-0089-2.05/17, CAPES and FAPEMA for providing the scholarship to PSB under the process 88887.159561/2017-00. This paper benefited from sugges¬ tions provided by P. Bragan^a and R. Collins. 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Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons. org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow us¬ ers to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Link: https://doi.org/10.3897/zse.95.34069.suppll zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 361-372 | DOI 10.3897/zse.95.33707 4>yEnsPFr. BERLIN Type specimens of Aplysiida (Gastropoda, Heterobranchia) in the Academy of Natural Sciences of Philadelphia, with taxonomic remarks Carlo M. Cunha^’^’^, Gary Rosenberg^ 1 Universidade Metropolitana de Santos. Ave. Conselheiro Nebias 536, 11045-002, Santos, SP, Brazil 2 Academy of Natural Sciences of Philadelphia, Drexel University. 1900 Benjamin Franklin Parkway, Philadelphia, PA 19103, USA 3 CAPES - Foundation, Ministry of Education of Brazil, Brasllia/DF 70040-020, Brazil http://zoobank.org/AA43756B-4521-4FA3-A9C3-ABB2CFFBCDC6 Corresponding author: Carlo M. Cww/za (carlomagenta@gmail.com) AcdidQYmc Q6.\ioY. Thomas von Rintelen ♦ Received 15 February 2019 ♦ Accepted 30 May 2019 ♦ Published 19 June 2019 Abstract The type specimens of 15 nominal species of Aplysiida (Gastropoda, Heterobranchia) at the Academy of Natural Sciences of Phil¬ adelphia are listed and their primary types are illustrated. Type materials described by the following authors are present; Henry A. Pilsbry (9 names), Angelo Heilprin (2), Charles N. E. Eliot (1), William H. Pease (2) and Elizabeth J. Eetson (1). Some taxonomic notes are provided. Aplysia perviridis (Pilsbry, 1895), comb, nov., Aplysia pilsbryi (Eetson, 1898), and Aplysia pulmonica Gould, 1852 are reinstated as valid. Key Words Mollusca, Anaspidea, Aplysiomorpha, morphology, new combinations, biodiversity. Curatorial methods, malacology, nomencla¬ ture, type specimens Introduction Since the publication of Pilsbry’s Manual of Conchology (1895, 1896a, 1896b), where a number of new taxa were described, little has been published about the types of Ap¬ lysiida (e.g. Valdes and Heros 1998). Systematic revisions of several aplysiidan groups are needed (Gosliner 1994; Valdes et al. 2006; Uribe et al. 2013), and information about type materials is essential for determining the sta¬ tus of species-group taxa. As the aplysiidan material that Pilsbry studied as well as that of other authors is housed at the Academy of Natural Sciences (ANSP), evaluation of the type materials held there is important. We follow Bouchet et al. (2017) for the name of the order; it has also been called Anaspidea and Aplysiomorpha. Material and methods To find putative type material of aplysiidans, we searched the ANSP collection database for items indicated as hav¬ ing a type status; this database includes digitized records from the handwritten ledgers. We also searched the dry collection for specimens indicated as type specimens on their labels and relevant literature (e.g. Pilsbry 1896; Bal¬ es 1960) for indications of type materials held by ANSP. Where ethanol-preserved material was verified as part of a type series, the animals were dissected by standard techniques, under a stereomicroscope, with the specimen immersed in ethanol. Digital photography of the whole animals was obtained with the specimen immersed in ethanol an acrylic container with rubber bottom. After Copyright Carlo M. Cunha, Gary Rosenberg. This is an open access articie distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the originai author and source are credited. 362 Cunha, C.M. & Rosenberg, G.: Type specimens of Aplysiida in the Academy of Natural... dissections, photographs were taken of the specimen carefully pinned into the container in the desired position. Digital photographs of shells were made using either a standard digital macro-lens system with single pho¬ tos of each view for specimens larger than ~10 mm in maximum dimension, or a computer-operated motorized StackShotTM system (Cognisys, Inc.), coupled with Hel¬ icon Remote and Helicon Focus software (HeliconSoft Ltd.), allowing digital combination (i.e., “stacking”) of multiple individual shots of each view for specimens less than 10 mm. The figures of shells are not shown to scale, but the maximum dimension of each specimen is given in the corresponding figure legend. The photographs of types published here are also available in the online database for the ANSP Mollusca collection. Catalogue numbers in the ANSP malacology collec¬ tion were originally assigned in a single sequence, started in 1889 by Henry Pilsbry. Around 1970, George M. Da¬ vis established a separate number sequence prefixed with “A” for alcohol-preserved material. Many samples with catalogue numbers in the original sequence were given new numbers in the A sequence. This renumbering affect¬ ed all of the nominal taxa of Aplysiida for which ANSP has type material in alcohol. In some cases, the shell is stored under the dry collection under the original number and body from which it was dissected is preserved in the alcohol collection under an “A” number. We have traced these associations to ensure that we have recognized the components of each lot. The list of nominal taxa is arranged alphabetically by the original epithet (in bold) with the taxon name written as in the original description, except that capitalization is adjusted to current standards. Institutional Acronyms: ANSP, Academy of Natural Science of Philadelphia, USA; CPIC, research collection of the Department of Biological Sciences of the Califor¬ nia State Polytechnic University, Pomona, USA; MCZ, Museum of Comparative Zoology, Harvard University, Boston, USA. Results and discussion Search of the ANSP collection, the collection database and relevant literature found potential type material of 18 names. Three of these proved to be false leads: a lot of Dolabrifera maillardi Deshayes, 1863 (ANSP 114205) from Reunion was erroneously flagged in the database as a type; one was a manuscript name of Pilsbry (ANSP 84337, A7044, current¬ ly Aplysia sp.) from Zanzibar [we do not mention the name to avoid creating a nude name]; and one labeled as “Co¬ type?” of Dolabrifera marmorea Pease (ANSP 247112), but Sowerby (1868), not Pease introduced this name. For 15 nominal taxa, we verified the presence of type material in ANSP. This includes holotypes for 3 taxa and syntypes for 11, along with 1 {Aplysia badistes) where we could not determine in the specimen was the holo- type or a paratype. The names were introduced by Henry A. Pilsbry (9), Angelo Heilprin (2), Charles N. E. Eliot (1), William H. Pease (2), and Elizabeth J. Eetson (1). The type localities are Western Atlantic (7), South Pacific Ocean (3), North Pacific Ocean (4), and Indian Ocean (2). Currently 18 valid species with 10 synonyms are recog¬ nized for Aplysiida in the Western Atlantic. This catalog is especially important to that region, as it represents 25% of the 28 nominal taxa. In the following section, the type material and current status of each nominal species is included. Two species were previously considered valid and 13 held as synon¬ ymous. Aplysia perviridis (Pilsbry, 1895) new comb., Aplysia pilsbryi (Eetson, 1898), and Aplysia pulmonica (Gould, 1852) are herein are reinstated as valid and Teth- ys pulmonica var. tryoniana is returned to the synonymy of Aplysia pulmonica Gould, 1852 rather than A. argus Riippell & Eeuckart, 1830. Nominal taxa Aplysia aequorea Heilprin, 1888 Figure lA, B Aplysia aequorea Heilprin, 1888: 325-327, pi. 16, figs 2-2b. Type locality. South side of Castle Harbor, opposite Tucker’s Town, Bermuda. Type material. Holotype (monotypy), ANSP A7030, 1 specimen (dehydrated but subsequently put back on al¬ cohol, 33.2 mm long preserved); ANSP 66519, 1 shell (35 mm) (A. Heilprin coll. 1888), removed from body of holotype by Heilprin. Remarks. Heilprin (1888) stated that he examined a sin¬ gle specimen. We do not illustrate the rehydrated alcohol specimen as it is a blackened mass without visible distin¬ guishing features. Current systematic position. Aplysia dactylomela Rang, 1828 (fide Valdes et al. 2013). Aplysia (Metaplysia) badistes Pilsbry, 1951 Figure IC-G Aplysia (Metaplysia) badistes Pilsbry, 1951: 1-6, figs 1-9. Type locality. Venetian Causeway, Biscayne Bay, Flori¬ da, USA. Type material. Holotype and paratype: ANSP A7028, 2 specimens (one whole except for removal of buccal mass, measuring 45.8 mm long preserved and the other dissect¬ ed, measuring 46.7 mm long preserved); ANSP 187712,1 shell (18.7 mm) and gizzard plates of dissected specimen of A7028 (H. A. Pilsbry coll, iv/1951), 1 slide with radu- las of dissected specimens. zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 361-372 363 Figure 1. Aplysiida types. A. Holotype of Aplysia aequorea (=A. dactylomeJa), shell, L 35 mm, ANSP 66519 in dorsal view. B. Same, in ventral view. C-G. Types of A. badistes, C. Shell, L 18.7 mm, ANSP 187712 in dorsal view. D. Same, in ventral view. E. Same species, whole specimen, L 45.8mm, ANSP A7028 in dorsal view. F. Same, in ventral view. G. Same, in right lateral view. H-J. Syntypes of A. benedicti {=Aplysia argus), whole specimens ANSP A7027. H. Specimen L 79 mm in dorsal view. I. Same, in right lateral view. J. Specimen L 76 mm in right lateral view. zse.pensoft.net 364 Cunha, C.M. & Rosenberg, G.: Type specimens of Aplysiida in the Academy of Natural... Remarks. Aplysia badistes Pilsbry, 1951 is the type species of subgenus Metaplysia Pilsbry, 1951. Pilsbry stated “Type and another specimen are 187712ANSP”. Pilsbry described the internal anatomy from one specimen but extracted the radula from both. Someone separated the shell and gizzard plates as ANSP 188279, and labeled them “Paratype”, a term not used in the publication. We have restored them to the original catalogue number (187712) as it is not possible to tell which specimen Pilsbry intended to be the holotype. Current systematic i^os\i\on. A. Juliana Quoy & Gaimard, 1832 Bales 1960). Aplysia (Tethys) benedicti Eliot, 1900 Figure IH-J Aplysia {Tethys) benedicti Eliot, 1900: 513-515, pi. 19, fig. 2a, b. Type locality. Apia Harbor, Upolu, Samoa. Type material. Syntypes: ANSP A7027, 2 specimens + 1 slide with radulae and jaw mounted (leg., C. N. E. Eliot, 19-21/vii/1899) [1 specimen (Eliot 1900: fig 2a), pre¬ served 76 mm long, whole, not dissected; another speci¬ men, preserved 79 mm long, with radula removed]. Current systematic position. Aplysia argus Riippell & Eeuckart, 1830 (^95. Aplysiaper- viridis Pilsbry, 1895 pilsbryU Tethys Letson, 1898. Aplysia pilsbryi (Let- son, 1898) robertsi^ Tethys Pilsbry, 1895. Aplysia robertsi (Pilsb¬ ry, 1895) swiftiU Dolabrifera Pilsbry, 1896. Dolabrifera ascifera (Rang, 1828) tryoniana^ Tethys pulmonica var. Pilsbry, 1895. Aplysia pulmonica Pease, 1860 willcoxi, Aplysia Heilprin, 1887. Aplysia fasciata Poiret, 1789 Acknowledgements We thank Amanda Lawless, Francisco Borrero, and Paul Callomon (ANSP) for help with photography and in de¬ termining the type status of the specimens and Dr An¬ gel Valdes (California State Polytechnic University) for sending the pictures of Aplysia dactylomela, CPIC 00297 and A. pulmonica, CPIC 0031. This work was support¬ ed by Capes Foundation, Bolsista da CAPES proc. no. 8739/13-7 (C.M. Cunha) and by National Science Foun¬ dation grant DBI 1203605, to G. Rosenberg, for digital imaging of type specimens at ANSP. 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Aquatic Invasions 8(4): 427-436. https://doi.org/10.3391/ ai.2013.8.4.06 Valdes A, Breslau E, Padula V, Schrbdl M, Camacho Y, Malaquias MAE, Alexander J, Bottomley M, Vital XG, Hooker Y, Gosliner TM (2017) Molecular and morphological systematics of Dolabrif¬ era Gray, 1847 (Mollusca: Gastropoda: Heterobranchia: Aplysiom- orpha). Zoological Journal of the Einnean Society 184(1): 31-65. https://d 0 i. 0 rg/l 0.1093/zoolinnean/zlx099 zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 373-390 | DOI 10.3897/zse.95.34486 4>yEnsPFr. BERLIN A glimpse in the dark? A first phylogenetic approach in a widespread freshwater snail from tropical Asia and northern Australia (Cerithioidea, Thiaridae) Dusit Boonmekam\ Duangduen Krailas\ France Gimnich^, Marco T. Neiber^, Matthias Glaubrecht^ 1 Department of Biology, Faculty of Science, Silpakorn University, Nakhon Fathom, 73000, Thailand 2 Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, 53113 Bonn, Germany 3 Center of Natural History (CeNak), Universitdt Hamburg, Martin Luther King-Platz 3, 20146 Hamburg, Germany http://zoobank.org/BC529DE5-DF53-41EF-A803-383E08E04721 Corresponding author: Marco T. (marco-thomas.neiber@uni-hamburg.de) Academic editor: Thomas von Rintelen ♦ Received 12 March 2019 ♦ Accepted 30 May 2019 ♦ Published 3 July 2019 Abstract Thiaridae are a speciose group of freshwater snails in tropical areas including a high number of described nominal taxa for which modern revisions are mostly lacking. Using an integrative approach, the systematic status of a group of thiarids from the Oriental region, including the nominal species Melania aspera and M rudis, is reassessed on the basis of shell morphology and biometry, radula dentition patterns, and reproductive biology along with molecular genetic methods. Our results suggest that populations from the Oriental region cannot be distinguished on the basis of shell morphology, radula characters and their reproductive mode and are monophyletic based on mitochondrial sequences. Hence, M rudis with M aspera are regarded as belonging to the same species along with several other nominal taxa that were previously included in M rudis. Moreover, populations from Thailand and Australia, from where the species was not previously recorded, could be shown to form a monophyletic group together with samples from Indonesia. However, a generic affiliation with Thiara, in which the investigated taxa were often included in the past, was not supported in our phylogenetic analyses, highlighting the need for a comprehensive revision of the genus-group systematics of Thiaridae as a whole. Key Words Cerithioidea, evolutionary systematics. Oriental region, Thailand Introduction Despite advances in the understanding of the family-level phylogeny of Cerithioidea Fleming, 1822, the taxonom- ical diversity in Thiaridae Gill, 1871 (1823) is still not well understood, and evolutionary systematic research in the sense of Glaubrecht (2010) in this particular family is still in its infancy. The Thiaridae, in earlier treatments subsumed under the name Melaniidae Children, 1823, have been used as a “rubbish bin” to accommodate all freshwater lineages belonging to the Cerithioidea. Only after the removal of the families Pachychilidae Fischer & Crosse, 1892, Melanopsidae Adams & Adams, 1854, Pa- ludomidae Stoliczka, 1868, Pleuroceridae Fischer, 1885 (1863), and Semisulcospiridae Morrison, 1952) (Camp¬ bell 2019; Neiber and Glaubrecht 2019b, 2019c, 2019d; Strong and Lydeard 2019 and references therein) and recently the Neotropical Hemisinidae Fischer & Crosse, 1891 (Glaubrecht and Neiber 2019a), a more accurate cir¬ cumscription of “core” Thiaridae began to emerge on the basis of molecular and/or morphological evidence (e.g., Glaubrecht 1993, 1996, 2011; Holznagel and Lydeard 2000; Lydeard et al. 2002; Glaubrecht et al. 2009; Strong 2011; Strong etal. 2011). In addition to uncertainties in the delimitation of gen¬ era, research on thiarids is further complicated by the large disparity of shell characters among species, a large phenotypic plasticity within species and a high ecologi- Copyright Dusit Boonmekam et al. This is an open access articie distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the originai author and source are credited. 374 Dusit Boonmekam et al.: Evolutionary systematics of Thiara aspera cal adaptability that is, however, also known from other limnic Cerithioidea. This conchological variability has certainly led to an overestimation of the number of spe¬ cies in the past, as specifically shown for limnic lineages in the superfamily (Glaubrecht 1993, 1996; Kohler and Glaubrecht 2001, 2003, 2006; Glaubrecht and Kohler 2004; Glaubrecht et al. 2009), but may also cause prob¬ lems in delimiting species resulting in an underestimation of the actual morphological disparity versus the taxonom- ical diversity, at least in some cases. These problems are exacerbated by the putatively widespread occurrence of parthenogenesis in different lineages of Thiaridae (Glau¬ brecht 1996) and the associated problems of what is actu¬ ally meant by “species” in this case (e.g. Hausdorf 2011). Additionally, Thiaridae have also realised different life history strategies that were characterised by Glaubrecht (1996, 1999, 2006, 2011) by the duration of ontogenet¬ ic stages to remain within a specialised structure of the female, viz. the subhaemocoelic brood pouch. While in some thiarids only very early ontogenetic stages, i.e. embryos without shell, develop and are released as veligers (ovoviviparity), other thiarid species brood and even transform their subhaemocoelic brood pouch into a matrotrophic organ or “pseudoplacenta” that apparently nourishes the developing juveniles, as e.g. in the South¬ east Asian thiarid Tarebia granifera (Lamarck, 1816) (eu- viviparity, see Glaubrecht 1996; Glaubrecht et al. 2009; MaaB and Glaubrecht 2012; Veeravechsukij et al. 2018b). Finally, some thiarids also have an extraordinarily high invasive potential, such as Melanoides tuberculata (Muller, 1774) and Tarebia granifera and today have an almost pantropical distribution (e.g.. Brown 1994; Glau¬ brecht 1996). To date, only few of the several dozen thiarid taxa have seen closer investigation. Glaubrecht et al. (2009) and MaaB and Glaubrecht (2012) surveyed the thiarid fauna of Australia. Dechruska et al. (2013) evaluated the status and identity of the nominal taxon Melania jugicostis Han¬ ley & Theobald, 1876 from the Southeast Asian main¬ land, and Veeravechsukij et al. (2018a, 2018b) investi¬ gated the phylogeography and reproductive biology of T. granifera and its trematode parasites. However, many other named taxa have been rarely studied and, thus, re¬ main enigmatic and even pure nomenclatorial “ghosts” with highly questionable status as evolutionary relevant entities, which hampers further insights into the systemat¬ ics, biogeography, and evolution of these freshwater gas¬ tropods otherwise under scrutiny, e.g., in speciation and/ or radiation studies. Melania aspera Lesson, 1831, which was originally described from New Guinea (Lesson 1830-1831), is such an “enigmatic” taxon (Fig. 1), which Glaubrecht and Pod- lacha (2010) regarded as a possible senior synonym of the nominal species Melania rudis Lea & Lea, 1851. The latter taxon is usually regarded as belonging to Thiara Roding, 1798 and thought to be relatively widespread, be¬ ing reported from several countries, occurring from India and Sri Lanka to Southeast Asia and the Indo-Australian archipelago (Schepman 1892, 1915; Rensch 1934; van Benthem Jutting 1937; Subba Rao 1989; Ramakrishna and Dey 2007; Budha 2010; Path and Talmale 2011, see also Fig. 2). However, actual distribution records are rel¬ atively scarce in the literature and the distinction from other nominal thiarid taxa remains uncertain so far. As a further contribution towards a better understand¬ ing of thiarid diversity, we here re-evaluate the identity of M. aspera and M. rudis on the basis of museum sam¬ ples including available type material as well as material collected during ongoing surveys in Southeast Asia using shell morphology and biometry, radula dentition patterns, and reproductive biology along with molecular genetic methods. Nomenclatural issues and the synonymy of the genus Thiara are also discussed. Material and methods This study is mainly based on the examination of spec¬ imens in the collections of the Parasitology and Medi¬ cal Malacology Research Unit, Department of Biology, Faculty of Science, Silpakorn University, Thailand and the Museum fur Naturkunde, Berlin, Germany, and sup¬ plemented by material from other museums (see below). Additionally, new samples were collected using hand picking and scooping methods in Thailand and Australia. Specimens were fixed in 75-96% ethanol. Collection acronyms MNHN Museum National d’Histoire Naturelle de Paris, France SUT Silpakorn University, Nakhon Pathom, Thailand USNM National Museum of Natural History, Wash¬ ington, USA ZMB Museum fur Naturkunde, Berlin, Germany (formerly Zoologisches Museum Berlin) Coordinates (WGS84) of localities were taken with a GPS device or determined as accurately as possible from a map. Sampling sites were then mapped on a dot-by- dot basis to a digitally reduced version of the drainage pattern map of the Indo-Australian region. This map was prepared using a relief map on the basis of the Glob- al30-Arc-Second Elevation Data (GTOPO30) from the U.S. Geological Survey and a river map from the map server Aquarius Geomar; and then compiled using Adobe Photoshop CS3 and Adobe Illustrator. For the exact local¬ ity data, see the material examined section. Shell characters Specimens were photographed using a digital EOS 350D camera (Canon, Tokyo, Japan). Standard biometric pa¬ rameters were taken from each shell using electronic cal- zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 373-390 375 Figure 1. Shells of ''Thiara” aspera (Lesson, 1831). A. Holotype of Melania aspera Lesson, 1831, MNHN 21098, ‘La Nou- velle-Guinee’ [more specifically Manokwari on New Guinea Island, West Papua, Indonesia, see Glaubrecht and Podlacha 2010]; B. Syntype of Melania rudis Lea & Lea, 1851, USNM 119778, Amboyna; C. Syntype of Melania microstoma Lea & Lea, 1851, USNM 119722, mountain streams, isle of Negros, Philippines; D. ZMB 107002, Calcutta, India; E. ZMB 107003, Ceylon, Sri Lanka; F. ZMB 127534, Don Ko Canal, Nakhon Pathom, Thailand; G. ZMB 127535, Don Ko Canal, Nakhon Pathom, Thailand; H. ZMB 127535, Don Ko Canal, Nakhon Pathom, Thailand; 1. ZMB 191279, Yehembang River, Bali, Indonesia; J. ZMB 191279, Ye- hembang River, Bali, Indonesia; K. ZMB 106472, Yehembang, Bali, Indonesia; L. East of Mendaya, stream southwest of Gumicik, Bali, Indonesia; M. ZMB 191278, stream at Tembeeha, road Tirobus-Kendari, Southwest Sulawesi, Indonesia; N. ZMB 107378, Banggai Islands, Peleng Island, West of Peninsula, Tataban river. Central Sulawesi, Indonesia; O. ZMB 107377, Banggai Islands, Peleng Island, West of Peninsula, Tataban river. Central Sulawesi, Indonesia.river; P, Q. ZMB 107617, Wabalarr, Roper River, Northern Territory, Australia; R. ZMB 106599, Berry Springs, Northern Territory, Australia. Scale bar; 1 cm. lipers (accuracy 0.1 mm): shell height (H), shell width (W), aperture length (AL; measured from the upper ap- ertural angle to the farthest point on the basal margin of the aperture), aperture width (AW; measured perpendic¬ ular to AL as the widest distance between outer apertural margin and outer margin of parietal callous), height of the body whorl (BW), and number of whorls (NW) as shown in Figure 3A. To reduce dimensionality a principal component analysis was conducted on log-transformed shell measurements using R 3.3.2 (R Core Team 2016). Only the minimal number of PC A axes that aceounted for more than 95% of the cumulative variation were used for further testing. The Shapiro-Wilk test was conducted in R to test for normal distributions of PCA 1 and PCA 2 values, respee- tively, for the here proposed geographic subgroups, i.e., samples from 1) Thailand, 2) Indonesia, 3) India, and Sri Lanka, and 4) Australia. Since some of the Shapiro-Wilk tests were significant {p < 0.05), the non-parametric Kru- skal-Wallis rank sum test was conducted for PCA 1 and zse.pensoft.net 376 Dusit Boonmekam et al.: Evolutionary systematics of Thiara aspera Figure 2. Distribution and reproductive strategy of Thiara’' aspera (Lesson, 1831). Stars: type localities of a) Melania aspera Les¬ son, 1831, Monokwari, New Guinea, b) Melania rudis Lea & Lea, 1851, Amboyna and c) Melania microstoma Lea & Lea, 1851, mountain streams, isle of Negros, Philippines. Pie charts show the percentages of offspring in the brood pouch of female T. aspera in different size classes as defined in Glaubrecht et al. (2009), see inset. The numbers near the pie charts refer to the number of individuals examined per population. Filled circles: material preserved in ethanol; open circles: dry shells. PCA 2 assuming the grouping of specimens according to geography followed by Dunn’s test (Bonferroni-cor- rected) as post-hoc test as implemented in the R package “dunn.test 1.3.5” (Dinno 2017) in case that the Krus- kal-Wallis-rank-sum tests were significant. Radula preparation Shells of representative specimens were cracked with a small vice and removed from the soft body parts, which were afterwards examined and dissected with the aid of a Leica Wild MZ 9.5 stereo microscope (Leica Microsys¬ tems, Wetzlar, Germany). Radulae were extracted follow¬ ing the protocol of Holznagel (1998), fixed on aluminium stubs, and coated with platinum using a Polaron SC 7640 Sputter Coater (Quorum Technologies, East Grinstead, UK). Radulae were then viewed and photographed (ori¬ ented so that denticles on the teeth were well visible) with a scanning electron microscope (SEM) EVO ESIO (Zeiss, Oberkochen, Germany). Content of brood pouch The brood pouch was opened after removing the mantle and its content was counted under a Eeica Wild MZ 9.5 stereo microscope. Both, shelled juveniles and embryos. were grouped into standard size classes as described in Glaubrecht et al. (2009). Embryos and juveniles from representative specimens were fixed on aluminium stubs, air-dried, coated with platinum using a Polaron SC 7640 Sputter Coater, and then viewed, photographed, and mea¬ sured (Fig. 3B) with a EVO ESIO SEM. Parameters of the embryonic shell were measured from SEM images as shown in Figure 3B: diameter of first half whorl (de; measured as the maximal witdth of the shell after 0.75 turns of the suture line), width of first quarter whorl (he; measured parallel to de as the distance from the starting point of the suture to the point after 0.25 turns of the su¬ ture line), width of first half whorl (we; measured perpen¬ dicular to de as the distance from the starting point of the suture to the point after 0.5 turns of the suture line). Molecular methods and phylogenetic analyses Total genomic DNA was extracted from ethanol-pre¬ served foot muscle tissue using a CTAB protocol as de¬ scribed by Winnepenninckx et al. (1993) from 31 thiarid specimens and Paludomus siamensis Blanford, 1903 as outgroup representing one of the cerithioidean families, which have been shown to be closely related to the Thia- ridae (Wilson et al. 2004; Strong et al. 2011). For phylogenetic analyses, fragments of the mitochon¬ drial cytochrome c oxidase subunit 1 (coxl) gene and the zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 373-390 377 A w de B Figure 3. Measured shell parameters. A: H - shell height; W - shell width; BW - body whorl height; AL - aperture length; AW - aperture width. B: he - height of embryonic shell; we - width of embryonic shell; de - maximum diameter at one whorl. 16 S rRNA (16S) gene were amplified by polymerase chain reaction (PCR) using the primer pairs LCO1490 (5’-GGT CAA CAAATC ATAAAGATATTG G-3’; Fol- mer et al. 1994) plus HC02198var (5’-TAW ACT TCT GGG TGK CCA AAR AAT-3’; Rintelen et al. 2004) and 16S_F_Thia2 (5’-CTT YCG CAC TGA TGA TAG CTA G-3’; Neiber and Glaubrecht 2019a, see also Gimnich 2015) plus H3059var (5’-CCG GTY TGA ACT CAG ATC ATG T-3’; Wilson et al. 2004), respectively. Am¬ plifications were conducted in 25 pi volumes containing 50-100 ng DNA, 1 x PCR buffer, 200 mM of each dNTP, 0.5 mM of each primer and 1 U of Taq polymerase. After an initial denaturation step of 3 min at 94 °C, 35 cycles of 30 s at 94 °C, 60 s at 45-62°C and 60-120 s at 72°C were performed, followed by a final extension step of 5 min at 72°C. PCR products were purified using a NucleoSpin Extract II Kit (Macherey-Nagel, Bethlehem, PA, USA). Both strands of the amplified gene fragments were cy¬ cle-sequenced using the primers employed in PCR with the Big Dye Terminator chemistry version 1.1 (Applied Biosystems, Inc., Waltham, MA, USA). Sequences were visualised on an ABI 3130x1 or ABI 3730x1 Genetic Ana¬ lyzer (Applied Biosystems, Inc.). Forward and reverse sequence reads were assembled with CODONCODE AEIGNER v. 3.7.1 (CodonCode Corporation, Dedham, MA, USA) and corrected by eye. For information on vouchers, see Table 1. The protein coding coxl sequences were aligned with MUSCEE (Edgar 2004) as implemented in MEGA 6 (Tamura et al. 2013) under default settings. The 16S sequences of were aligned with MAFFT (Katoh and Standley 2013) using the Q-INS-i iterative refinement algorithm and otherwise de¬ fault settings, because this algorithm has been described to perform better for the alignment of sequence data sets that may contain deletions and insertions than alternative mul¬ tiple sequence alignment methods (Golubchik et al. 2007). Maximum likelihood (ME), Bayesian Inference (BI), and maximum parsimony (MP) approaches were used to reconstruct the phylogenetic relationships. The sequence data set was initially divided into four partitions for the nucleotide model-based ME and BI approaches: 1) co¬ don positions of coxl, 2) 2"^^ codon positions of coxl, 3) 3’'^ codon positions of the coxl, and 4) the 16S. To se¬ lect an appropriate partitioning scheme and evolutionary models the sequence data set was analysed with PARTI- TIONFINDER V. 1.1.1 (Eanfear et al. 2012) conducting an exhaustive search and allowing for separate estimation of branch lengths for each partition using the Bayesian in¬ formation criterion as recommended by Euo et al. (2010). Models to choose from were restricted to those available in MRBAYES v. 3.2.6 (Ronquist et al. 2012) as well as in GAREI v. 2.1 (Zwickl 2006). As best-fit partitioning scheme, the PARTITIONFINDER analysis suggested to combine the and 2"‘^ codon positions of coxl and the 16S sequences together in one partition (GTR + G model) and the 3'^'* codon positions of coxl in a second partition (HKY+G model). The BI analysis was performed using MRBAYES v. 3.2.6. Metropolis-coupled Monte Carlo Markov chain (MC^) searches in MRBAYES were run with four chains in two separate runs for 50,000,000 generations with de¬ fault priors, trees and parameters sampled every 1000 gen¬ erations under default heating using the best-fit model as suggested by PARTITIONFINDER. Diagnostic tools in MRBAYES, including Estimated Sample Size (ESS) val¬ ues > 200, were used to ensure that the MC^ searches had reached stationarity and convergence. The first 5,000,000 generations of each run were discarded as burn-in. Heuristic ME analysis was performed with GAREI using the best-fit models as suggested by PARTITION¬ FINDER. Support values were computed by bootstrap¬ ping (BS) with 1,000 replicates. Heuristic MP searches were carried out with PAUP v. 4.0bl0 (Swotford 2002) using 100 random-addition-se¬ quence replicates and TBR branch swapping. Support values were computed by bootstrapping with 1,000 replications. Alternative phylogenetic hypotheses were tested using the approximately unbiased (AU) test (Shimodeira 2002) as implemented in the program CONSEE (Shimodeira and Hasegawa 2001). Information on vouchers and GEN- BANK accession numbers are listed in Table 1. zse.pensoft.net 378 Dusit Boonmekam et al.: Evolutionary systematics of Thiara aspera Table 1. Museum registration numbers, GenBank accession numbers and locality data for the specimens used in the molecular phylogenetic analyses. Abbreviations for countries: AUS - Australia, IDN - Indonesia, IND - India, THA- Thailand. Taxon Museum number Extraction number Country Latitude Longitude GenBank accession number coxl 16 S rRNA gene “Thiara" aspera SUT 0311020 11449 THA 13°38'08"N 100‘’05'03"E MK879291 MK879427 SUT 0312070 11446 THA 13°48'08"N 100°02'06"E MK879292 MK879428 SUT 0311044 9603 THA 13°38'08"N 100°05'03"E MK879290 - ZMB 191268 2200 IDN 03°39'28"S 122°13'52"E MK879296 MK879434 ZMB 191488 4558 IDN 08‘’38'39"S 115°16'38"E MK879297 MK879435 ZMB 107377 6494 IDN 0r32T8"S 122°5r28"E MK879293 MK879429 ZMB 107378 6495 IDN 0r32T8"S 122°5r28''E MK879294 MK879430 ZMB 107617 7586 AUS 14°56'02"S 133°10'26"E MK879295 MK879433 ZMB 107617 8743 AUS 14‘’56'02"S 133°10'26"E - MK879431 ZMB 107617 8744 AUS 14°56'02"S ISSnO'SET - MK879432 “Stenomelania" denisoniensis ZMB 106682 7599 AUS 14°55'47"S 133°08'44"E MK879288 MK879425 ZMB 106632 7602 AUS 15°00'42"S 133°14'25"E MK879287 MK879424 Thiara amaruia ZMB 191489 2886 IDN 0r26'43"S IST-SS'Or'E MK879289^ MK879426^ ZMB 107472 6496 IDN 03‘’35'28"S 128°08'42"E MK094074 MK098355 Thiara winteri ZMB 106554 1043 IDN 08‘’23'38"S 114°45'04"E MK879301 MK879439 ZMB 190261 1055 IDN 02°35'34"S ISO^SATOT MK879302 MK879440 Thiara cf. winteri ZMB 106472 1001 IDN 08‘’23'38"S 114°45'04"E MK879298 MK879436 ZMB 191279 2232 IDN 08°23'36"S 114°45'04"E MK879299 MK879437 ZMB 191279 4559 IDN 08°23'36"S 114‘’45'04"E MK879300 MK879438 Mienipiotia scabra ZMB 107382 6514 IDN 00°48'33"N 127°17'40"E MK879279 MK879416 ZMB 107564 7340 AUS 14‘’55'38"S 133°07'06"E MK879280 MK879417 ZMB 127495 9574 THA 07‘’55T5''N 099‘’15'47"E MK879285 MK879422 SUT 0312060 9578 THA 12°51T5"N 099°59'49"E MK879278 MK879415 SUT 0311024 9580 THA 14°54'04"N 100°03'48"E MK879276 MK879413 SUT 0311040 9582 THA 13°25'07"N OSS-STTST MK879277 MK879414 ZMB 127470 9589 THA 08°27'09"N 098°28'01"E MK879284 MK879421 ZMB 127468 9599 THA 12°56'54"N 099°28'52"E MK879283 MK879420 ZMB 107962 9779 THA 16‘’37'38"N 100‘’56'43"E MK879282 MK879419 ZMB 107869 9781 THA 08°38T8"N 099°44'59"E MK879281 MK879418 SUT 0311009 9787 THA 16°ir33"N 099°15'51"E MK879275 MK879412 Meianoides tubercuiata ZMB 200313 7530 IND ir34'45"N OTO-SA'SET MK879274 MK879411 Paiudomus siamensis ZMB 107721 7334 THA 14°26T5"N 098°5rH"E MK879286 MK879423 “ From Neiber and Glaubrecht (2019a). Results Biometric analyses The first two principal components (PCA 1 and PCA 2) account for > 95% of the cumulative variation in shell parameters. The plot of PCA 1 vs PCA 2 (Fig. 4A) shows that the clusters of specimens that were grouped accord¬ ing to geographic origin widely overlap. Especially the clusters of specimens from Thailand and Indonesia (cor¬ responding to mitochondrial Clades A and B, Fig. 4) and the clusters of specimens from Australia (corresponding to mitochondrial Clade C, Fig. 4) also widely overlap. The Kruskal-Wallis rank sum tests were significant for PCA 1 (p < 5.0 X 10-6) PCA2 (p < 2.0 x lO-’^), i.e., at least one group stochastically dominates one other group in each of the tests. Dunn’s test for PCA 1 found signif¬ icant differences between the groups including samples from Indonesia and Australia (p < 0.0001) as well as be¬ tween the groups including samples from Australia and Thailand (p < 0.0075), respectively, but not for pairwise comparisons of the other groups (Fig. 4B). Dunn’s test for PCA 2 found significant differences between the fol¬ lowing groups: Indonesia vs Australia (p < 0.0031), Indo¬ nesia vs Thailand (p < 0.0001), Australia vs Thailand (p < 0.0007), Australia vs India/Sri Lanka (p < 0.0004), and Thailand vs India/Sri Lanka (p < 0.0001), but not for In¬ donesia vs India/Sri Lanka (Fig. 4C). However, both for PCA 1 and PCA 2 the comparison of ranges shows that the ranges of all pairs of geographic groups overlap and therefore do not allow a diagnostic separation of these groups on the basis of the biometric data. The included type specimens of the nominal taxa M. rudis and M. mi¬ crostoma fall within the convex hull spanned by speci¬ mens sampled from Thailand, Indonesia, Australia, India, and Sri Lanka in the PCA 1 vs PCA 2 plot; only the ho- lotype of the nominal taxon M. aspera lies outside this area (Fig. 4A), although closely resembling the examined syntypes of M. rudis and M. microstoma with respect to shell sculpture and overall shape. Phylogenetic analyses A clade including Thiara amarula (Linnaeus, 1758) (the type species of Thiara Roding, 1798), T winteri (Busch, 1842) in Philippi (1842-1844), T. cf. winteri from Bali, and the specimens identified as Thiara aspera from Thai- zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 373-390 379 land, Indonesia, and Australia as well as ''Stenomelanid’' denisoniensis (Brot, 1877) in Brot (1874-1879) was re¬ covered in all three analyses (BI: 1.00, BS (ML): 92, BS (MP): 96). However, Thiara is paraphyletic with respect to “5.” denisoniensis. Thiara amarula grouped together with T. winteri and T. cf. winteri from Bali in a clade (BI: 1.00, BS (ML): 97, BS (MP): 93). A sister group relation¬ ship of T. amarula and T. winteri was recovered in the BI and ML analyses (BI: 0.99, BS (ML): 83) but not in the MP analysis (BS (MP): < 50). Within this clade, the clades including specimens of T. amarula, T winteri, and T. cf winteri from Bali, respectively, were supported (BI: 1.00, BS (ML): 96-100, BS (MP): 100). The clade containing T. amarula, T winteri, and T. cf winteri was recovered as the sister group of a clade containing “5.” denisoniensis and specimens from Thailand, Australia, and Indonesia (including also a single specimen from Bali) assigned to the T. aspera on basis of conchological similarity with rather high support (BI: 1.00, BS (ML): 92, BS (MP): 96). Within this clade, “5.” denisoniensis was recovered as the sister group (BI: 1.00, BS (ML): 94, BS (MP): 90) ofT. as¬ pera, which in turn formed a rather well-supported clade (BI: 1.00, BS (ML): 90, BS (MP): 79). Thiara aspera specimens from Australia (Clade C) formed a maximally supported clade. The T aspera specimens from Thailand grouped together with a single individual from Bali in a well-supported clade (Clade A; BI: 1.00, BS (ML): 92, BS (MP): 100), which was sister to another supported clade (Clade B; BI: 1.00, BS (ML): 77, BS (MP): 100) that in¬ cluded T aspera specimens from Sulawesi. The included specimens of Mieniplotia scabra (Muller, 1774) formed the sister clade of a specimen of Melanoi- des tuberculata from India in the BI analysis, albeit without support. To test alternative phylogenetic hypotheses, we conducted four AU tests: 1) the monophyly of M. scabra (p = 0.252) and 2) the monophyly of T winteri plus the T. cf winteri specimens from Bah (p = 0.156) could not be reject¬ ed, whereas 3) the monophyly of T aspera, T winteri and T cf winteri (p < 0.001) and 4) the monophyly of Thiara excl. “5.” denisoniensis (p = 0.033) but including the T aspera specimens was rejected at a confidence level of a = 0.05. Systematic account Thiaridae Gill, 1871 (1823) Thiara Roding, 1798* Vesica Humphrey, 1797: 58 [unavailable, published in a work rejected for nomenclatural purposes, see International Commission on Zoo¬ logical Nomenclature 1912; 116-117; among the mentioned species is Vesica thiara Humphrey, 1797 (unavailable) = Helix amarula Lin¬ naeus, 1758], TTz/araRoding, 1798: 109 [type species: HelixamarulaVmmQm, 1758, by subsequent designation of Herrmannsen 1849 in Herrmannsen 1847-1849: 576]. a question mark indicates a tentative synonymisation Melania Lamarck, 1799: 75 [type species; Helix amarula Linnaeus, 1758, by monotypy]. Melanigenus Renier, 1807; pi. 8 [unavailable, published in a work re¬ jected for nomenclatural purposes, see International Commission on Zoological Nomenclature 1956: 290], Melas Montfort, 1810: 322-324 [unjustified emendation of Melania Lamarck, 1799]. Melanidia Rafinesque, 1815: 144 [unjustified emendation of Melania Lamarck, 1799]. Melanea- Sowerby 1818 in Sowerby 1818-1822: 33 [incorrect subse¬ quent spelling of Melania Lamarck, 1799]. ? Spirilla Gray, 1824: 254 [unavailable, published in synonymy; men¬ tioned as Spirilla spinosa (quoting a label or note attributed to G. Humphrey as "^Spirilla spinosa, freshwater spiral spined shell, from Admirality Island, New Guinea”) under Melania setosa Swainson, 1824 (= Thiara cancellata Roding, 1798, see Swainson 1824: 13-15 and Wilkins 1957; 167-169) and as being conspecific with the nomen- claturally unavailable Buccinum aculeatum Lister, 1692: pi. 1055, fig. 8. Mentioned as a synonym by Ferussac 1824: 318, Gray 1825: 524, Oken 1833: 133, Gray 1847: 152, Wilkins 1957: 167 as well as by Agassiz 1842: 84, Agassiz 1847: 348 and Herrmannsen 1848 in Herr¬ mannsen 1847-1849: 491 in nomenclators, with the name attributed to Humphrey 1797 (where it could not be found). Used by Favre 1869: 79 (attributing the name to G. Humphrey but without reference to the work of Gray 1824) for a subgenus of Fusus Bruguiere 1789 in Bm- guiere 1789-1792 and in a very different meaning from that of Gray 1824 and therefore not regarded here as having been made available from that work]. Spirella - Oken 1833: 61 [incorrect subsequent spelling of the unavailable Spirilla Gxdiy, 1824]. Melacantha Swainson, 1840: 341 [type species: Helix amarula Linnaeus, 1758 by subsequent designation of Herrmannsen 1849 in Herrmannsen 1847-1849:26], Thaira - Gray 1840: 148 [incorrect subsequent spelling of Thiara Roding, 1798]. Amarula Sowerby, 1842: 61 [type species: Helix amarula Linnaeus, 1758, by monotypy], Melanium - Busch 1842 in Philippi 1842-1845: 4 [incorrect subsequent spelling of Melania Lamarck, 1799]. Tiara - Gray 1847: 152 [incorrect subsequent spelling of Thiara Roding, 1798]. Thaera - Agassiz 1847:367 [unavailable, emendation for Thaira as used by Gray 1840; 148 proposed in synonymy in a nomenclator], Lithoparches Gisiei, 1848: ix [nom. nov. pro Melania Lamarck, 1799; type species; Helix amarula Linnaeus, 1758, by typification of the replaced name]. Hydrognoma Gistel, 1848: 169 [nom. nov. pro Melania Lamarck, 1799, type species: Helix amarula Linnaeus, 1758, by typefication of the re¬ placed name], Tiaropsis Brot, 1871: 298 [non Agassiz 1849: 289-298; type species: Mel¬ ania winteri Busch, 1842 in Philippi 1842-1844: Melania, 1, pi. 1 figs 1,2 by subsequent designation of Brot 1874 in Brot 1874—1879: 7]. Cerithomelania Moore, 1899; 233-234 [type species; Helix amarula Lin¬ naeus, 1758 by original designation]. ? Ripalania Iredale, 1943: 209 [type species: Melania queenslandica Smith, 1882 by monotypy], ? Setaeara Morrison, 1952: 8 [type species; Thiara cancellata Roding, 1798 by original designation]. zse.pensoft.net 380 Dusit Boonmekam et al.: Evolutionary systematics of Thiara aspera Australia Indonesia India/Sri Lanka Thailand T Melania aspera, holotype ♦ Melania microstoma, syntype ■ Melania rudis, syntype -5.0 A -2.5 0.0 25 5.0 standardized PC 1 (93.8% explained variance) 1 -r ^ ^<2- ^e. • rN B Figure 4. Results of the analysis of biometric data of Thiara" aspera (Lesson, 1831) specimens from Australia (yellow), Indonesia (green), Thailand (red) and India/Sri Lanka (blue) and type material of Melania aspera Lesson, 1831 (holotype, triangle), Melania rudis Lea & Lea, 1851 (syntype, square) and Melania microstoma Lea & Lea, 1851 (syntype, diamond). A. Scatter plot of the first two axes of the principal component analysis (PCA) of biometric data. Coloured lines indicate the outline of the convex hull for each geographic group; B, C. Boxplots of PCA 1 (B) and PCA 2 (C); bars above the box plots indicate significant differences of groups resulting from testing with Dunn’s test. zse.pensoft.net 0.1 Figure 5. Bayesian 50% majority-rule consensus tree based on partial sequences mitochondrial cytochrome c oxidase subunit 1 {coxl) and 16S rRNA (16S) genes. Support values at nodes refer to Bayesian posterior probabilities (left), Maximum Likelihood (middle) and Maximum Parsimony (right) bootstrap values. AUS; Australia, IDN: Indonesia, THA: Thailand. Numbers at tips refer to DNA vouchers in the collection of the ZMB, see also Table 1. zse.pensoft.net 382 Dusit Boonmekam et al.: Evolutionary systematics of Thiara aspera Remarks. Many names have been proposed for the group of Thiaridae that is currently regarded as repre¬ senting Thiara Roding, 1798. Several of these names are objective junior synonyms of Thiara having the same type species {Helix amarula Linnaeus, 1758), and several others are nomenclaturally unavailable. A few, like Ripalania Iredale, 1943 or Setaeara Morri¬ son, 1952, may actually be synonyms of Thiara. How¬ ever, those hypotheses should be further tested using molecular genetic approaches. Therefore, these nom¬ inal genera were only tentatively included in the syn¬ onymy of Thiara. '‘‘‘Thiara^^ aspera (Lesson, 1831)* Figs 1, 6, 7 Melania aspera Lesson, 1831 in Lesson (1830-1831: 357-358) [type locality: “La Nouvelle-Guinee” (= New Guinea), restricted to Ma- nokwari by Glaubrecht and Podlacha (2010)]. Melania rudis Lea & Lea, 1851: 186 [type locality: ‘Amboyna’ (= Am¬ bon)]. Melania microstoma Lea & Lea, 1851: 186 [type locality: mountain streams, isle of Negros, Philippines]. ? Melania armillata Lea & Lea, 1851: 195-196 [type locality: India], ? Melania broti Reeve, 1859 in Reeve (1859-1861: pi. 22 fig. 160) [type locality: Ceylon (= Sri Lanka)]. ? Melania hybrida Reeve, 1859 in Reeve (1859-1861: pi. 13 fig. 163) [type locality: not given], ? Melania chocolatim Brot, 1860: 256-257, pi. 16, fig 2 [type locality: “Ceylon” (= Sri Lanka)]. ? Melania (Tiaropsis) rudis var. spinosa Brot, 1877 in Brot (1874-1879: 306) [type locality: not given, see also Brot (1868: 33, pi. 1, fig. 7)]. ? Melania {Tiaropsis) drilliiformis Martens, 1897: 305 [nomen nudum], ? Melania fortitudinis Fulton, 1904: 51-52, pi. 4, fig. 3 [type locality: “Soekaboemi, Java” (= Sukabumi, Java)]. ? Melania rudis var. cylindrica Schepman, 1915: 27 [type locality: West Ceram, Kairatu (= West Seram Island, Kairatu)]. Diagnosis. Thiarid with a turreted, subcylindrical to elon¬ gate-ovoid, strongly ornamented high-spired shell with usually rather flattened whorls and a narrowly pyriform aperture that at most reaches half the total shell height, but usually less. Ornamentation of the shell consisting of sinuous axial ribs that usually reach to the base of the body whorl and spiral chords that form nodes where they intersect the ribs; spiral chords usually present on the en¬ tire whorl but strongest at the base of body whorl. Remarks. The examined type specimens of M. aspera, M. rudis, and M. microstoma correspond well to each other in overall shell shape and sculpture and are here regarded as conspeciflc because of this. As already noted by Brot (1874-1879: 307) and Glaubrecht and Podlacha (2010: 200), the name Melania aspera Les¬ son, 1830 has priority over the somewhat more fre¬ quently used name Melania rudis (e.g., van Benthem Jutting 1937, 1956; Subba Rao 1989; Ramakrishna and Dey 2007; Budha 2010; Path and Talmale 2011 as T rudis). The holotype of Melania aspera is unusual in possessing a very small aperture in relation to overall shell height, possibly explaining its isolated position in the PCA 1 vs PCA 2 scatter plot (Fig. 4A). The nomi¬ nal taxa M. armillata, M. broti, and M. chocolatum de¬ scribed from India or Sri Lanka were regarded by Brot (1874-1879) as closely related to M. rudis and are here tentatively synonymised with M. aspera, largely fol¬ lowing the views of Rensch (1934) and van Benthem Jutting (1937, 1956) who synonymised these taxa with M. rudis. According to Brot (1874-1879: 307-308) Melania hybrida is based on a teratological specimen with an unusual aperture formation and is here tenta¬ tively synonymised with M. aspera. The nominal taxon Melania {Tiaropsis) rudis var. spinosa Brot, 1877 is an individual variation of M. aspera with somewhat longer shoulder spines. The original figures and descriptions of Melania fortitudinis and M. {Tiaropsis) rudis var. cylindrica from Java and West Seram Island also cor¬ respond well with the holotype of M. aspera and are herein treated as synonyms of the former. Type material examined. Holotype of Melania aspera Lesson, 1831, MNHN 21098, “La Nouvelle-Guinee”; syntype of Melania rudis Lea & Lea, 1851, USNM 119778, “Amboyna”; syntype of Melania microstoma Lea & Lea, 1851, USNM 119722, ‘mountain streams, isle of Negros, Philippines’. Additional material examined (w: ethanol preserved samples). India: Kolkata, ZMB 107002. Sri Lanka: Co¬ lombo, ZMB 107003. Thailand: Samut Sakhon Provin¬ ce, Klong Don Ko, SUT 0311020, ZMB 127535, SUT 0311044, SUT 0311053, ZMB 127534, w; Nakhon Pa- thom province. Pond in Silpakom University campus, SUT 0312069, SUT 0312070 = ZMB 127536, w. Indo¬ nesia: Bali: South Bali, Yehembang River, ZMB 191279, ZMB 191279a, w. South Bali, at Yehembang, ZMB 106472, w; east of Mendaya, stream southwest of Gu- micik, ZMB 191488; Sulawesi: South Sulawesi, Kalena catchment, Angkona river, ZMB 192751, w; southeast Sulawesi, Pohara river, at Pohara, road Kendair to Kola- ka, ZMB 191261, w; southeast Sulawesi, Simbune river, 1 km northeast of Raterate, road Kendari to Kolaka, ZMB 191262, ZMB 191262a, w; southeast Sulawesi, stream at Tembeeha, road Tirobus to Kendari, ZMB 191278, w; central Sulawesi, Banggai Islands, Peleng Island, West Peninsula, Tataban river, ZMB 107378, w, ZMB 107377,w. Australia: Northern Territory: Berry Springs, ZMB 106704, w, ZMB 106599a, w, ZMB 127616, w; Wabalaar, Roper River, ZMB 107617, w, ZMB 107614, w, ZMB 127645, w; Salt creek, ZMB 127619, w, ZMB 127636, w, ZMB 127637, w; Roper Bar, ZMB 127620, w; Queensland: O’Shanassy, ZMB 107280. a question mark indicates a tentative synonymisation zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 373-390 383 A B Figure 6. Juvenile and embryonic shells of "'Thiara" aspera (Lesson, 1831), SUT 0311020, Samut Sakhon Province, Klong Don Ko. A. Lateral view; B. Apical whorls, lateral, C. Apical view. D. Details of the protoconch. Scale bars; 450 pm (A); 250 pm (B); 200 pm (C); 100 pm (D). Shell. Turreted, subcylindrical to elongate-ovoid, cor¬ neous to dark brown, with up to nine whorls (the early whorls usually eroded) (Fig. 1; for juvenile shells, see Fig. 6). Whorls rather flat to convex, separated by a slightly impressed to distinctly impressed, undulating suture. Whorls slightly constricted below the suture, or¬ namented with sinuous ribs and spiral chords that usu¬ ally form nodules at their intersections. Radial sculpture usually strongest on the upper half of the whorls, with the nodules at the shoulder of the whorls usually largest, sometimes forming spines. Towards the lower part of the body whorl the spiral sculpture often becomes the domi¬ nant sculptural element, forming distinct parallel chords. Aperture pyriform, angled in its upper part and rather narrow, wider at the base and appearing truncated in fron¬ tal view. Columella thickened, almost straight to curved, abruptly terminating basally. Shell size H = 7.6^8.0 mm, W = 3.1-22.0 mm (Table 2). Operculum. The operculum is typical for thiarids, oval and paucispiral, light to dark brown, and with the nucleus being excentric in the lower left corner. Juvenile shell. The shells of the juveniles in the brood pouch had up to flve whorls, with a maximum height of about 2.5 mm. The protoconch is smooth, with the radi¬ al and spiral sculpture developing on the first teleoconch whorls (Fig. 6). For measurements of the embryonic shell, see Table 3. Radula. Taenioglossate (Fig. 7), resembling other thiarids. As in all thiarids the central tooth or rachidian is significantly wider than tall; all specimens have a cen¬ tral cusp flanked by three to six triangular denticles on both sides, resulting in up to 12 denticles and a typically 4-5/1/4-5 pattern at the upper cutting edge (Fig. 7A, C, E, Table 4). The laterals are equipped with three to six zse.pensoft.net 384 Dusit Boonmekam et al.: Evolutionary systematics of Thiara aspera Table 2. Shell parameters ofThiard' aspera (Lesson, 1831) specimens from Thailand, Indonesia and Australia, with min./max. values, mean, standard deviation (SD), and number of whorls. Voucher Country, region n Measurements (mm) NW H W AL AW BW USNM 119778 Indonesia, Ambon Island 1 23.7 9.7 9.8 5.1 15.5 4 USNM 119722 Philippines, Negros island 1 20.3 7.7 7.1 3.3 12.3 6 MNHN 21098 Indonesia, West Papua 1 25.0 7.8 6.9 3.5 12.6 7 GSUBg 14265 Indonesia, Java 1 48.0 22.0 22.0 10.0 28.1 7 ZMB 107002 India, Calcutta 1 17.4 7.5 5.7 2.5 12.0 3 ZMB 107003 Sri Lanka, Colombo 5 Range 13.3-16.6 5.3-6.6 4.3-5.3 2.3-2.5 9.0-11.3 4-5 Mean 14.3 5.9 4.8 2.4 9.8 SD 1.2 0.4 0.3 0.1 0.8 SUT 0311053 Thailand, Samut Sakhon 30 Range 7.6-12.8 3.1-5.5 2.8-6.2 1.6-3.4 4.4-7.8 4-7 Mean 9.5 3.9 4.0 2.3 5.7 SD 1.1 0.6 0.6 0.4 0.9 SUT 0311020 Thailand, Samut Sakhon 52 Range 14.1-24.3 7.1-11.1 7.0-11.1 3.2-5.4 9.7-16.2 6-7 Mean 17.7 8.7 8.7 4.2 12.0 SD 2.5 1.0 1.0 0.5 1.5 SUT 0311044 Thailand, Samut Sakhon 1 22.9 10.9 10.5 4.8 15.2 6 SUT 0312070 Thailand, Nakhon Pathom 12 Range 14.4-19.8 5.3-7.9 5.7-6.8 2.4-4.2 7.6-12.0 5-8 Mean 17.1 6.8 7.0 3.4 9.9 SD 1.5 0.7 0.9 0.5 1.2 ZMB 191488 Indonesia, Bali 2 Range 19.8-22.1 8.3-9.4 8.0-8.8 4.1-4.7 12.8-14.5 5 Mean 20.9 8.9 8.4 4.4 13.7 SD 1.1 0.6 0.4 0.3 0.9 ZMB 191278 Indonesia, Sulawesi 19 Range 14.3-18.2 6.2-7.8 6.1-8.9 3.1-3.9 9.3-12.3 4-5 Mean 16.1 6.7 7.2 3.6 10.4 SD 1.0 0.3 0.6 0.2 0.7 ZMB 107377 Indonesia, Sulawesi 10 Range 13.6-20.3 5.5-7.5 4.5-7.3 2.7-4.0 7.7-11.7 4-5 Mean 16.8 6.5 6.2 3.4 9.6 SD 2.1 0.7 0.8 0.4 1.3 ZMB 107378 Indonesia, Sulawesi 19 Range 12.9-19.6 5.2-7.1 5.1-7.8 2.6-3.9 7.7-11.2 4-5 Mean 16.9 6.3 6.3 3.2 9.9 SD 1.5 0.4 0.7 0.4 0.8 ZMB 191279 Indonesia, Bali 17 Range 16.8-27.0 7.0-9.9 7.9-12.0 3.3-5.1 11.0-17.2 4-6 Mean 22.7 8.6 9.9 4.3 14.5 SD 3.7 1.0 1.4 0.6 2.2 ZMB 106472 Indonesia, Bali 16 Range 17.9-22.8 6.4-9.3 7.0-10.6 3.1-5.0 11.4-16.2 4-7 Mean 20.1 7.5 8.5 3.8 12.7 SD 1.4 0.6 0.9 0.4 1.2 ZMB 127538 Indonesia, Bali 20 Range 18.^30.4 8.6-13.7 9.3-14.9 4.4-7.4 12.8-20.7 4-6 Mean 24.8 11.0 11.8 5.6 16.8 SD 3.0 1.2 1.4 0.7 1.9 ZMB 191268 Indonesia, Sulawesi 5 Range 29.3-41.5 11.5-16.0 12.1-18.7 5.7-8.4 12.3-26.8 4-6 Mean 35.3 13.5 15.3 6.9 20.9 SD 4.9 2.0 2.5 1.1 5.3 Table 3. Measurements of parameters of the juvenile protoconch ofThiard" aspera (Lesson, 1831) of specimens obtained from the brood pouch. Voucher Country, region n Measurements (^m) he we de ZMB 127534 Thailand, Samut Sakhon 3 Range 48.0-63.2 96.0-120.0 312.7-395.7 Mean 54.0 106.7 352.8 ZMB 127535 Thailand, Samut Sakhon 2 Range 56.3-71.4 107.0-114.3 354.3-366.2 Mean 63.9 110.7 352.8 ZMB 191278 Indonesia, Sulawesi 2 Range 34.0-72.4 76.0-91.1 202.0-252.4 Mean 53.2 84.6 227.2 ZMB 191488 Indonesia, Bali 1 - 83.3 95.2 259.5 zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 373-390 385 Figure 7. Radulae ofThiard’’ aspera (Lesson, 1831) from Thailand. A, B. SUT 0312070, Nakhon Fathom province, pond at Silpa- korn University campus; A. Central and lateral teeth; B. Marginal teeth; C, D. SUT 0311020, Samut Sakhon province, Klong Don Ko; C. Central and lateral teeth; D. Marginal teeth. E, F: SUT 0311053, Samut Sakhon Province, Klong Don Ko; E. Central and lateral teeth; F. Marginal teeth. Scale bars; 35 pm (A, E); 5 pm (B, F); 25 pm (C); 10 pm (D). Table 4. Variation of cusps on the radula teeth ofThiard’’ aspera (Lesson, 1831) specimens. Voucher Country, region n Marginal teeth Lateral teeth (left) Lateral teeth (right) Rachidian SUT 0311053 Thailand, Samut Sakhon 4 6-8 3-1-3 3-1-3 4-5-1-4-5 SUT 0311020 Thailand, Samut Sakhon 4 6-8 3-1-3 3-1-3 4-5-1-4-5 SUT 0312070 Thailand, Nakhon Fathom 2 7-8 5-1-5 4-1-4 4-1-4 SUT 0312069 Thailand, Nakhon Fathom 2 9-10 3-1-3 3-1-3 3-1-3 ZMB 191278 Indonesia, Sulawesi 2 7-8 3-1-3 3-1-3 4-1-4 ZMB 191488 Indonesia, Bali 1 10 6-1-6 6-1-6 6-1-5 ZMB 191279 Indonesia, Bali 3 6-7 4-1-4 4-1-4 4-5-1-4-5 ZMB 106472 Indonesia, Bali 2 6-7 3-1-3 3-1-3 4-1-4 zse.pensoft.net 386 Dusit Boonmekam et al.: Evolutionary systematics of Thiara aspera smaller denticles on the inner side, and three to six den¬ ticles outside from the large main cusps (Fig. 7A, C, E, Table 4). The marginal teeth are moderately long, spoon¬ shaped, with a varying number of 6-10 denticles (Fig. 7B, D, F, Table 4). Reproductive strategy. The results of the analysis of brood pouch content are summarised in Figure 2. Juveniles of up to 2 mm (rarely also larger) were found in the populations from Thailand, Indonesia (Bah and Sulawesi) and Australia (Northern Territory) suggesting an euviviparous reproduc¬ tive strategy for‘T.” aspera, i.e, the taxon was found to give birth to crawling and shelled juveniles in accordance with the definitions in Glaubrecht et al. (2009). In a few popula¬ tions in Thailand and on Bah, gravid females with only early embryos, i.e., veliger larvae in the brood pouch were found. Distribution. ''Thiara'' aspera as here understood is a widespread species, with records from Sri Lanka and India (Subba Rao 1989), Myanmar and Cambodia (van Benthem Jutting 1956), Indonesia (Rensch 1934; van Benthem Jutting 1956), and the Philippines (Lea and Lea 1851; van Benthem Jutting 1956). As our results indicate, the taxon is also present in Thailand and northern Austra¬ lia, from where it was not previously reported (Fig. 2). Discussion The results of our phylogenetic analyses show that the nominal taxon Melania winteri Busch, 1842 is closely related to Thiara amarula and can be classified with the same genus. However, the nominal species Melania as¬ pera Lesson, 1830 (= Melania rudis Lea & Lea, 1851), which has often been classified as a member of Thiara (e.g., van Benthem Jutting 1937, 1956; Subba Rao 1989; Ramakrishna and Dey 2007; Budha 2010; Patil and Tal- male 2011 under the name T rudis) cannot be includ¬ ed within that genus on the basis of our data without broadening the concept of Thiara to an extent that it en¬ compasses almost the entire conchological diversity of Thiaridae because "Stenomelania" denisoniensis Brot, 1877, which is conchologically similar to Stenomela¬ nia Fischer, 1885 or Melanoides Olivier, 1804, clusters within Thiara s. lat. in our phylogenetic analyses and an approximately unbiased test rejected the monophyly of T amarula, T winteri, T. cf winteri, and "T'. aspera, i.e., excluding "Stenomelania" denisoniensis. Pending a phy¬ logenetic analysis of the entire family, we here retain the species in Thiara, but indicate the tentative placement by quotation marks. Our phylogenetic analyses further show that "T." as¬ pera exhibits little genetic variation throughout the In- do-Malayan Archipelago and the Southeast Asian main¬ land, although populations vary considerably with regard to shell shape, and especially sculpture, confirming previ¬ ous surveys on thiarid species, which showed also an ex¬ traordinary plasticity of the shell (Glaubrecht et al. 2009). We here report the presence of "T." aspera in Thailand for the first time, albeit in anthropogenic habitats. Previ¬ ous surveys of the Thai freshwater snail fauna, e.g., by Brandt (1974) did not record the species. Thus, as his years-long surveys were exhaustive it is safe to assume that the species is probably introduced but additional sur¬ veys should be carried out to clarify whether the taxon also occurs in natural habitats in this country and was only overlooked in the past. We also report "T." aspera here for Australia for the first time, where the taxon was found in natural habitats in the Northern Territory and in north-western Queens¬ land. The populations from Australia were found to be somewhat differentiated genetically from the remaining specimens of "T." aspera from Thailand and Indonesia included in the phylogenetic analyses and also slightly differ conchologically, i.e., the spiral sculpture almost disappears on the upper half of the teleoconch whorls. Further analysis should therefore confirm whether these differences are constant and would allow a taxonomic separation of the Australian populations. Unfortunately, no samples could be included in the phylogenetic analyses from either India or Sri Lanka. However, as the examined material closely resembles the holotype of Melania aspera in shell characters (al¬ though this specimen is exceptional because of its very small aperture in relation to total shell height which may explain its isolate position in Fig. 4A), the populations from these two countries are here regarded as belonging to the species. Therefore, "T." aspera has to be consid¬ ered as a widespread species, ranging from India and Sri Lanka across the Southeast Asian mainland and islands into Australia (Fig. 2). At present, our data on "T" aspera do not allow to assess whether the observed differences of juvenile stages in the brood pouch of the female indicate differences in the reproductive strategy, or rather individual or seasonal variations. The close phylogenetic relationships among these populations (Fig. 5), however, let the latter two ex¬ planations appear more likely in our opinion. Therefore, we consider "T." aspera a euviviparous species, although it has to be stated that repeated periodic sampling would be necessary to resolve this issue conclusively. Conclusions These results highlight the need for a comprehensive revision of the genus-group systematics of Thiaridae as a whole. However, mitochondrial DNA markers are fraud with difficulties in some freshwater cerithioide- ans (Kohler and Deein 2010; Whelan and Strong 2015; Kohler 2016) and probably also in Thiaridae. Likewise, there appears to be a confusing variability in shell and reproductive features in thiarids, which is in stark con¬ trast to a conserved radular morphology as compared to some other cerithioidean families (Glaubrecht 1996). A stable system of the family, which ought to include the zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 373-390 387 type species of all named genus-group taxa, can be ex¬ pected to emerge only after phylogenetic analyses based on suitable molecular markers and/or detailed morpho¬ logical data become available. A stable system of the family then could serve as a basis for a better under¬ standing of the evolutionary systematics and phyloge- ography of the group. Acknowledgements We thank Thomas von Rintelen and Christine Zorn for access to the collection housed at the Museum fur Naturkunde, Berlin. We are indebted to the German Ac¬ ademic Exchange Service DAAD for grants in support of research in Thailand and the Deutsche Forschungs- gemeinschaft DFG for a grant (DFG GL 297/19-1) mak¬ ing research in Australia possible. We are also indebted to the support fund of the Faculty of Science, Silpa- korn University, Thailand and the Thailand Research Fund (The Royal Golden Jubilee Ph.D. Programme PHD/0195/2551) for funding. We thank Vince Kessner (Adelaide River) and Richard Willan (Darwin) for help¬ ing with the fieldwork and handling of collections in Aus¬ tralia. We also thank Frank Kohler (Sydney) and Zoltan Feher (Budapest) for constructive comments on a draft version of the manuscript. References Adams H, Adams A (1854) The genera of Recent Mollusca; arranged according to their organization. London, van Voorst. Vol. 1, Part 10, Van Voorst, London, 289-320. [pis 37-40] Agassiz L (1842) Nomina systematica generum molluscorum, tarn viven- tium quam fossilium, secundum ordinem alphabeticum disposita, ad- jectis auctoribus, libris in quibus reperiuntur, anno editionis, etymo- logia et familiis ad quas pertinent. In: Agassiz L (Ed.) Nomenclator zoologicus, continens nomina systematica generum animalium tarn viventium quam fossilium. 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BERLIN Establishment of a new shrimp family Chlorotocellidae for four genera previously assigned to Pandalidae (Decapoda, Caridea, Pandaloidea) Tomoyuki Komai\ Tin-Yam Chan^, Sammy De Grave^ 1 Natural History Museum and Institute, Chiba, 955-2 Aoba-cho, Chuo-ku, Chiba 260-8682, Japan 2 Institute of Marine Biology and Center of Excellence for the Oceans, National Taiwan Ocean University, 2 Pei-Ning Road, Keelung 20224, Taiwan 3 Oxford University Museum of Natural History, Parks Road, Oxford, UK http://zoobank.org/86895CA3-596A-4015-8350-82EEF10F9885 Corresponding author: Tin-Yam Chan (tychan@mail.ntou.edu.tw) Q6.\ioY. Kristina von Rintelen ♦ Received 8 May 2019 ♦ Accepted 17 June 2019 ♦ Published 3 July 2019 Abstract A new caridean shrimp family, Chlorotocellidae, is established to accommodate four genera previously assigned to Pandalidae, viz., Chlorotocella Balss, 1914 (type genus), Chlorocurtis Kemp, 1925, Anachlorocurtis Hayashi, 1975, and Miropandalus Bruce, 1983, which represents the sister clade to a clade consisting of all other pandalid genera (including the two genera previously assigned to Thalassocarididae) in a recent comprehensive phylogenetic analysis of Pandaloidea. Diagnoses are provided for the new family and its constituent genera, and a comparison with Pandalidae is provided, for which a new diagnosis is given. Key Words Anachlorocurtis, ASR analysis, Chlorotocella, Chlorocurtis, Miropandalus Introduction The caridean family Pandalidae Haworth, 1825 is pre¬ dominantly composed of cold-water and deep-sea taxa, including several species of commercial importance (Holthuis 1980). An extensive multimarker molecular phylogenetic analyses of the family and the closely re¬ lated Thalassocarididae by Liao et al. (2019) clarified that Thalassocarididae, represented by two genera, Tha- lassocaris Stimpson, 1860 and Chlorotocoides Kemp, 1925, are deeply nested within Pandalidae, and that four genera, Chlorotocella Balss, 1914, Chlorocurtis Kemp, \925, Anachlorocurtis YidiydiAhi, 1975, dcnd Miropandalus Bruce, 1983, comprise the sister clade to the remaining clade consisting of all other pandalid genera. These four genera are represented by small-sized species inhabiting shallow subtidal waters in tropical to subtropical and often associated with cnidarians (Hayashi and Miyake 1968; Hayashi 1975; Bruce 1983; Okuno and Yokota 1995; Minemizu 2000, 2013; Kato and Okuno 2001; Kawamoto and Okuno 2003; Humann and DeLoach 2010; Horka et al. 2014; Anker and De Grave 2016) in sharp contrast to the ecologies of most Pandalidae. To incorporate the documented phylogenetic pattern into the formal classification of Caridea, together with consider¬ ations on the morphological distinctness and ecological traits of these taxa, we hereby propose a new family, Chlorotocellidae fam. nov. for these four genera and re¬ define the family Pandalidae. Materials and methods The morphological data assembled following an exam¬ ination of the literature and direct examination of speci¬ mens of relevant taxa formed the basis of the phylogenet¬ ic analysis by Liao et al. (2019) and can be found in the online supplementary material for that study. Aside from the family diagnosis, diagnoses are provided for each ge¬ nus in the new family; to shorten diagnoses, synapomor- phies for the family are not repeated in generic diagnosis; autapomorphies are in bold italics. Copyright Tomoyuki Komai et al. This is an open access articie distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the originai author and source are credited. 392 Tomoyuki Komai et al.: New caridean family Chlorotocellidae Illustrations showing diagnostic characters are given for Chlorotocella (C. gracilis Balss, 1914) and Chlorocurtis (C. jactans (Nobili, 1904)), as no published modem illustrations are available in easily accessible literature for those taxa. The three species of Anachlorocurtis and the monotypie Miro- pandaliis have been well illustrated in their respective type description, and thus, no additional figures are presented. Details of specimens used for preparation of draw¬ ings are listed below. These specimens are deposited in the Laboratory of Marine Zoology, Faculty of Fisheries, Hokkaido University, Hakodate, Japan (HUMZ), Nation¬ al Research Institute for Far Seas Fisheries, Shizuoka, Japan (NRIFSF), and the Natural History Museum and Institute, Chiba, Japan (CBM). Chlorotocellidae fam. nov. Chlorocurtis jactans (Nobili, 1904): CBM-ZC 11596, 1 ovigerous female (cl 1.3 mm), Uehara, Iriomote Island, Yaeyama Islands, Ryukyu Islands, sea grass beds, 0.5-1 m at low tide, 18 July 2007, dip net, coll. T. Komai. Chlorotocella gracilis Balss, 1914: CBM-ZC 12534, 1 male (cl 5.0 mm), TRV “Toyoshio-maru”, 2001-6 cruise, stn 4, W of Tanegashi- ma Island, Ohsumi Islands, 30°33.50'N, 130°53.30'E, 47 m, 26 May 2001, dredge, coll. T. Komai; HUMZ-C 1556, 1 male (cl 4.1 mm), Tosa Bay, 40 m, 22 May 1960, coll. M. Toriyama. Pandalidae Haworth, 1825 Chlorotocus novaezealandiae (Borradaile, 1916): NRIFSF 578, 1 male (cl 17. 0 mm). New Zealand, no other data. Heterocarpus ensifer A. Milne-Fdwards, 1881: HUMZ-C 255, 1 male (cl 33.0 mm), no data. Pandalus montagui Feach, 1814: CBM-ZC 3422,1 transitional male (cl 10.5 mm), Texel, The Netherlands, 8 April 1991. Systematics Infraorder Caridea Dana, 1852 Superfamily Pandaloidea Haworth, 1825 Family Pandalidae Haworth, 1825 Type genus. Pandalus Leach, 1814, by original designation. Composition. Atlantopandalus Komai, 1999, Austropan- dalus Holthuis, 1950, Bitias Fransen, 1990, Chelonika Fransen, 1997, Chlorotocoides Kemp, 1925, Chloroto¬ cus A. Milne-Edwards, 1882, Dichelopandalus Caullery, 1896, Dorodotes Bate, 1888, Heterocarpus A. Milne-Ed¬ wards, 1881, Heteronika Hendrickx, 2019, Notopanda- lus Yaldwyn, 1960, Pandalina Caiman, 1899, Pandalus Eeach, 1814, Pantomus A. Milne-Edwards, 1883, Peri- pandalus de Man, 1917, Plesionika Bate, 1888, Procletes Bate, 1888, Pseudopandalus Crosnier, 1997, and Thalas- socaris Stimpson, 1860. Diagnosis. Rostrum well developed, usually ven- trally with teeth or rows of setae. Thoracic sternites 6-8 each with paired conspicuous prominences, teeth or protuberances (Fig. 3A). Pleomere 6 posterolateral process usually terminating in small tooth. Telson with longitudinal row of spiniform setae located on dorso¬ lateral ridges. Eyestalks subpyriform or kidney-shaped, cornea distinctly longer and wider than eyestalk. Anten- nular stylocerite with proximolateral projection, distal- ly acuminate or rounded; article 2 usually with minute spiniform setae (Fig. 6A); outer fiagellum with distal portion (distal to aesthetasc-bearing portion) usually well developed, consisting of numerous articles. Article 1 of mandibular palp with prominent expansion on inner distal margin (Fig. 6B). Maxilliped 2 with podobranch. Maxilliped 3 with or without exopod. Pereopod 1 fingers minute or completely reduced. Pereopod 2 subequal or unequal; basis with small process on lateral surface (Fig. 6C); carpal articulation greatly variable, but never tri-ar- ticulated. Arthrobranchs usually present on maxilliped 3 and pereopods. Chlorotocellidae fam. nov. http://zoobank.org/CBE45390-043A-46E8-9743-7F5B167255AD Type genus. Chlorotocella Balss, 1914, by present designation. Composition. Chlorotocella Balss, 1914 (two species), Chlorocurtis Kemp, 1925 (monotypie), Anachlorocurtis Hayashi, 1975 (three species) and Miropandalus Bruce, 1983 (monotypie). Diagnosis. Rostrum, if present, without teeth or fringe of setae on ventral margin (Figs lA, 4A). Thoracic sternites without conspicuous ornamentation, such as prominenc¬ es, teeth or protuberances (Fig. 3B). Pleomere 6 postero¬ lateral process rounded or truncate (Figs 1C, 4B). Telson with dorsolateral spiniform setae located adjacent to lateral margins (Figs ID, 4C). Eyestalks subcylindrical, cornea distinctly shorter than eyestalk (Figs IE, 4D). An- tennular stylocerite devoid of proximolateral projection, distally obliquely truncate, bi- or tridentate (Figs lA, F, 4E); outer fiagellum with distal portion (distal to aesthe¬ tasc-bearing portion) reduced, consisting only of few ar¬ ticles (Figs lA, 4E). Maxilliped 2 without podobranch ( Figs IE). Maxilliped 3 with no exopod (Figs 2A, 5A). Pereopod 1 fingers absent (Figs 2B, 5B, C). Pereopod 2 always subequal; basis without small process on lateral surface of basis; carpus consistently divided into three articles (Figs 2C, 5D, 6D). Arthrobranchs always absent from maxilliped 3 and pereopods. Remarks. Characters differentiating Chlorotocellidae fam. nov. and Pandalidae are summarized in Table 1, with the character states of Chlorotocellidae fam. nov. being synapomorphic against Pandalidae (see Eiao et al. 2019). zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 391-402 393 Figure 1. ChJorotocella gracilis Balss, 1914, male (cl 5.0 mm), CBM-ZC 12534. A. Carapace, antennule and antenna, lateral view (left eye removed); B. Anterior part of carapace, lateral view; C. Pleon, lateral view; D. Telson, dorsal view; E. Left eye, dorsal view; F. Left antennular peduncle, dorsal view; G. Left antenna, ventral view; H. Left mandible, outer view; inset, palp, outer view; 1. Left maxillule, outer view (coxal endite missing); J. Left maxilla, outer view; K. Left maxilliped 1, outer view; L. Left maxilliped 2, outer view; M. Endopod of left pleopod 1, ventral view. zse.pensoft.net 394 Tomoyuki Komai et al.: New caridean family Chlorotocellidae Figure 2. Chlorotocella gracilis Balss, 1914, male (cl 5.0 mm), CBM-ZC 12534. Left thoracic appendages in lateral view. A. Max- illiped 3; B-F. Pereopods 1-5, respectively. zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 391-402 395 Figure 3. Thoracic sternites 6-8 and coxae (cx) of pereopods 3-5, ventral view. A. Chlorotocus novaezealandiae (Borradaile, 1916), male (17.0 mm), NRIFSF 578 (arrows indicating prominences on sternites 6-8); B. Chlorotocella gracilis Balss, 1914, male (cl 4.1 mm), HUMZ-C 1556. Table 1. Comparison of diagnostic characters between Chlorotocellidae fam. nov. and Pandalidae. Character states of Chlorotocel- lidae are apomorphic against those of Pandalidae. Chlorotocellidae fam. nov. Pandalidae 1 Rostrum, if present, without teeth or fringe of setae on ventral margin (Figs lA, 4A). Rostrum usually well developed, ventral margin armed with a few or series of teeth accompanied by row(s) of short setae. 2 Thoracic sternites without conspicuous ornamentation, such as keels, teeth or protuberances (Fig. 3B). Thoracic sternites 6 and 7 each with paired prominences on either side of median keel; thoracic sternite 8 with transverse carina bearing submedian spines or teeth (Fig. 3A). 3 Pleomere 6 without posteroventral tooth; posterolateral process rounded or truncate (Figs 1C, 4B). Pleomere 6 with small posteroventral tooth; posterolateral process terminating in small tooth. 4 Telson with dorsolateral spiniform setae located adjacent to lateral margins (Figs ID, 4C). Telson with dorsolateral spiniform setae located on dorsolateral ridges. 5 Eyestalks subcylindrical, cornea distinctly shorter than eyestalk (Figs IE, 4D). Eyestalks pyriform or subpyriform, cornea distinctly wider than eyestalk. 6 Antennular stylocerite distally obliquely truncate, bi- or tri- dentate, devoid of proximolateral projection (IF, 4E). Antennular stylocerite acuminate or rounded, usually having small proximolateral projection (Fig. 6A). 7 Outer antennular flagellum with distal portion (distal to thickened aesthetasc-bearing portion) reduced, consisting of few articles (Figs lA, 4E). Outer antennular flagellum with distal portion (distal to aesthetasc- bearing portion) long and slender, consisting of 10 or more articles. 8 Maxilliped 2 without podobranch (Fig. IL). Maxilliped 2 with podobranch. 9 Maxilliped 3 without exopod (Figs 2A, 5A). Maxilliped 3 with or without exopod. 10 Pereopod 1 fingers absent (Figs 2B, 5B, C). Pereopod 1 fingers minute or completely reduced. 11 Pereopods 2 always subequal; basis without small process on lateral surface of basis; carpus consistently divided into 3 articles (Figs 2C, 5D, 6D). Pereopods 2 subequal or unequal; basis without small process on lateral surface of basis; division of carpus highly variable, but never 3-articulated. 12 Arthrobranchs absent from maxilliped 3 and pereopods. Arthrobranchs usually present on maxilliped 3 and pereopods 1-4. Supplementary figures of diagnostic characters can be found in Hayashi (1975: figs 1-3), Bruce (1983: figs 1-5), Hayashi (2007a: figs 538, 539, 542a-f), Hayashi (2007c: figs 557-559a-e), Horka et al. (2014: figs 1-8), and Ahyong (2015: figs 9, 10). Amongst these characters, the division of the carpus of pereopod 2 and quite possibly the absence of ventral rostral teeth can readily be used to differentiate the two families, although determination of their polarity is not straightforward. In Chlorotocellidae the pereopod 2 car- zse.pensoft.net 396 Tomoyuki Komai et al.: New caridean family Chlorotocellidae Figure 4. Chlorocurtis jactans (Nobili, 1904), ovigerous female (cl 1.3 mm), CBM-ZC 11596. A. Carapace, lateral view; B. Pleon, lateral view; C. telson, dorsal view; D. Left eye, dorsal view; E. Left antennule, dorsal view (inner flagellum damaged); F. Right anten- nular peduncle, distal 2 articles, mesial view; G. Left antenna, ventral view (flagellum missing); H. Left mandible, outer view; 1. Left maxillule, outer view; J. Left maxilla, outer view; K. Left maxilliped 1, outer view; L. Left maxilliped 2, outer view (epipod broken ofl). zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 391-402 397 Figure 5. Chlorocurtis jactans (Nobili, 1904), ovigerous female (cl 1.3 mm), CBM-ZC 11596, left thoracic appendages in lateral view (except for C). A. Maxilliped 3; B. Pereopod 1; C. Same, propodus, mesial view; D-G. Pereopods 2-5, respectively. pus is consistently divided into three articles, whereas in Pandalidae, the number of the carpal articles is quite variable according to taxa, but none are tri-articulate (cf. Komai 1994). The absence of ventral rostral teeth is also reported in three taxa of the pandalid genus Plesionika (Chace 1985) but with doubt (see Chace 1985; Komai et al. 2005; Hayashi 2009; Komai 2011; Li and Chan 2013). Such a similarity, if really present, can be resulted from homoplasy (Liao et al. 2019). According to the Ancestral State Reconstruction (ASR) analysis by Liao et al. (2019), Pandalidae is char¬ acterized by the following synapomorphic features: (1) second article of the antennular peduncle with a few min¬ ute spiniform setae on the dorsodistal margin (Fig. 6A); (2) mandibular palp eonsisting of three articles (Fig. 6B); and (3) basis of pereopod 2 bearing a small process on the lateral surface (Fig. 6C). In these regards, Chloro- tocellidae shows the following plesiomorphic states: (1) mandibular palp tends to be reduced, being absent or consisting of two articles at most (Figs IH, 4H); (2) second article of the antennular peduncle is unarmed on the dorsodistal margin (Figs lA, F, 4E); and (3) basis of pereopod 2 being unarmed (Fig. 6D). Nevertheless, an assessment of the polarity of the de¬ velopment of the mandibular palp is fraught with diffi¬ culty and heavily dependent on outgroup selection. In Caridea in general, however, a reduction of the mandib¬ ular palp is considered to be derived (e.g., Christoffersen 1987, 1989), as compared to the well-developed palp in Dendrobranchiata and most other Decapoda. Furthermore, the other two eharacters are subject to re¬ versal within Pandalidae (Komai 1994; Liao et al. 2019). In species of Thalassocaris and Chlorotocoides (previ¬ ously in their own family Thalassocarididae, but now considered part of Pandalidae), the second article of the antennular peduncle is devoid of spiniform setae and the basis of the second pereopods unarmed (Komai 1994). In addition to the three aforementioned characters, the possession of a rounded laminar expansion at the inner distal angle of the first article of the mandibular palp zse.pensoft.net 398 Tomoyuki Komai et al.: New caridean family Chlorotocellidae Figure 6. A, B. Pandalus montagui Leach, 1814, transitional male (cl 10.5 mm), CBM-ZC 3422; C. Heterocarpus ensifer A. Milne-Edwards, 1881, male (cl 33.0 mm), HUMZ-C 255; D. Chlorotocella gracilis Balss, 1914, male (cl 4.1 mm), HUMZ-C 1556. A. Left antennular peduncle, dorsal view; B. Left mandible, outer view; inset, palp, lateral view; C. Basis of left pereopod 2; D. Basis of right pereopod 2. (Fig. 6B, inset) might be synapomorphic to Pandalidae (Komai 1994; Liao et al. 2019), although a secondary loss of this structure is observed in Thalassocaris and Chlo- rotocoides (Komai 1994; Liao et al. 2019). It is impos¬ sible to evaluate the homology of this character for the taxa assigned to Chlorotocellidae, because in those taxa, the mandibular palp only comprises two articles {Chloro¬ tocella) or is absent {Chlorocurtis, Anachlorocurtis and Miropandalus), and the homology of the articles has not been established in taxa with different numbers. Genus Chlorotocella Balss, 1914 Chlorotocella Bahs 1914: 33; Holthuis 1955: 118, 127; 1993: 263, 266; Hayashi 2007a: 150. Type species. Chlorotocella gracilis Balss, 1914. Diagnosis. Rostrum elongate, very slender, gently up¬ turned, exceeding far beyond distal margin of antennal scaphocerite, dorsally armed with two teeth around ros¬ tral base (one postrostral); ventral margin unarmed (Fig. lA). Carapace without projections on dorsal midline; supraorbital tooth present, suborbital lobe prominent, longer than antennal tooth, distally rounded, slightly constricted at base, pterygostomial tooth moderately small (Fig. lA, B). Pleomeres 1-6 dorsally rounded; pleomeres 4 and 5 each with pair of posterolateral teeth, pleomere 5 with deep transverse groove near posterodorsal margin, pleuron with small posteroven- tral tooth (Fig. 1C). Pleomere 6 with minute postero¬ median tooth, posteroventral angle with minute tooth (Fig. 1C). Telson with additional anterior pair of spin- iform setae located more mesial to other lateral series of spiniform setae, posterior margin narrow, slightly produced medially, with two pairs of unequal spiniform setae (Fig. ID). Eye with ocellar spot (nebenauge) (Fig. IE). Antennular peduncle article 1 armed with tooth on dorsodistal margin (Fig. lA, F). Mandible with two-ar¬ ticulated palp (Fig. IH). Maxillule palp without distal outer lobule (Fig. II). Maxilla with short, moderately slender endopod (Fig. IJ). Maxilliped 1 with coxal and basial endites well developed, both with row of setae on mesial margin; exopodal flagellum well developed (Fig. IK). Maxilliped 2 endopod with dactylus located at distal portion of propodus; exopod well developed (Fig. IE). Pereopod 1 Angers completely reduced (Fig. 2B). Pereopods 3-5 propodi each with closely spaced, short to long spiniform setae in distal 0.2; carpi each with few spiniform setae on lateral surface; meri usually with spiniform setae arranged in two rows; ischia each with spiniform seta on ventral surface in pereopods 3 and 4 (Fig. 2D-F). Male pleopod 1 endopod without ap¬ pendix interna (Fig. IM). Composition. Chlorotocella gracilis', C. spinicaudus (H. Milne Edwards, 1837). Distribution. Indo-West Pacific, South Australia; shal¬ low subtidal to 60 m; free living in algal-rich habitats or facultatively associated with gorgonarians and hydroids. zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 391-402 399 Remarks. At present, two species are assigned to Chlo- rotocella (De Grave and Fransen 2011), viz., C. graci¬ lis (type species) and C. spinicaudus. Holthuis (1995) clarified that Hippolyte spinicaudus H. Milne Edwards, 1837 was a senior subjective synonym of Pandalus lep- torhynchus Stimpson, 1860. In addition, a third taxon, which was placed in the synonymy of C. spinicaudus by De Grave and Fransen (2011), Pandalus (Parapanda- lus) leptorhynchus var. gibber Hale, 1924, was described from Gulf St Vincent, South Australia, characterized mainly by the prominently crested tergite of pleomere 3 (see Hale 1927). This taxon has been seldom mentioned in more recent literature. Ledoyer (1984) illustrated a specimen with a weakly crested tergite from Noumea (New Caledonia), which he assigned to C. gracilis, but left it open as to whether this should be a distinct species or merely a “forme gibbef of C. gracilis. In contrast, Poore (2004) treated the taxon as a distinct species, C. gibber (Hale), noting it was restricted to the Gulf St Vin¬ cent (South Australia). Because no modern descriptions are available for C. spinicaudus, the above generic diagnosis is largely based on C. gracilis and the summary information available on the other species. It seems possible that Hale’s (1924) tax¬ on might be distinct from C. gracilis and C. spinicaudus as it is characteristic by having a highly crested tergite of the pleomere 3 (Hale 1924: pi. 4, fig. 6; 1927: fig. 35). Re¬ assessment of the taxonomic status of C. spinicaudus and Pandalus (Parapandalus) leptorhynchus var. gibber will be necessary to fully clarify the taxonomy of the genus. Genus Chlorocurtis Kemp, 1925 Chlorocurtis Kemp 1925: 272, 279; Holthuis 1955: 118, 127; 1993: 263, 265; Hayashi 2007b: 248. Type species. Chlorocurtis miser Kemp, 1925. Diagnosis. Rostrum short but well developed, direct¬ ed forward, reaching midlength of article 1 of antennu- lar peduncle; dorsal margin crested, with five to seven teeth including two or three postrostral; ventral margin unarmed (Fig. 4A). Carapace without conspicuous pro¬ jections on dorsal midline; no supraorbital tooth; subor¬ bital lobe absent; pterygostomial tooth moderately small (Fig. 4A). Pleomeres 1-5 dorsally rounded; pleomeres 1-3 with long, erect setae on dorsal surface, pleomeres 4 and 5 each without pair of posterolateral teeth; pleo¬ mere 5 without deep transverse groove near posterodorsal margin, pleuron rounded posteriorly; pleomere 6 without posteromedian tooth, posteroventral angle unarmed (Fig. 4B). Telson posterior margin rather broad, convex, with three pairs of unequal spiniform setae (Fig. 4C). Eye without ocellar spot (nebenauge) (Fig. 4D). Antennular peduncle article 1 unarmed on dorsodistal margin; stylo- cerite obliquely truncate distally, distolateral angle termi¬ nating in tooth, distomesial angle subacute or blunt; outer flagellum shorter than peduncle, distal portion reduced to single article (Fig. 4E). Short, club-like, modified setae present at ventrodistal margin of article 2 of antennu¬ lar peduncle (one seta) and distal margin of antennal scaphocerite (two setae) (Fig. 4F, G.). Mandible with¬ out palp (Fig. 4H). Maxillule palp with well-developed distal outer lobule (Fig. 41). Maxilla with short, distal¬ ly tapering endopod (Fig. 4J). Maxilliped 1 with coxal and basial endites well developed, both with row of setae on mesial margin; exopodal flagellum well developed (Fig. 4K). Maxilliped 2 endopod with dactylus located at distal portion of propodus; exopod well developed (Fig. 4E). Pereopods 3-5 propodi broadened distally, oblique flexor distal margins each with short rows of narrowly spaced long spiniform setae flanking field of short setae, forming prehensile structure together with dactylus folded back, carpi without spiniform setae on lateral surface; meri without spiniform setae; ischia with¬ out spiniform seta on ventral surface (Fig. 5E-G). Male pleopod 1 endopod without appendix interna. Composition. Monotypic. Distribution. Indo-West Pacific, intertidal to 10 m; sea- grass beds. Remarks. Chlorocurtis was originally established for Chlorocurtis miser by Kemp (1925). Eater, Holthuis (1947) synonymized Chlorocurtis mi¬ ser with Virbius (?) jactans (Nobili, 1904) without any argumentation, although clearly correct. This synonymy has since been widely adopted (e.g., Holthuis 1955; Ee- doyer 1968, 1984; Bruce 1976; Hayashi 2007b; Holthuis 1993; De Grave and Fransen 2011; Gan and Ei 2018). Gtmis Anachlorocurtis Hayashi, 1975 Anachlorocurtis Hayashi 1975: 173; 2007a; 147; Holthuis 1993: 263; Horkaetal. 2014: 12. Type species. Anachlorocurtis commensalis Hayashi, 1975. Diagnosis. Rostrum short, ascending in adults, reaching midlength of article 1 of antennular peduncle, terminating in acute tip or obliquely truncate distally with irregular dentition; dorsal and ventral margins usually unarmed. Carapace without supraorbital tooth; dorsal midline with two prominent processes, anterior one postrostral, irregu¬ larly denticulate anteriorly, posterior one cardiac in posi¬ tion, directed forward, acuminate; suborbital lobe absent; pterygostomial angle rounded, unarmed. Pleomeres 1-5 dorsally rounded; pleomeres 4 and 5 each without pair of posterolateral teeth. Pleomere 5 without deep transverse groove near posterodorsal margin; pleuron rounded pos¬ teriorly. Pleomere 6 without posteromedian tooth; pos- teroventral angle without tooth. Telson posterior margin truncate or rounded, with five pairs of unequal spiniform zse.pensoft.net 400 Tomoyuki Komai et al.: New caridean family Chlorotocellidae setae. Eye without ocellar spot (nebenauge); cornea with papilla-like tubercle. Antennular peduncle article 1 un¬ armed on dorsodistal margin; stylocerite obliquely trun¬ cate distally, both distal angles dentate; outer flagellum shorter than peduncle, distal portion reduced to one or two articles. Mandible without palp. Maxillule palp with well-developed distal outer lobule bearing apical seta. Maxilliped 1 with coxal and basial endites poorly de¬ veloped, narrow; exopodal flagellum absent. Maxilliped 2 endopod with dactylus located at mesial portion of propodus or fused to propodus; exopod absent. Articula¬ tion between carpal article 1 and 2 of pereopod 2 strongly oblique. Pereopods 3-5 propodi slightly narrowing dis¬ tally, with few widely spaced minute spiniform setae on flexor margin; carpi without spiniform setae on lateral surface; meri of pereopods 3 and 4 each with one spin¬ iform seta distolaterally and one minute spiniform seta at midlength of ventral surface. Male pleopod 1 endopod with small rounded lateral lobe far exceeded by well-de¬ veloped appendix interna. For illustrations see Hayashi (1975: flgs 1-3), Hayashi (2007a: flgs 538, 539, 542a-f); Horka et al. (2014: flgs 1-8) and Ahyong (2015: flgs 9, 10). Composition. Anachlorocurtis commensalis, A. occiden- talis Horka, De Grave & Duris, 2014, and A. australis Ahyong, 2015. Distribution. Indo-West Paciflc, shallow subtidal to 40 m; associated with antipatharian corals. Genus Miropandalus Bruce, 1983 Miropandalus Bruce, 1983: 482; Holthuis 1993; 263, 269; Hayashi 2007c: 585. Type species. Miropandalus hardingi Bruce, 1983. Diagnosis. Rostrum absent. Carapace without supra¬ orbital tooth; dorsal midline with two very prominent, erect processes, anterior one postrostral, tapering, pos¬ terior one cardiac in position, slightly curved, anteri¬ orly, blunt, suborbital lobe absent; pterygostomial angle rounded or angular. Pleomere 1 with prominent protu¬ berance, pleomere 3 with triangular crest on posterior half of dorsal midline, pleomeres 4 and 5 each without pair of posterolateral teeth; pleomere 5 without deep transverse groove near posterodorsal margin; pleuron rounded posteriorly. Pleomere 6 without posteromedian tooth; posteroventral angle unarmed. Telson posterior margin rounded, with several short spiniform setae. Eye without ocellar spot (nebenauge); cornea without papil- la-like tubercle. Antennular peduncle article 1 unarmed on dorsodistal margin; stylocerite subtruncate distally, bi- or tridentate, outer flagellum shorter than peduncle, distal portion completely reduced. Mandible without palp. Maxillule palp with well-developed distal outer lobule, without apical seta. Maxilliped 1 with coxal and basial endites poorly developed, narrow; endopod stout; exopodal flagellum absent. Maxilliped 2 endopod with dactylus fused to propodus; exopod absent. Articulation between carpal article 1 and 2 of pereopod 2 strongly oblique. Pereopods 3-5 propodi narrowing distally, with few minute spiniform setae on flexor margin; carpi with¬ out spiniform setae on lateral surface; meri of pereopods 3 and 4 unarmed. Male pleopod 1 endopod with small rounded lateral lobe far exceeded by well-developed ap¬ pendix interna. For illustrations, see Bruce (1983: flgs 1-5) and Ha¬ yashi (2007c: flgs 557-559a-e). Composition. Monotypic. Distribution. West Paciflc, subtidal to 58 m; associated with antipatharian corals. Acknowledgements This work was supported by grants from the Ministry of Science and Technology, Taiwan, R.O.C., and the Cen¬ ter of Excellence for the Oceans (National Taiwan Ocean University), which is flnancially supported by The Fea¬ tured Areas Research Center Program within the frame¬ work of the Higher Education Sprout Project by the Min¬ istry of Education in Taiwan, R.O.C. References Ahyong ST (2015) Decapod Crustacea of the Kermadec Biodiscovery Expedition 2011. Bulletin of the Auckland Museum 20; 405-442. Anker A, De Grave S (2016) An updated and annotated checklist of marine and brackish caridean shrimps of Singapore. Raffles Bulletin of Zoology 34 (Supplement): 343-454. Balss H (1914) Ostasiatische Decapoden II. Die Natantia und Reptan- tia. Abhandlungen der Mathematisch-Physikalischen Klasse der Kbniglich Bayerischen Akademie der Wissenschaften 10 (Supple¬ ment 2): 1-101. 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BERLIN Eulimacrostoma gen. nov., a new genus of Eulimidae (Gastropoda, Caenogastropoda) with deseription of a new speeies and reevaluation of other western Atlantic species Leonardo Santos de Souza\ Alexandre Dias Pimenta^ 1 Malacologia, Departamento de Invertebrados, Museu Nacional, Universidade Federal do Rio de Janeiro, Quinta da Boa Vista, Sdo Cristdvao, 20940-040, Rio de Janeiro, RJ, Brazil http://zoobank.org/954FD888-055A-4151-947A-9055B02DA514 Corresponding author; Leonardo Santos de Souza (Ieosouza2301@gmail.com) AQdLdQViixc Q6.\iQY. Matthias Glauhrecht ♦ Received 14 February 2019 ♦ Accepted 28 June 2019 ♦ Published 25 July 2019 Abstract Anew genus and a new species of Eulimidae are described based on the shell morphology and on the host-parasite relationship of the type species. Eulimacrostoma microsculpturata Souza & Pimenta gen. nov. and sp. nov. parasitizes a starfish of the genus Luidia and has an elongated, conical, straight, or slightly curved shell, a protoconch with a brownish spiral band and convex whorls, a peculiar large and broad aperture with a strongly protruding outer lip, and microsculpture of axial lines on the teleoconch. Four other species are included in the genus, all from the western Atlantic: Eulimacrostoma chascanon (Watson, 1883), comb, nov., Eulimacrostoma fu- sus (Dali, 1889), comb, nov., Eulimacrostoma lutescens (Simone, 2002), comb, nov., and Eulimacrostomapatula (Dali & Simpson, 1901), comb. nov. Newly available material of Eulimacrostoma patula expands the known geographic distribution of this species in the Caribbean to the north coast of Brazil. Eulimacrostoma chascanon and Eulimacrostoma fusus and Eulimacrostoma lutescens are known only by the type series which was re-examined. A redescription is provided for Eulimacrostoma chascanon and Eulimac¬ rostoma fusus. Species within Eulimacrostoma differ mainly by teleoconch sculpture, the presence or absence of an umbilical fissure, and shell dimensions. Lectotypes are designated for Eulimacrostoma chascanon, Eulimacrostoma fusus, and Eulimacrostoma patula. Key Words biodiversity, micromolluscs, parasitic snails shell morphology, taxonomy, Vanikoroidea Introduction Eulimidae Philippi, 1853 is a diverse group of marine gastropods comprising more than 1,000 valid species and more than 90 valid genera (MolluscaBase 2018). A mas¬ sive collection effort in New Caledonia included eulimids as one of the most species-rich families of marine gastro¬ pods and about 80% of the species were possibly new to science (Bouchet et al. 2002). Eulimids are parasites of the five extant classes of Echinodermata and shows dif¬ ferent life strategies (e.g., ectoparasitism, endoparasitism, gall formers) (Waren 1983; Takano and Kano 2014). Waren (1983) suggested that all species in a given genus parasitize a single class of Echinodermata, with the exception of Vitreolina Monterosato, 1884, which is known to exploit ophiuroids and echinoids. Takano et al. (2018: 215) suggested that Vitreolina contains distantly related lineages and, thus, it is probably not monophyletic and Waren’s (1983) rule would apply without exceptions. Genera of Eulimidae that parasitize echinoderms of the class Asteroidea are Apicalia A. Adams, 1862, Aster- olamia Waren, 1980, Asterophila Randall & Heath, 1912, Niso Risso, 1826, Paramegadenus Humphreys & Eiitzen, 1972, Parvioris Waren, 1981, Stilifer Broderip, 1832, and Thyca H. Adams & A. Adams, 1854 (Waren 1983). The shell shape in these genera varies from the usual conical format to capuliform, or the animal is shell-less. All kinds of life strategies are present among them. Copyright Leonardo Santos de Souza, Alexandre Dias Pimenta. This is an open access articie distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the originai author and source are credited. 404 de Souza, L.S. & Pimenta, A.D.: Eulimacrostoma gen. nov., a new genus of Eulimidae... Recently, one eulimid attached to the starfish Luidia ludwigi scotti Bell, 1917 (Luidiidae), collected in the up¬ per bathyal zone otf Florida, USA, was studied by the au¬ thors and did not correspond to any other species and ge¬ nus known to parasitize asteroids or other echinoderms. The availability of a single specimen hampered the de¬ scription of the taxa, but recently we identified shells of the same species from nearby localities in malacological collections, which enabled the present description. Material and methods The material examined is housed in the following mal¬ acological collections: Academy of Natural Sciences of Philadelphia, Drexel University, Philadelphia, Pennsyl¬ vania, USA (ANSP); Florida Museum of Natural History, Florida University, Gainesville, Florida, USA (FLMNH); Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, USA (MCZ); Museu Na- cional, Universidade Federal do Rio de Janeiro, Rio de Janeiro, Rio de Janeiro, Brazil (MNRJ); Natural History Museum, London, United Kingdom (NHMUK); Smith¬ sonian National Museum of Natural History, Washington DC, USA(USNM). During the course of this study, a fire destroyed sever¬ al collections of MNRJ (Zamudio et al. 2018), including material cited herein. Some vouchers and types from oth¬ er institutions on loan were also at the MNRJ and were destroyed. These lots are indicated with an asterisk (*) in the material examined. Some samples were collected by large projects and oceanographic expeditions, such as: (1) Eolis Cruises, organized by John B. Henderson, collected by the yacht Eolis between 1910-1916 in the Florida Keys, USA (see Bieler and Mikkelsen 2002 for approximate coordinates of the collecting stations); (2) Johnson-Smithsonian Deep- sea expedition, collected by the yacht Caroline in 1933, in deep waters of Puerto Rico, and sponsored by Eldridge R. Johnson (see Bartsch 1933); (3) “A Multidisciplinary Amazon Shelf Sediment Study” (AMASSEDS), collect¬ ed by the Research Vessel (RW) Columbus Iselin in 1990 near the mouth of the Amazon River, northern Brazil; (4) “Terres Australes et Antarctiques Fran^aises, cruise MD55” (TAAF MD55), a Joint project of the “Museum National d’Histoire Naturelle”, France, and the “Universi¬ dade Santa Ursula”, Brazil, collected by the R/V Marion Dufresne between May and June 1987 (Tavares 1999). Taxonomic identifications were based on conchologi- cal features in comparison with original descriptions and illustrations and examination of type material. Terminol¬ ogy of shell features follows Bouchet and Waren (1986) and Souza and Pimenta (2019). The outer lip may be opisthocline (connection at the suture behind the distal region), prosocline (connection in front of the distal re¬ gion) or orthocline (in the same plane). Incremental scars are older positions of the outer lip and when present are usually deep and well demarcated; they are formed dur¬ ing the periodical growth of the eulimids (see Bouchet and Waren 1986: 310 for details). Microsculpture of ax¬ ial lines usually appears in regular intervals, from suture to suture in each whorl and do not interrupt the suture as do the incremental scars. Growth lines are usually pres¬ ent at irregular intervals and do not reach from one su¬ ture to the other. Measurements of the shell are based on Souza and Pimenta (2019): shell length (SE); body whorl length (BWE); aperture length (AE); shell width (SW); aperture width (AW). Abbreviations of generic names: Eulima Risso, 1826 (E.); Eulimacrostoma gen. nov. {Eu.). Systematics Family Eulimidae Philippi, 1853 Genus Eulimacrostoma Souza & Pimenta, gen. nov. http://zoobank.org/8C4C4750-0C97-4ABl-B092-lD37ADBAE3FE Type species. Eulimacrostoma microsculpturata Souza & Pimenta, sp. nov. Recent, northwestern Atlantic and Caribbean. Diagnosis. Eulimids parasitic on asteroids. Shell elon¬ gated, conical, straight or slightly curved. Protoconch subcylindrical, smooth. Teleoconch with slightly convex whorls, several incremental scars and microsculpture of axial lines, wide brownish spiral bands, a large and spread aperture, occupying between 60-70% of the body whorl length, and an orthocline outer lip, strongly protruding. Etymology. Eulima, due to the systematic affinity and for being one of the most common names of the family, in combination with Macros, Gr. = long; and Stomatos, Gr. = mouth; in reference to the broad shape of the shell aperture. Eulimacrostoma microsculpturata Souza & Pimenta, sp. nov. http://zoobank.org/F45DF003-4D73-4689-A283-20B0CD5131A6 Figures lA-G, 2A-G Melanella patula auct. non. (Dali & Simpson, 1901): Dali (1927, in part.: 67). Type material. Holotype: USNM 429762. Para- types: USA: Florida: Eolis stn. 307, off Fowey Eight (~25°35'26"N, 80°05'48"W, 128 m): USNM 417624 [1 shell]; Eolis stn. 362, off Fowey Eight (~25°35'26"N, 80°05'48"W, 174 m): USNM 417511* [3 shells]; Eolis stn. 370, off Ajax Reef (~25°24'00"N, 80°08'00"W, 128- 165 m): USNM 417497 [3 shells]; Eolis stn. 376, off Cae¬ sars Creek (~25°23'02"N, 80°12'37"W, 165 m): USNM 433081* [7 dd]; off Alligator Reef Eight, Eower Florida Keys (183 m), coll. 21/iv/1967: ANSP 312431* [1 shell]; zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 403-415 405 Figure 1. Eulimacrostoma microsculpturata gen. nov. and sp. nov. A. Holotype (USNM 429762); B, C. Paratype (USNM 433081); D-G. Paratype (FLMNH uncatalogued.) A, B, D. Shell in frontal view; C, E. Shell in lateral view; F. Detail of apical whorls in frontal view; G. Specimen attached to the host Luidia ludwigi scotti Bell, 1917 (Echinoderm collection, FLMNH 5042), red ellipse indicates the eulimid. Scale bars; 1 mm (A-E); 100 pm (F). Eolis stn. 300, off Sand Key (~24°27'16"N, 81°52'28"W, 132 m): USNM 433039 [1 shell]; Florida Straits (24°30'48.60"N, 83°30'1.44"W, 280 m), attached around the mouth of Luidia ludwigi scotti Bell, 1917 (FLMNH 5042, Echinodermata collection), coll. F. Michonneau, 13/iii/2007: FLMNH* (uncatalogued) [1 specimen]. Etymology. The epithet alludes to the microsculpture characteristic of the present species. Type locality. Puerto Rico: ofF San Juan, Johnson-Smithso- nian Deep-sea expedition stn. 10 (18°29'20"N-18°30'24"N, 66°05'30"W-66°04T5"W, 219-293 m), coll. Yacht Caro¬ line, 02/ii/1933. Diagnosis. Eulimid parasitic on starfish, with an elongat¬ ed shell, presenting microsculpture of axial lines, dome¬ shaped apex, a narrow brownish spiral band close to the suture in the protoconch and a wide brownish spiral band along the teleoconch, a high, wide and spread aperture. Umbilicus absent. Description. Shell conical with an obtuse apex, reaching about 7.3 mm long and 2.0 mm wide. Protoconch vit- zse.pensoft.net 406 de Souza, L.S. & Pimenta, A.D.: Eulimacrostoma gen. nov., a new genus of Eulimidae... Figure 2. Eulimacrostoma microsculpturata gen. nov. and sp. nov. A-E. Paratype (ANSP 312431); F. Paratype (USNM 433081); G. Paratype (USNM 417511). A, C. whole shell in ventral view; B. Shell in lateral view; D. Detail of body whorl in frontal view; E. Detail of teleoconch surface, white arrows indicates the micro sculpture of axial lines; F. Detail of apical whorls in frontal view; G. Detail of protoconch in apical view, white arrow indicates the transition protoconch-teleoconch. Scale bars; 1 mm (A-C); 500 pm D); 100 pm (E, G); 200 pm (F). zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 403-415 407 Table 1. Measurements (in mm, except for number of whorls) of Eulimacrostoma species studied. Taxon Catalog number Status Wh SL BWL AL SW AW Eu. microsculpturata USNM 429762 Holotype 8 3.16 1.60 1.06 1.08 0.60 ANSP 312431 Paratype 12 7.28 3.34 2.21 1.97 1.17 USNM 433081 Paratype 11 6.50 3.09 2.10 1.88 1.10 USNM 1273894 Paratype 11 6.55 3.04 1.99 1.79 1.15 USNM 1273894 Paratype 10.5 6.32 2.88 1.90 1.80 1.06 USNM 433081 Paratype 10 6.03 2.94 1.97 1.75 1.03 USNM 433081 Paratype 10 6.10 2.94 1.97 1.75 1.03 Eu. chascanon NHMUK 1887.2.9.1587 Lectotype 13 4.49 1.96 1.23 1.15 0.65 Eu. fusus USNM 87273 Lectotype 12 11.24 5.28 3.55 2.78 1.55 MCZ 7515 Para lectotype 13 12.29 5.63 3.81 2.75 1.62 Eu. lutescens MNHN-IM 2000-5662 Holotype 10 8.88 4.47 3.05 2.90 1.60 MZSP 34514 Paratype lot 10.62 5.00 3.22 3.15 1.82 MNHN-IM 9 7.58 3.75 2.69 2.28 1.41 Eu. patula USNM 160202 Lectotype 9 4.53 2.20 1.50 1.50 0.91 MNRJ 34479 8.5 3.99 2.07 1.42 1.50 0.89 MNRJ 34476 9 4.24 2.07 1.37 1.42 0.83 t Apex broken. Abbreviations: AL; aperture length; AW: aperture width; BWL: body whorl length; SL: shell length; SW; shell width.; Wh; whorls reous, with a brownish spiral band close to the suture, subcylindrical, about 2.5 whorls, 300 pm wide, 400 pm in height, smooth, transition to teleoconch marked by a subtle incremental scar and the end of the brownish spi¬ ral band. Teleoconch with about nine whorls of sinuous outline, convexity more attenuated on the lower region of each whorl; suture deep, well impressed, sloping; sub- sutural zone occupying about 1/5 of the whorl height; surface glossy, presenting several incremental scars and micro sculpture of opisthocline axial lines; incremental scars well developed, appearing in irregular intervals; micro sculpture of axial lines present at fairly regular in¬ tervals between 80-110 pm. Last whorl occupying about 45% of the shell length; base rounded in outline, elon¬ gated. Aperture high, occupying about 70% of the body whorl length, wide, pear-shaped, expanded laterally, acute posteriorly, rounded and spread anteriorly; outer lip thin, very sinuous, orthocline, retracted near the suture, after strongly protruding, and retracted in the distal re¬ gion, maximum projection at the middle of the outer lip height; inner lip sinuous, sloping and well demarcated. Umbilicus absent. Teleoconch whitish or vitreous, fresh specimens usually have the region near the suture unc¬ oloured and the rest of the whorl yellowish to brownish. Measurements. Holotype USNM 429762: whorls = 8; SL = 3.16 mm; BWL = 1.60 mm; AL = 1.06 mm; SW = 1.08 mm; AW = 0.60 mm. Paratype USNM 433081: Whorls = 11; SL = 6.50 mm; BWL = 3.09 mm; AL = 2.10 mm; SW= 1.88 mm; AW =1.10 mm. Geographic distribution. USA: Florida; Puerto Rico. Bathymetric distribution. From 128 m to 293 m. Remarks. Dali (1927: 67) cited 20 specimens of “Mc/- anellapatuld' from “off Georgia”. The USNM houses two lots (USNM 108031, USNM 108380, respectively from Femandina, Florida, USFC stn. 2668, and from Georgia, USFC stn. 2415), that can be attributed to the material stud¬ ied by Dali (1927) due to the labels. USNM 108031 con¬ tains a single shell with a broken protoconch and an eroded surface. Although the shell is not in perfect condition, it is possible to aflhrm that it is actually Eulimacrostoma mi- crosculpturata sp. nov. by the general shape and teleoconch sculpture. USNM 108380 contains seven shells, most of them immature, which can neither be identified with cer¬ tainty as Eulimacrostoma patula (Dali & Simpson, 1901), comb. nov. nor as Eulimacrostoma microsculpturata. Most specimens of Eu. microsculpturata have a straight spire (Figs lA, 2A), but individuals of the lot USNM 433081 (Fig. IB) have a slightly curved spire. The curvature is a growth phenomenon related with the position of the incremental scars and usually helps to dis¬ tinguish species (Bouchet and Waren 1986: 312). How¬ ever, in the case of Eu. microsculpturata the protoconch, color pattern, shape of the aperture, and dimensions are very similar in all individuals and we consider all of them belonging to the same taxon. The holotype USNM 429762 and paratype FLMNH are young individuals and shows a more rhomboid aperture and angulated body whorl, which is a common feature at this stage of growth (Bouchet and Waren 1986: 310; Souza et al. 2018: 926). Eulimacrostoma chascanon (Watson, 1883), comb, nov. (Fig. 3A-C) differs by the colorless shell, an aperture not so gaping anteriorly and by the proportionally small¬ er dimensions (Table 1). The lectotype of E. chascanon (Fig. 3A, B) is not fresh and the color pattern may have disappeared over time despite the good preservation of the shell surface. Eulimacrostoma fusus (Dali, 1889), comb. nov. (Fig. 3D-I) differs by being proportionally greater in shell size: the lectotype with about 12 whorls is 11.24 mm long, whereas the holotype of Eu. microsculpturata with about 11 whorls is 6.50 mm long. Eulimacrostoma fusus has no color pattern and the protoconch is wider than in Eu. microsculpturata. Eulimacrostoma lutescens (Simone, 2002), comb. nov. (Fig. 3J-N) differs by the fiatter teleoconch whorls and by the relatively wider and longer shell (holotype MNHN- zse.pensoft.net 408 de Souza, L.S. & Pimenta, A.D.: Eulimacrostoma gen. nov., a new genus of Eulimidae... IM 2000-5662, 10 whorls, 8.88 mm long, 2.90 mm wide vs paratype USNM 433081 of Eu. microsculpturata, 11 whorls, 6.50 mm long). Eulimacrostoma patula (Fig. 4A-J) has a more trun¬ cated base, like some specimens of Eu. microsculpturata (Fig. lA-E), but differs by the presence of an umbilical fissure and by the slightly wider spire angle. Eulimacrostoma chascanon (Watson, 1883), comb. nov. Figure 3A-C Eulima chascanon’Watson 1883: 114-115. E. chascanon: Watson 1886: pi. 35, fig. 4. Type material. Lectotype (herein designated) NHMUK 1887.2.9.1587. Paralectotype: from type locality: NHMUK Norman Coll. 1979225 [1 shell]. Material examined. Type material. Type locality. Puerto Rico: North of Culebra Island, off St. Thomas, Challenger Expedition stn. 24 (18°38'30"N, 65°05'30"W, 713 m). Redescription. Shell whitish, conical with an obtuse apex, reaching about 4.5 mm long and 1.2 mm wide, about 13 whorls. Protoconch vitreous, subcylindrical. Shell with about 13 whorls of slightly sinuous outline, with convex¬ ity more attenuated on the lower region of each whorl; suture slightly impressed, sloping; subsutural zone not visible; surface glossy, showing axial lines and incremen¬ tal scars; incremental scars slightly impressed, appearing in irregular intervals. East whorl occupying about 45% of the shell length; base rounded, elongated. Aperture high, occupying about 60% of the body whorl length, wide, pear-shaped, expanded laterally, acute posteriorly, round¬ ed and spread anteriorly; outer lip thin, very sinuous, or- thocline, strongly retracted near the suture, after strongly project, and retracted in the distal region, maximum pro¬ jection below the middle of the outer lip height; inner lip sinuous, sloping and well demarcated. Umbilicus absent. Measurements. Eectotype NHMUK 1887.2.9.1587: whorls= 13; SE = 4.49 mm; BWE = 1.96 mm; AE = 1.23 mm; SW= 1.15 mm; AW = 0.65 mm. Geographic distribution. Known only from the type locality. Bathymetric distribution. Known only from 713 m. Remarks. Eulimacrostoma chascanon comb. nov. (Fig. 3A-C) presents some similarities to the new genus, such as the elongated shell with moderately convex whorls and the wide aperture with an orthocline outer lip, strongly protruding. The shell surface is very polished, but some axial lines can be observed in the types. An analysis under SEM would confirm whether these impressions are like the kind of sculpture present in Eu. microsculpturata. The type material of Eu. chascanon formerly consist¬ ed of two syntypes (Fig. 3A-C) housed in the NHMUK collection. The shell of NHMUK 1887.2.9.1587 is the best preserved one and seems to be the shell figured by Watson (1886: pi. 35, fig. 4), which is here selected as the lectotype (Fig. 3A, B). Eulimacrostoma fusus (Fig. 3D-I) differs main¬ ly by the proportionally longer shell (lectotype USNM 87273, 12 whorls, 1E24 mm long vs lectotype NHMUK 1887.2.9.1587 of Eu. chascanon, 13 whorls, 4.49 mm long) and wider apex. Eulimacrostoma lutescens (Fig. 3J-N) differs by the much wider protoconch and by the wider and longer tel- eoconch (holotype MNHN-IM 2000-5662, 10 whorls, 8.88 mm long, 2.90 mm wide vs lectotype NHMUK 1887.2.9.1587,13 whorls, 4.49 mm long, 1.15 mm wide). Eulimacrostoma fusus (Dali, 1889), comb. nov. Figure 3D-I Eulima fusus Dali 1889: 329, pi. 19, fig. IIB. Strombiformis fusus: Abbott 1974: 127, fig. 1392 [reproduced from original illustration]. Type material. Eectotype (herein designated) USNM 87273. Paralectotype: Cuba: off Morro Eight, “Blake” stn. 100 (457-732 m), coll. RIV Blake, xii/1878: MCZ 7515. Material examined. Type material. Type locality. Yucatan Strait (1170 m). Redescription. Shell whitish, conical with an obtuse apex, reaching about 12.4 mm long and 2.8 mm wide, about 12 whorls, apical whorls slightly bent, aperture broad. Proto¬ conch whitish, subcylindrical. Shell with about 12 whorls of slightly sinuous outline, with convexity more attenu¬ ated on the lower region of each whorl; suture well im¬ pressed, sloping; subsutural zone occupying about 1/5 of the whorl height; surface glossy, showing microsculpture of axial lines and incremental scars; incremental scars well impressed, appearing in irregular intervals. East whorl occupying about 45% of the shell length; base rounded, slightly truncated. Aperture high, occupying about 65% of the body whorl length, wide, slightly acute and spread an¬ teriorly and acute posteriorly; outer lip thin, very sinuous, orthocline, strongly retracted near the suture, after strong¬ ly protruding, and retracted in the distal region; inner lip almost straight, well demarcated. Umbilicus absent. Measurements. Eectotype USNM 87273: whorls = 12; SE = 11.24 mm; BWE = 5.28 mm; AE = 3.55 mm; SW = 2.78 mm; AW = 1.55 mm. Geographic distribution. Yucatan Strait and off Cuba. zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 403-415 409 Figure 3. Eulimacrostoma spp. A-C. EuUmacrostoma chascanon (Watson, 1883), comb, nov.; A, B. Lectotype herein desig¬ nated (NHMUK 1887.2.9.1587); C. Paralectotype (NHMUK Norman Coll. 1979225); D-L Eulimacrostoma fusus (Dali, 1889), comb, nov.: D, E, G-L Lectotype herein designated (USNM 87273); G. Paralectotype (MCZ 7515). J-N. Eulimacrostoma lutescens (Simone, 2002), comb, nov.; J, K. Holotype (MNHN-IM 2000-5662); L-N. (MNHN-IM uncatalogued), from type locality. A, C-E, G, K, M. Shell in frontal view; B. Shell in dorsal view; F, L, N. Shell in lateral view; H, O. Detail of apical whorls in frontal view; I, J. Detail of teleoconch surface, white square in I indicates detail in J, white arrows in J indicates microsculpture of axial lines. Scale bars: 1 mm (A-G, I, K-N); 400 pm (H); 50 pm (J); 200 pm (O). Credits: A-C: NHMUK; H, I; USNM; J, K; MNHN-IM. zse.pensoft.net 410 de Souza, L.S. & Pimenta, A.D.: Eulimacrostoma gen. nov., a new genus of Eulimidae... Bathymetric distribution. From 457 to 1170 m (Dali 1889). Remarks. The types of Eulimacrostoma fusus comb. nov. (Fig. 3D-I) have a partially broken aperture, resembling a channeled aperture (Dali 1889). This condition does not allow for the aperture to be perfectly described, but it is very elongated. The shell is conical, very elongated, with moderately convex and slightly distorted whorls. This species has well-demarcated axial lines in the teleoconch surface (Fig. 3H, I) and a broad subcylindrical protoconch (Figure 3G), which resembles Eu. microsculpturata. Dali (1889) referred to two specimens from different localities, and thus these shells are originally syntypes. Here, USNM 87273 is selected as the lectotype (Fig. 3D, E, G-I). Dali (1927: 68) recorded 18 shells off Fernandina, Florida, USA, without illustrations. The material cited by Dali was not found in the USNM and is considered du¬ bious. There is no other additional record of the species. Ode (1989: 67) recorded ''Polygireulima fusus (Dali, 1889)” without illustrations from Texas, USA, at a depth of 110 m. This record was based on a single specimen with 2.9 mm long, a very small length for the species. We could not find the voucher of Ode’s record. Because there are no illustrations and the length reported by him is small in comparison to the types, we consider this a dubious record. Eulimacrostoma lutescens (Fig. 3J-N) differs by the wider spire angle of the teleoconch in comparison to the slowly increasing diameter of the teleoconch in Eu. fusus. Eulimacrostoma lutescens (Simone, 2002), comb. nov. Figure 3J-N Batheulima lutescens Simone 2002: 56, figs 5-8. B. lutescens: Rios 2009: 194 (text figure reproduced from original illus¬ tration); Dornellas and Simone 2011: 25. Type material. Holotype MNHN-IM 2000-5662. Para- types: Brazil: Espirito Santo state: MD55 stn. 42 CB76 (18°58'59"S, 37°49'59'’W, 637 m), coll. RW Marion Dufresne, 27/v/1987: MNHN-IM 2000-5664 [1 shell]; Rio de Janeiro state: MD55 stn. 64 CB105 (23°46'59"S, 42°10'00"W, 610 m), coll. RTV Marion Dufresne, 02/ vi/1987: MNHN-IM 2000-5663 [2 shells]; Sao Pau¬ lo state: Off Sao Sebastiao Island (23°47'S, 42°10'W, 610 m): MZSP 34514 [1 shell], MZSP 34515 [1 shell]. Material examined. Type material. Brazil: from type lo¬ cality: MNHN-IM* (uncatalogued) [2 shells]. Type locality. Brazil: Espirito Santo: MD55 stn. 54 CB93 (19°36'00"S, 38°53T8"W, 640 m), coll. R/V Marion Du- fresne, 30/v/1987. Measurements. Holotype MNHN-IM 2000-5662: whorls = 10; SE = 8.88 mm; BWE = 4.47 mm; AE = 3.05 mm; SW = 2.90 mm; AW = 1.60 mm. Geographic distribution. Brazil: Espirito Santo, Rio de Janeiro, Sao Paulo (Simone 2002). Bathymetric distribution. From 610 to 640 m. Remarks. Simone (2002) originally included this spe¬ cies in Batheulima Nordsieck, 1968, but the species do not present the typical dark-brown protoconch and the sigmoidal axial lines in the protoconch like Bateulima fuscoapicata (Jeffreys, 1884), type species of the genus. Eulimacrostoma lutescens comb. nov. fits the shape of the genus erected here by the very elongated and slender shell, with an elongated, anteriorly spread aperture, and a similar color pattern. The protoconch and teleoconch of the latter species has a weak brownish spiral band (Fig. 3E, N). Additionally, the shell of Eu. lutescens has several axial lines (Simone 2002). In addition to the type series of Eu. lutescens, we find two other shells collected from the type locality. One of them is figured here (Fig. 3E-N), and it is slightly smaller than the holotype but has one less whorl (Table 1). The initial whorls of the holotype (Fig. 3J) are straighter than in the topotype (Fig. 3E, M), but it may be variable. The topotype has a similar conical shape, with a spire angle of 21° (the holotype has 23°), presence of axial lines very evident, faint brownish spiral bands in the protoconch (Fig. 3N) and teleoconch (Fig. 3E, M), a similar aperture (Fig. 3E) and a strongly protruding outer lip (Fig. 3M). Eulimacrostoma patula (Dali & Simpson, 1901), comb. nov. Figure 4A-K Eulima (Leiostraca) patula Dali and Simpson 1901: 413, pi. 57, fig. 3. Strombiformis patula: Abbott 1974: 127, fig. 1390 (reproduced from original illustration). E. patula: Lamy and Pointier 2017: 277, pi. 88, fig. 3A, B. Type material. Eectotype USNM 160202a (herein des¬ ignated). Paralectotypes USNM 160202b [4 shells], from type locality. Material examined. Type material. Brazil: Amapa state: mouth of Amazonas River, outer shelf (~03°58'42"N, 49°33'24"W), coll. 30/vii/2001: MNRJ 34476* [4 shells], MNRJ 34477* [2 shells], MNRJ 34478* [3 shells], MNRJ 34479* [8 shells]; AMASSEDS stn. 4134 (02°21T2"N, 48°29'54"W, 72 m^ coll. R/V ColumbusIselin, 05/xi/1990: MNRJ 34589* [2 shells], MNRJ 35284* [1 shell]. Type locality. Puerto Rico: Mayaguez Harbour, Fish Hawk stn. 6062 (46-55 m). Redescription. Shell vitreous or with wide brownish spi¬ ral bands, conical with an obtuse apex, reaching about 4.5 mm long and 1.5 mm wide, about eight whorls, ap¬ erture broad. Protoconch vitreous, subcylindrical, about two whorls, 330 pm in width, smooth, transition to teleo- zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 403-415 411 Figure 4. Eulimacrostomapatula (Dali & Simpson, 1901), comb. nov. A. Lectotype herein designated (USNM 160202); B, F. (MNRJ 34479); C. (MNRJ 34476). D-E, G-K. (MNRJ 34589). A-D. Shell in frontal view; E. Shell in lateral view; F. Detail of teleoconch surface, white arrows indicates microsculpture of axial lines; G. Detail of body whorl in ventral view; H. detail of body whorl in lat¬ eral view; 1. Detail of umbilical fissure; J. Detail of apical whorls in frontal view; K. Detail of protoconch in apical view, white arrow indicates the transition from protoconch to teleoconch. Scale bars: 1 mm (A-E); 200 pm (F-H); 100 pm (I-K). Credits: A: USNM. zse.pensoft.net 412 de Souza, L.S. & Pimenta, A.D.: Eulimacrostoma gen. nov., a new genus of Eulimidae... conch marked by a subtle incremental scar. Teleoconch with about six whorls of sinuous outline, with convexity more attenuated on the lower region of each whorl; suture deep, well impressed, sloping; subsutural zone occupying about 1/5 of the whorl height; surface glossy and smooth, except for microsculpture of axial lines and incremental scars; incremental scars well developed, appearing in in¬ tervals of about 0.5 whorl. Last whorl occupying about 50% of the shell length; base slightly abrupt in outline. Aperture high, occupying about 70% of the body whorl length, wide, pear-shaped, expanded laterally, acute pos¬ teriorly, rounded and spread anteriorly; outer lip thin, very sinuous, orthocline, strongly retracted near the suture, af¬ ter strongly protruding, and retracted in the distal region, maximum projection at the middle of the outer lip height; inner lip sinuous, sloping and well demarcated. Umbilical fissure present. Shell whitish or vitreous, usually with a wide brownish spiral band along the teleoconch. Measurements. Lectotype USNM 160202: whorls = 9; SL= 4.53 mm; BWL = 2.20 mm; AL = 1.50 mm; SW =1.50 mm; AW = 0.91 mm. Geographic distribution. Puerto Rico (Dali and Simp¬ son 1901); Guadeloupe (Lamy and Pointier 2017); Bra¬ zil: Amapa (this study). Bathymetric distribution. From 46 m (Dali and Simp¬ son 1901) to 300 m (Lamy and Pointier 2017). Remarks. Eulimacrostoma patula comb. nov. (Fig. 4A- K) has a combination of shell features that fit the morphol¬ ogy of the newly described genus, such as the elongated aperture spread anteriorly, a orthocline outer lip strongly protruding (Fig. 4G, H) and moderately convex teleoconch whorls (Fig. 4A-E). Additionally, this species usually has a weak brownish spiral band in the teleoconch (Fig. 4C). Dali and Simpson (1901) referred to five specimens in the original description without the selection of a holo- type and, thus, all shells are syntypes. We select the shell of USNM 160202 (Fig. 4A) as the lectotype. The material from the north coast of Brazil fits perfect¬ ly with the lectotype of Eu. patula in shape and reaches a similar size (lectotype USNM 160202, 9 whorls, 4.53 mm long vs MNRJ 34479, 8 whorls, 3.99 mm long). The record of Eu. patula from the north coast of Brazil represents the southernmost record of this species and is more than 2,500 km from the type locality and about 1,800 km from the recent record from Guadeloupe (Lamy and Pointier 2017). This species is currently known only by empty shells. As commented above, one shell recorded by Dali (1927: 67) is actually Eu. microsculpturata and the re¬ maining shells found cannot be identified with certainty due to their poor condition. We did not find more material to reassess the record of Eu. patula by Dali (1927). Parker and Curray (1956) recorded Eu. patula (as ''Melanella patula'') from calcareous banks off Texas and Louisiana, USA, in a depth of about 55 m, in a checklist of species without illustrations. Ode (1989) recorded Eu. pat¬ ula (as 'Eabinella patula") from Texas also with no illus¬ trations. The source of material of these records could not be traced in malacological collections and the presence of this species in the Gulf of Mexico is considered dubious. Eulimacrostoma patula differ from the congeners mainly by the presence of an umbilical fissure and by the more truncated base. Discussion Eulimacrostoma gen. nov. is erected to group the eulimids with an elongated, conical shell, possessing a smooth pro¬ toconch, a teleoconch with slightly convex whorls and an enlongate and strongly anteriorly spread aperture. The spe¬ cies usually share a similar color pattern of brownish spiral bands in the protoconch and teleoconch, and microsculpture of axial lines in teleoconch. These latter features are more developed in the type species and in Eu. fusus and Eu. lute- scens. The anatomy of Eu. microsculpturata is not described, as only one young specimen with soft parts was identified. However, the kind of host (a starfish) and the combination of shell features corroborate the distinction of this genus. Among the eulimid genera that parasitize asteroids, the most similar to Eulimacrostoma is Niso Risso, 1826, due to its conical shape, presence of brownish spiral bands in several species and axial lines on the shell surface. Howev¬ er, Eulimacrostoma can be distinguished from Niso by the shape of the aperture, which gapes anteriorly and is more elongated in the former, whereas it is usually rhomboid in the latter. In addition, Niso usually has a well-developed umbili¬ cus. The type species of Eulimacrostoma does not present an umbilicus, hui Eu. patula has an umbilical fissure, which is a variable feature in relation to its presence or absence in some Eulimidae genera (e.g., EulimettaWarm, 1992, Fusceulima Easeron, 1955) (Souza and Pimenta 2014,2015). Other similar genera, in relation to conchological fea¬ tures, that parasitize other classes of echinoderms or of unknown hosts are Batheulima, Eulima, and Haliella Monterosato, 1878. Eulimacrostoma is similar to Batheulima, whose host is unknown (Bouchet and Waren 1986); they have an elongated aperture, with a strongly protruding outer lip and sigmoidal axial lines in the teleoconch. However, Eulimacrostoma can be distinguished by features of the protoconch, which is smooth, not completely colored. Eulimacrostoma is also similar to Eulima, but the more typical forms of this species-rich genus that para¬ sitizes ophiuroids have a narrower aperture, that is not so spread anteriorly, and the outline of teleoconch whorls of Eulima is usually fiat. zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 403-415 413 50-'N 40'^N JO^N 20®N |[|“N QO lO^S m\\ 80AV l&W COW 50-\V 40'W i(f\\ .10‘N 20"N I0"N 0^ iirs loonv 00" W 80^VV 70nv 60°\V 50=W 40= W 30= W 8{l=W 70=-%' ftO=W 40= W 30’W 20=W lO'W 50=N 4[1=N 30=N 2t)“N lO^N 0 “ I0=S 100°W 00°W' 80=W 70°W 60°\V 50=W' 40°W 30°W Figure 5. Geographical distributions of Eulimacrostoma spp. based on confirmed identifications. A. EuUmacrostoma microsculp- turata sp. nov.; B. Eulimacrostoma chascanon (Watson, 1883), comb, nov.; C. Eulimacrostoma fusus (Dali, 1889), comb, nov.; D. Eulimacrostoma lutescens (Simone, 2002), comb, nov.; E. Eulimacrostomapatula (Dali & Simpson, 1901), comb. nov. White circle: type locality; black circle: additional records. Records without geographic coordinates are approximate. ATLANTIC OCEAN V N c 10 °.s- ATLANTIC OCEAN ATLANTIC OCEAN A 5Cr'N- 40 N- .Ui'-N- ••• 2 crN- B ATLANTIC OCEAN —I-1-1-i-1-1-1-1— 100=Vv' 90=W 80=W 70 AV 60=W 50=\V 40nV 30'=W D ATLANTIC OCEAN • T-1-1-1-1-1-r zse.pensoft.net 414 de Souza, L.S. & Pimenta, A.D.: Eulimacrostoma gen. nov., a new genus of Eulimidae... Eulimacrostoma can also be compared to Haliella Monterosato, 1878 by possessing a high aperture, but the latter can be distinguished by having a more cylindrical shell, a more obtuse apex, and by being colorless. The host of Haliella species is doubtful, Waren (1983) com¬ mented that the genus possibly includes parasites of ophi- uroids (Waren 1983: 47), while Hasegawa (2009: 276) noted irregular sea urchins. Hasegawa (2009) identified Haliella sp. collected in the same haul of several irregular sea urchins and ophiuroids. Despite these hypotheses, the species of Haliella have never been found on the host. Delimitation of genera is a hard task in Eulimidae (Bouchet and Waren 1986: 310), with some exceptions. Several species and genera are known only from empty shells and/or have no data about the host and/or anatomy (e.g., Batheulima, Eulimetta, Oceanida de Folin, 1870) (Waren 1983, 1992; Bouchet and Waren 1986; Souza and Pimenta 2014, 2015). In a revision of the family in the northeast Atlantic, Bouchet and Waren (1986) established eight new genera (including Acrochalix, Bulimeulima, Campylorhaphion, and Halielloides) known only from shell morphology and argued that they can introduce 10 more genera based exclu¬ sively on shell morphology. The introduction of these gen¬ era was justified by consistent features of the shell and the author’s experience on the eulimid fauna of other regions. The description of Eulimacrostoma is based on the combination of shell features and on data about the host of Eu. microsculpturata. Albeit only one species has the host known, all species included in Eulimacrostoma have very similar conchological features. Recent efforts with molecular data are elucidating many aspects of the evolution of Eulimidae (Takano and Kano 2014; Takano et al. 2018), but the family is too species-rich and to reach a high percentage of DNA-se- quenced taxa is a challenge. Takano et al. (2018) pro¬ posed a new methodology to identify the hosts of eulim- ids by extracting DNA from the proboscis of the eulimid in an attempt to find DNA traces of the host and reached successful results. This is an interesting technique, espe¬ cially if the record of the eulimid attached to the host is not available. We expect that the description of Eulimacrostoma calls the attention of researchers to congeners from other parts of the world. By now, Eulimacrostoma is known only from the western Atlantic (Fig. 5A-E). Several genera of Eulimidae have a worldwide distribution (e.g., Eulima, Fusceulima, Melanella Bowdich, 1822). The only known host of Eulimacrostoma is a starfish of the genus Luidia Forbes, 1839, which has a worldwide distribution (Mah 2019), occurring mainly in shallow waters of tropical and subtropical seas and inhabiting soft bottoms (Clark and McKnight 2000). The current state of knowledge is not sufficient to establish a specificity in the parasite-host re¬ lationship at the species or genus level. Thus, more data is necessary to reach a better comprehension about the biology of Eulimacrostoma. Acknowledgements We are grateful to the curators/collection managers for the loans, photographs and support during research: Gary Rosenberg, Nasreen Phillips and Paul Callomon (ANSP), Mandy Bemis and John Slapcinsky (FEMNH), Adam Baldinger (MCZ), Fuiz Simone (MZSP), and El¬ len Strong and Yolanda Villacampa (USNM). The first author is much indebted to his friends Carine Galvao and Gustavo Miranda for assistance during a research trip to the collections in the USA. Thanks to Nataly Slivak and Carlos Renato R. Ventura (MNRJ) for calling our atten¬ tion to the eulimid attached to the starfish. Beatriz Olivei¬ ra and Camila Messias (MNRJ) for having conducted the SEM micrographs. Mauricio Fernandes (UNIRIO) who reviewed and improved an early version of this manu¬ script. Also, we wish to thanks Dr Jose H. Eeal (BMSM), an anonymous reviewer. Dr Matthias Glaubrecht (Ed¬ itor) and Robert Forsyth for a detailed revision of the manuscript. This work was supported by “Funda^ao Carlos Chagas Filho de Amparo a Pesquisa do Esta- do do Rio de Janeiro” (FAPERJ) under grant numbers E-26/110.325/2014 and E-26/110.068/2014. The first author is grateful to “Conselho Nacional de Desenvolvi- mento Cientifico e Tecnologico” (CNPq) and to “Coorde- na^ao de Aperfei^oamento de Pessoal de Nivel Superior” (CAPES) for a scholarship, and to Conchological Society of Great Britain and Ireland for a research grant. References Abbott RT (1974) American seashells. The marine Mollusca of the At¬ lantic and Pacific coasts of North America. Van Nostrand Reinhold Company, New York, 663 pp. Bartsch P (1933) Station records of the first Johnson-Smithsonian deep- sea expedition. Smithsonian Miscellaneous Collections 91(1): 1-31. Bieler R, Mikkelsen PM (2002) The Cruises of the Eolis\ John B. Henderson’s mollusk collections off the Florida Keys, 1910-1916. American Malacological Bulletin 17(1/2): 125-140. Bouchet P, Waren A (1986) Revision of the northeast Atlantic bathyal and abyssal Aclididae, Eulimidae, Epitoniidae (Mollusca, Gastrop¬ oda). 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BERLIN Three new speeies of Macrophya Dahlbom (Hymenoptera, Tenthredinidae) with a key to species of the Macrophya imitator group in China Mengmeng Lm\ Zejian Meicai WeP 1 College of Ecology, Lishui University, Lishui, Zhejiang, China 2 Postdoctoral Work Station, Lishui Academy of Forestry, Lishui, Zhejiang, China 3 College of Life Science, Jiangxi Normal University, Nanchang, Jiangxi, China http://zoobank.org/7F0D0518-F845-43BB-924A-7B4AF0222126 Corresponding author: Meicai Wei (weimc@126.com); Zejian Li (Iizejian2006@163.com) AcdidQmic Qdiiior. Martin Husemann ♦ Received 20 April 2019 ♦ Accepted 10 July 2019 ♦ Published 25 July 2019 Abstract The Macrophya imitator group was proposed by Liu et al. in 2015. Three new species, Macrophya longlingensis sp. nov., M niesh- uaiguoi sp. nov. and M zejiani sp. nov. from China are described. A key to all Chinese species and a geographical distribution map of the M. imitator group in China are provided. Key Words Hymenoptera, Macrophya imitator group, Sawflies, taxonomy, Tenthredinoidea Introduction Macrophya Dahlbom, 1835 is the third largest genus in the subfamily Tenthredininae (Hymenoptera, Tenthredinidae). It contains 306 species worldwide, of which 167 have been recorded for China up to April 2019 (Li et al. 2019a, 2019b; Liu et al. 2019a, 2019b). The taxonomy and distribution of the genus in China has been studied by the last author and his co-workers since 1994 and a division of Macrophya in species groups was given by Liu et al. (2015,2018,2019b). The Macrophya imitator group is the second largest species group in Macrophya, with 17 species worldwide, all of which are present in China. Among them, M. imita¬ tor Takeuchi is also distributed in Japan, Korea and Rus¬ sia (Takeuchi 1937), and M. postscutellaris Malaise in Myanmar (Malaise 1945). The species of the Macrophya imitator group are all similar in general morphology and constitute a clearly defined species group in Macrophya. In this study, three new species belonging to this species group are described from China, namely: M. longlingen¬ sis Li, Liu & Wei, sp. nov., M. nieshuaiguoi Li, Liu & Wei, sp. nov. and M. zejiani Liu & Wei, sp. nov. A key to all species found in China is provided. Materials and methods All specimens of the newly described species were ob¬ tained by sweeping in wooded bog and forest fringe zones in Yunnan Province (southern China) from 1994 to present. Eight specimens of three new species and 788 specimens of known species were examined and studied Copyright Mengmeng Liu etal. This is an open access articie distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the originai author and source are credited. 418 Mengmeng Liu et al.: Three new species of Macrophya imitator group for this work. The specimens were examined with a Mot- ic-SMZ-171 stereomicroscope. Images of the imagines were taken with a Nikon D700 digital camera and a Lei- ca Z16APO. The genitalia were examined with a Motic BA410E microscope and photographed with Motic Moti- cam Pro 285A. Images were focus-stacked using Helicon Focus (HeliconSoft, Kharkiv, Ukraine) and further pro¬ cessed with Adobe Photoshop CS 11.0. The terminology of genitalia follows Ross (1945) and that of general morphology follows Viitasaari (2002). For a few terms (e.g. middle fovea and lateral fovea), we fol¬ low Takeuchi (1952). All types are deposited in the Insect Collection of Cen¬ tral South University of Forestry and Technology, Chang¬ sha, Hunan, China (CSCS). Abbreviations: OCL The distance between a lateral ocellus and the occipital Carina, or the hind margin of the head where this carina would be if it were developed (Benson 1954). OOL The shortest distance between an eye and a later¬ al ocellus. POL The distance between the mesal margins of the two lateral ocelli. Results Macrophya imitator species group Remarks. The Macrophya imitator group is morpho¬ logically very similar to the M. maculitibia group, a diagnosis to which was provided by Li et al. (2019b). Species of the M. imitator group can be recognized us¬ ing the diagnosis provided by Liu et al. (2015), here restated: the body mainly black, without metallic tinge; the antenna slender and black; the posterior margin of metepimeron straight or slightly concave, the append¬ age (= posterior corner of metepimeron) differentiated but not elongated, at least partly punctate and even¬ ly pilose, without basin; the abdominal tergum 1 not reticulate and the penis valve oval, narrowed toward apex, ergot short. Description. Body slender and mainly black, with¬ out metallic tinge; white maculae to varying extents on pronotum, hind trochanter and dorsal surface of hind tibia subapically; fore wing without smoky macula be¬ low pterostigma; clypeus at its greatest breadth slightly broader than the shortest distance between lower inner orbits of eyes; lateral margins convergent anteriorly, an¬ terior margin incised to approximately 1/5-1/3 length of clypeus, apex of lateral lobe obtuse; malar space narrower than diameter of an ocellus; postocellar area broader than long; vertex with minute and dense punc¬ tures, interspaces between punctures narrow usually; antenna slender and black, antennomere 3 clearly longer than antennomere 4; posterior margin of metepimeron straight or slightly concave, appendage (posterior cor¬ ner of metepimeron) differentiated but not elongated, at least partly punctate and evenly pilose, without basin; inner spur of hind leg slightly longer than half length of metabasitarsus, metabasitarsus always slender, slightly longer than following four tarsomeres together; claw with inner tooth slightly shorter than outer tooth; ab¬ dominal tergum 1 not reticulate; penis valve oval, nar¬ rowed towards apex, ergot short. Key to the Chinese speeies of the Macrophya imitator group 1 Female.2 Male.21 2 Ovipositor sheath much longer than middle tibia.3 Ovipositor sheath clearly shorter than middle tibia.5 3 Posterior margin of pronotum with narrow white band; dorsal surface of hind tibia with a large white macula subapically. Ohina (Beijing, Gansu, Hebei, Henan, Hubei, Ningxia, Qinghai, Shaanxi, Shanxi, Sichuan). M. iven/Wei, 1998 Pronotum entirely black; dorsal surface of hind tibia with a subapical white macula smaller than above.4 4 Postocellar area 2.5x broader than long; POL: OOL: OOL = 4.5: 10: 5.5 (Figure 3B); antennomere 3 approximately 1.4x longer than antennomere 4 (13: 9) (Figure 3D); distance between cenchri twice breadth of a cenchrus; middle serrulae with 2 or 3 proximal and 9 or 10 distal teeth (Figure 3H); cell 2Rs as long as cell IR^, petiole of anal cell in hind wing 0.6x as long as cross-vein cu-a (Figure 3A). Ohina (Yunnan). M. zejiani Liu & Wei, sp. nov. Postocellar area twice broader than long; POL: OOL: OOL = 3: 10: 7; antennomere 3 approximately 1.8x longer than antennomere 4 (11: 6); distance between cenchri 2.5x breadth of a cenchrus; middle serrulae with 2 proximal and 9-12 distal teeth; cell 2Rs clearly shorter than cell IR^, petiole of anal cell in hind wing only slightly shorter than cross-vein cu-a. Ohina (Sichuan). M. omeialpina Li, Jiang & Wei, 2018 5 Apex of middle tibia with a distinct white macula on dorsal surface; punctures on middle part of mesepisternum minute, much smaller than punctures on vertex.6 Apex of middle tibia without white macula on dorsal surface, but sometimes with a white spot or stripe on anterior surface; punctures on middle part of mesepisternum about as large as or somewhat smaller than punctures on vertex.12 6 Hind trochanter entirely white.7 Hind trochanter partly white, with a distinct black macula.8 zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 417-427 419 7 8 9 10 11 12 13 14 15 16 17 18 19 Ovipositor sheath longer than fore tibia, with lateral setae very short, not distinctly curved; middle serrulae with 20 fine distal teeth. China (Henan, Hubei, Hunan, Shaanxi). M. flactoserrula Chen & Wei, 2002 Ovipositor sheath shorter than fore tibia, with lateral setae long and curved; middle serrulae with 10-12 distal teeth. China (Gansu, Henan, Hubei, Shaanxi). M. funiushana Wei, 1998 Pronotum entirely black.9 Posterior margin of pronotum white.10 Postocellar area 1.7x broader than long; fore and middle trochanters entirely black; hind trochanter entirely black; subapical white macula on dorsal surface of hind tibia about 2/5 length of tibia; posterior margin of metepimeronal appendage with a distinct shiny and obtuse carina toward the longitudinal axis of body; ovipositor sheath as long as fore tibia; the middle serrulae with 13-16 distal teeth. China (Jllln, Shaanxi). M. bui Wei & Li, 2012 Postocellar area twice broader than long; fore and middle trochanters largely black; hind trochanter largely white, ventral surface with black macula; subapical white macula on dorsal surface of hind tibia shorter than 1/3 length of tibia; the inner side of metepimeronal appendage without a shiny and obtuse carina; ovipositor sheath distinctly longer than fore tibia; lancet oblique and weakly protruding, with several larger teeth, the middle serrulae with 5-7 distal teeth. China (Gansu, Hebei, Henan, Jilin, Liaoning, Ningxia, Shaanxi, Shanxi). M. par/m/fa tor Wei, 1998 Hind tibia with broad white ring at mid-length, as long as half length of hind tibia. China (Shaanxi). . M. circulotibialis LI, Liu & Heng, 2015 Hind tibia with subapical white macula shorter than half length of hind tibia.11 Abdominal tergum 1 entirely black, posterior margin without white macula; middle serrulae each with 1 or 2 proximal and 14 or 15 distal teeth, subbasal teeth small. China (Jllln). M. changbaina LI, Liu & Heng, 2015 Posterior margin of abdominal tergum 1 with 2 small, distinct and white maculae; middle serrulae flat, middle serrulae each with 2 proximal and 15-18 distal teeth, subbasal teeth minute. China (Jilin, Ningxia). . M. curvatitheca Li, Liu & Heng, 2015 Punctures on head and mesepisternum clearly defined, equal In size. Interspaces strongly shiny; punctures on metepi¬ meronal appendage clearly separated; hind tibia with white macula as long as half length of hind tibia. Myanmar; China (Chongqing, Guizhou, Hubei, Shaanxi, Tibet). M. postscutellaris Malaise, 1945 Punctures on mesepisternum smaller than punctures on head, punctures crowded on both. Interspaces very fine, partly obscure, less shiny; punctures on metepimeronal appendage hardly separated; hind tibia with subapical white macula distinctly shorter than half length of hind tibia.13 Frons distinctly convex and extending above top of eyes; posterior 1/3 of abdominal tergum 1 with white bands across its full breadth. China (Sichuan). M. kangdingensis Wei & LI, 2012 Frons flat and not extending above top of eyes; posterior margin of abdominal tergum 1 with very narrow white band, or with 2 small transverse white maculae.14 Posterior margin of pronotum white.15 Pronotum entirely black.16 Setae on ovipositor sheath short and straight in dorsal view; posterior margin of metepimeronal appendage without glabrous patch; middle serrulae with 9 or 10 fine distal teeth; annular spine bands narrow and remaining distant from each other. China (Gansu, Guizhou, Hubei, Hunan, Shaanxi, Sichuan). M. imitatoides Wei, 2007 Setae on ovipositor sheath long and evenly curved In dorsal view; posterior margin of metepimeronal appendage with a distinct glabrous patch; middle serrulae with 5 or 6 fine distal teeth; annular spine bands broadly meeting each other. China (Gansu, Hubei, Ningxia, Shaanxi, Sichuan). M. curvatisaeta Wei & Li, 2011 Hind trochanter entirely black.17 Ventral surface of hind trochanter with black macula.18 Postocellar area 1.7x broader than long; subapex In dorsal surface with a clear white macula; middle serrulae with 2 proximal and 5 or 6 distal teeth. Korea, Japan, Russia (East Siberia); China (Heilongjiang, Jilin, Liaoning). . M. /m/totor Takeuchi, 1937 Postocellar area twice broader than long (Figure 2B); hind tibia with subapical white macula on dorsal surface weak (Figure 2A); middle serrulae with 2 proximal and 7-10 distal teeth (Figure 2H). China (Yunnan). . M. nieshuaiguoi Li, Liu & Wei, sp. nov. Ventral surface of hind trochanter with a large, distinct black macula.19 Ventral surface of hind trochanter with a small, weak black macula.20 Postocellar area 2.2x broader than long; white band at center of posterior margin of abdominal tergum 1 narrow; sub¬ apical white macula on dorsal surface of hind tibia indistinct; petiole of anal cell In fore wing slightly shorter than vein Ir-m, about half length of vein cu-a; middle serrulae of lancet each with 2 proximal and 7 or 8 distal teeth, cypsella between the 8*^-9*^ serrulae slightly broader than length of the 9*^ serrula. China (Sichuan). . M. semipunctata Li, Liu & Wei, 2018 Postocellar area 2.5x broader than long; white band at center of posterior margin of abdominal tergum 1 broader than the former; subapical white macula on dorsal surface of hind tibia distinct and small, but oblique; petiole of anal cell in fore wing twice length of vein Ir-m, and about as long as vein cu-a; middle serrulae each with 2 proximal and 5-7 zse.pensoft.net 420 Mengmeng Liu et al.: Three new species of Macrophya imitator group 20 21 22 23 24 25 26 27 28 29 30 31 32 distal teeth, cypsella between the 8*^-9*^ serrulae as broad as length of the 9*^ serrula. China (Gansu, Ningxia, Shaanxi, Sichuan). M. nigromaculata Wei & Li, 2010 Anterior margin of clypeus incised to approximately 1/3 its length (Figure 1C); middle serrulae with 2 proximal and 8-11 distal teeth (Figure IH); petiole of anal cell in hind wing as long as cross-vein cu-a (Figure lA). China (Yunnan)... . M. longlingensis Li, Liu & Wei, sp. nov. Anterior margin of clypeus incised to approximately 1/5 its length; middle serrulae with 2 proximal and 5-7 distal teeth; petiole of anal cell in hind wing 0.5x longer than cross-vein cu-a. China (Hubei, Jilin, Shaanxi). . M. jiaozhaoae Wei & Zhao, 2011 Hairs on abdominal terga erect, approximately as long as diameter of middle ocellus; anterior margin of valviceps somewhat acute. M. i/i/en/'Wei, 1998 Hairs on abdominal terga oblique, much shorter than diameter of middle ocellus; anterior margin of valviceps more or less evenly rounded.22 Hind tibia with a white macula extending over approximately half its length.23 Hind tibia with a white macula clearly shorter than half its length.24 Hind tibia with a broad, white ring about its mid-length; all trochanters mostly black; white macula on posterior margin of pronotum broad; valviceps slightly narrowed toward apex, ergot long below. M. circulotibialis Li, Liu & Heng, 2015 Hind tibia with a white macula dorsally, but not forming; all trochanters entirely white; white macula on posterior margin of pronotum narrow; valviceps not narrowed toward apex, ergot short above. M. postscutellaris Malaise, 1945 Fore and middle trochanters mostly to entirely black.25 Fore and middle trochanters entirely white.29 Hind trochanter mostly to entirely black.26 Hind trochanter entirely white.28 Hind trochanter entirely black; pronotum entirely black. M. bui Wei & Li, 2012 Hind trochanter mostly black, marginal parts white; posterior margin of pronotum with white band more or less.27 Posterior margin of pronotum with broad white band; hind tibia with a distinct dorsal white macula; ergot of pennis valve long below. M. curvatitheca Li, Liu & Heng, 2015 Posterior margin of pronotum with narrow white band; hind tibia with dorsal white macula weak or distinct; ergot of penis valve short above. M. nigromaculata Wei & Li, 2010 White band in posterior margin of pronotum narrow but distinct; interspaces between punctures on vertex as broad as diameter of a puncture; valvula of penis valve not broadened toward apex. M. imitatoides Wei, 2007 Posterior margin of pronotum entirely black; interspaces between punctures on vertex narrower than diameter of a puncture; valvula of penis valve clearly broadened toward apex. M. imitator Jakeuch\, 1937 Labrum and clypeus entirely white.30 Labrum largely black and clypeus entirely black.32 Punctures on vertex large; ventral surface of hind femur black, without white band; valviceps approximately 2.3x longer than broad. M. kangdingensis Wei & Li, 2012 Punctures on vertex minute; ventral surface of hind femur with distinct white band; valviceps clearly 1.4-1.7x longer than broad.31 White band on posterior margin of abdominal tergum 1 very narrow; entire posterior margin of pronotum with narrow white macula; sterna of abdomen entirely black. M. flactoserrula Chen & Wei, 2002 White band submedially on posterior margin of abdominal tergum 1 approximately 2/5 of its breadth clearly; posterior margin of pronotum with a distinct, broad white macula; sterna of abdomen largely white. M. funiushana Wei, 1998 White band at posterior margin of pronotum distinct. M. curvatisaeta Wei & Li, 2011 White band at posterior margin of pronotum weak or indistinct. M. jiaozhaoae Wei & Zhao, 2011 Macrophya longlingensis Li, Liu & Wei, sp. nov. http://zoobank.org/6129209F-9628-4C4B-9CBC-F5B241B25 A4F Figure 1 Diagnosis. The new species is morphologically similar to M parimitatorWQi, 1998 in body and legs mainly black; antennae rather robust, middle antennomeres not inflated; anterior margin shallowly incised to approximately 1/3 its length, lateral corners somewhat short and broad; malar space linear, approximately 0.5x as broad as di¬ ameter of middle ocellus; lancet narrow and long, with 20 serrulae; but differs from the latter in having vertex shiny; frontal area coarsely and densely punctured, with smooth interspaces between punctures distinct; anterior 1/6 of katepimeron very smooth and shiny, without punc¬ tures or microsculpture, posterior 5/6 with some shallow large punctures, microsculpture indistinct; dorsal surface of middle tibia black, without white macula subapically; dorsal surface of hind tibia with a small, narrow white macula; cell 2Rs of fore wing clearly shorter than cell IR,, petiole of anal cell in hind wing as long as cross¬ vein cu-a; middle serrulae with 2 proximal and 8-11 dis¬ tal teeth, subbasal teeth clear and small. The new species is also morphologically similar to M. jiaozhaoae Wei & Zhao, 2011 in body and legs mainly black; antennae zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 417-427 421 rather robust, middle antennomeres not inflated; ventral surface of hind trochanter with a small, weak black mac¬ ula; pronotum entirely black; fronts flat and not extending above top of eyes; hind tibia with subapical white macula distinctly shorter than half length of hind tibia; ovipositor sheath clearly shorter than middle tibia; but differs from the latter in having anterior margin of clypeus incised to approximately 1/3 its length; middle serrulae with 2 proximal and 8-11 distal teeth; petiole of anal cell in hind wing as long as cross-vein cu-a. Description. Holotype: female. Body length 7 mm. Body and legs black; following parts pale brown: palp mostly, a small triangular macula on apical margin of clypeus, ventral surface of fore tarsomere mostly; following parts white: basal half of mandibles, transverse macula subme- dially on posterior margin of abdominal tergum 1, apical margins of all coxae, apical margins of fore and middle trochanters, hind trochanter except for ventral surface with a small black macula, apex of fore and middle fem¬ ora anteriorly, anterior surface of middle tibia, hind tibia with long, narrow subapical macula on dorsal surface. Body hairs short and dense, silver; setae on ovipositor sheath slightly long and curved, blackish brown. Wings hyaline, without smoky macula, pterostigma and veins mostly blackish brown (Figure lA). Vertex shiny; frontal area coarsely and densely punctured clearly, smooth interspaces distinct and smooth; interspaces of postocellar area with small areas and some large punctures, interspaces distinct and without microsculpture (Figure IB); labrum and clypeus less shiny, punctures on labrum sparse, punctures on clypeus denser, micro sculpture weak (Figure 1C). Mesonotum less shiny, punctures on mesonotum smaller and denser than punctures on head, interspaces distinct and smooth; center of mesoscutellum with some large punctures, interspaces broad, marginal area with denser punctures than center; mesoscutellar appendage mostly and metascutellum entirely smooth and shiny, but bottom of mesoscutellar appendage with weak microsculpture and without distinct punctures. Mesopleuron less shiny, mesepisternum with dense and coarse punctures, upper half with punctures large and interspaces broad, lower half with punctures small and interspaces narrow; anepimeron dull, with coarse wrinkles; anterior 1/6 of katepimeron very smooth and shiny, without punctures or microsculpture, posterior 5/6 of katepimeron with some shallow large punctures, dorsal half with some deep punctures; metepisternum dull, with minute punctures, microsculpture distinct; metepimeron less shiny, depressed area with some punctures and weak micro sculpture; metepimeronal appendage platform¬ shaped, with some minute punctures (Figure IE). All abdominal terga shiny, laterally abdominal tergum 1 with some shallow punctures, nearly smooth submedially; other abdominal terga less shiny, anterior 2/3 with some shallow punctures and weak microsculpture, posterior 1/3 smooth. Outer surface of hind coxa with somewhat dense and coarse punctures, outer surface of hind femur with sparse shallow punctures and fine microsculpture. Surface of ovipositor sheath coriaceous, with indistinct punctures and fine microsculpture. Labrum elevated medially, anterior margin slightly truncate; clypeus weakly elevated, base slightly broader than distance between lower inner orbits of eyes, lateral sides distinctly convergent apically, anterior margin shal¬ lowly incised to approximately 1/3 its length, lateral cor¬ ners somewhat short and broad, lobe margin subtriangular (Figure 1C); malar space linear, approximately 0.5x as broad as diameter of middle ocellus; frontal area and face flat, slightly higher than top of eyes in lateral view; mid¬ dle fovea weak, pot-shaped, lateral foveae clear, short fur¬ row-like; interocellar furrow shallow, postocellar furrow weak; POL: OOL: OCL = 8: 12: 9; postocellar area weak¬ ly elevated, approximately 2.2x broader than long; lateral furrow somewhat broad and shallow, divergent posterior¬ ly; head narrowed behind eyes in dorsal view, occipital Ca¬ rina complete. Antenna rather robust, approximately Fix longer than head and thorax together (16: 15), approxi¬ mately as long as abdomen; antennomere 2 approximate¬ ly 1.3 x as long as breadth; antennomere 3 approximately 1.5x as long as antennomere 4 (43: 29), approximately 0.8x as long as antennomeres 4 and 5 together (43: 55), middle antennomeres not inflated, subapical antennom¬ eres weakly compressed (Figure ID). Mesoscutellum el¬ evated, without median ridge or carina, as high as top of mesonotum in lateral view; mesoscutellar appendage with acute middle longitudinal carina; metascutellum with short and low carina; posterodorsal platform of mesepimeron as broad as diameter of middle ocellus; metepimeronal ap¬ pendage small platform-shaped; distance between cenchri 2.5x breadth of a cenchrus; mesopleuron and metapleuron as shown in Figure IE. Inner tibial spur of hind leg 0.6x length of metabasitarsus (20: 33); metabasitarsus slender, about Fix longer than following four tarsomeres together (IF 10); claw with inner tooth slightly shorter than outer tooth. Ovipositor sheath slightly shorter than metabasi¬ tarsus (10: 11), apical sheath longer than basal sheath (3: 2), setae on ovipositor sheath slightly curved and long in dorsal view, apical margin round in lateral view (Figure IF) . Fore wing with cross-vein cu-a joining cell IM in basal 1/3, cross-vein 2r joining cell 2Rs in apical 1/5, cell 2Rs clearly shorter than cell IR,, petiole of anal cell twice longer than cross-vein Ir-m and as long as cross-vein cu- a; petiole of anal cell in hind wing as long as cross-vein cu-a. Lancet narrow and long, with 20 serrulae (Figure IG) , serrulae slightly protruding and oblique, middle ser¬ rulae with 2 proximal and 8-11 distal teeth, subbasal teeth distinct and small, annular spine bands somewhat broad, the 7*-9* serrulae as shown in Figure IH. Male. Unknown. Type material. Holotype, $, China: Yunnan Prov¬ ince: Longling County, Mount Xiaohei, 24°4F713'N, 98°45.574'E, 2010 m, 2.vi.2009, leg. Zejian Li, ethylac- zse.pensoft.net 422 Mengmeng Liu et al.: Three new species of Macrophya Imitator group Figure 1. Macrophya longlingensis sp. nov., holotype. A. Female adult, dorsal view; B. Head of female, dorsal view; C. Head of female, frontal view; D. Antenna of female, lateral view; E. Mesopleuron and metapleuron of female; F. Ovipositor sheath, lateral view; G. Lancet; H. The 7*-9* serrulae. Scale bars: 2 mm (A); 100 pm (G); 50 pm (H). etate. Paratypes, 1$, same data, but leg. Gengyun Niu; 2$, Yunnan Province: Longling County, Mount Gaoli- gong, 24°49.700'N, 98°46.062'E, 2145 m, 2.vi.2009, leg. Yihai Zhong, ethylacetate; 1$, Yunnan Province: Yun- long County, Mount Daoren, 25°32.893'N, 99°11.267'E, 2265 m, 3.vi.2009, leg. Gengyun Niu, ethylacetate; 1$, Yunnan Province: Eushui County, Yaojiaping, 25°975'N, 98°710'E, 2550 m, 3.vi.2009, leg. Wei Xiao, ethylacetate. Host plants. Unknown. Distribution. China (Yunnan). Etymology. The specific name '"longlingensis’' is derived from Eongling County (Yunnan Province) where the ho¬ lotype was collected. Macrophya nieshuaiguoi Li, Liu & Wei, sp. nov. http://zoobank.org/DC7F944E-39C9-4E30-A022-F33927E38791 Figure 2 Diagnosis. The new species is morphologically similar to M. jiaozhaoae Wei & Zhao, 2010 in body and legs main¬ ly black; antennae rather robust, middle antennomeres not inflated; lancet narrow and long, serrulae slightly protrud¬ ing and oblique; hind tibia with subapical white macula distinctly shorter than half length of hind tibia; ovipositor sheath clearly shorter than middle tibia; but differs from the latter in having vertex less shiny, interspaces of postocular area and postocellar area with some large punctures, inter¬ spaces between punctures distinct; anterior margin of cly- peus deeply incised to approximately 2/5 its length; posto¬ cellar area about 1.6x broader than long; posterior margin zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 417-427 423 Figure 2. Macrophya nieshuaiguoi sp. nov., holotype. A. Female adult, dorsal view; B. Head of female, dorsal view; C. Head of female, frontal view; D. Antenna of female, lateral view; E. Mesopleuron and metapleuron of female; F. Ovipositor sheath, lateral view; G. Lancet; H. The 8*-10* serrulae. Scale bars: 2 mm (A); 100 pm (G, H). of abdominal tergum 1 with broad white band; petiole of anal cell in fore wing shorter than cross-vein cu-a, petiole of anal cell in hind wing as long as cross-vein cu-a; setae on ovipositor sheath slightly curved and sparse in dorsal view; middle serrulae with 2 proximal and 7-10 distal teeth. The new species is also morphologically similar to M. imitator Takeuchi, 1937 in body and legs mainly black; antennae rather robust, middle antennomeres not inflated; pronotum and hind trochanter entirely black; but differs from the lat¬ ter in having postocellar area twice broader than long; hind tibia with subapical white macula on dorsal surface weak; middle serrulae with 2 proximal and 7-10 distal teeth. Description. Holotype: female. Body length 8 mm. Body and legs black; palp mostly blackish brown; a small triangular macula on apical margin of clypeus pale brown; following parts white: basal half of mandibles, broad band on posterior margin of abdominal tergum 1 submedially, apical margins of fore and middle cox¬ ae, their outer surfaces with some longitudinal stripes, apical margin of hind coxa, fore and middle trochanters narrowly, hind trochanter entirely, anterior surface of fore and middle femora apically, anterior surface of fore tibia, hind tibia with small dorsal macula subapically; ventral surfaces of fore and middle tarsomeres mostly pale brown to pale blackish brown. Body hairs short and dense, silver; setae on ovipositor sheath slightly long and curved, blackish brown. Wings hyaline, without smoky macula, pterostigma and veins mostly blackish brown (Figure 2A). zse.pensoft.net 424 Mengmeng Liu et al.: Three new species of Macrophya imitator group Vertex less shiny; frontal area coarsely and densely punctured, interspaces smooth and narrow; interspaces of postocular area and postocellar area with sparse large punc¬ tures, interspaces between punctures distinct (Figure 2B); labrum and clypeus less shiny, labrum and clypeus with sparse shallow large punctures, microsculpture distinct; punctures on labrum shallow, punctures on clypeus denser toward apex (Figure 2C). Mesonotum less shiny, punc¬ tures smaller than those on head, interspaces smooth but indistinct and without microsculpture; center of mesoscu- tellum with sparse large punctures, interspaces broad, marginal area with dense punctures than center; mesoscu- tellar appendage somewhat shiny, with sparse minute punctures and weak microsculpture; metascutellum some¬ what shiny, with punctures indistinct and microsculpture weak. Mesopleuron less shiny, mesepisternum with dense and coarse punctures, upper half with punctures large and interspaces broad, lower half with punctures small and in¬ terspaces narrow; anepimeron dull, with coarse wrinkles; anterior 1/5 of katepimeron very smooth and shiny, with¬ out punctures or microsculpture, posterior 4/5 of katepi¬ meron with some large shallow punctures, dorsal half with sparse coarse punctures; metepisternum dull, with minute punctures, microsculpture clear; metepimeron less shiny, depressed area with sparse punctures and weak mi¬ crosculpture; metepimeronal appendage platform-shaped, with sparse minute punctures (Figure 2E). All abdominal terga somewhat shiny, two lateral sides of abdominal ter- gum 1 with sparse shallow punctures, central parts with fine but distinct microsculpture; other abdominal terga less shiny, anterior 3/5 of abdominal terga 2-8 with sparse shallow punctures, posterior 2/5 of abdominal terga 2-8 with weak microsculpture. Outer surface of hind coxa with somewhat dense and coarse punctures, outer surface of hind femur with some shallow punctures and fine mi¬ crosculpture. Surface of sheath coriaceous, with indistinct punctures and fine microsculpture. Labrum elevated medially, anterior margin slightly truncate; clypeus weakly elevated, base slightly broader than distance between lower inner orbits of eyes, lateral sides distinctly convergent apically, anterior margin deep¬ ly incised to approximately half its length, lateral corners short and broad, lobe margin roundly subtriangular (Fig¬ ure 2C); malar space linear, approximately 0.6x as broad as diameter of middle ocellus; frontal area and face flat, as high as top of eyes in lateral view; middle fovea weak, lat¬ eral foveae shallow, short furrow-like; interocellar furrow shallow, postocellar furrow weak; POL: OOL: OCL = 7: 20: 13; postocellar area weakly elevated, approximately twice as broad as long; lateral furrow somewhat narrow, divergent posteriorly; head narrowed behind eyes in dor¬ sal view, occipital carina complete. Antenna rather robust, approximately 1.3x longer than head and thorax together (4: 3), approximately 1.2x longer than abdomen (20: 17); antennomere 2 approximately 1.2x as long as breadth; an- tennomere 3 approximately 1.4x as long as antennomere 4 (33: 23), approximately 0.75x as long as antennomeres 4 and 5 together (33: 44), middle antennomeres not inflated. subapical antennomeres weakly compressed (Figure 2D). Mesoscutellum elevated roundly, without median ridge or carina, as high as top of mesonotum in lateral view; mesoscutellar appendage with slightly acute middle lon¬ gitudinal carina; metascutellum with short and low carina; posterodorsal platform of mesepimeron as broad as diam¬ eter of middle ocellus; metepimeronal appendage small platform-shaped; distance between cenchri 2.2x breadth of a cenchrus; mesopleuron and metapleuron as shown in Figure 2E. Inner tibial spur of hind leg 0.6x length of metabasitarsus (3: 5); metabasitarsus slender, about 1.3x longer than following four tarsomeres together (5:4); claw with inner tooth slightly shorter than outer tooth. Ovipos¬ itor sheath shorter than metabasitarsus (31: 45), apical sheath clearly longer than basal sheath (20: 11), setae on ovipositor sheath slightly curved in dorsal view, apical margin round in lateral view (Figure 2F). Fore wing with cross-vein cu-a joining cell IM in basal 1/3, cross-vein 2r joining cell 2Rs in apical 1/3, cell 2Rs clearly slightly shorter than cell IR,, petiole of anal cell 1.5x longer than cross-vein Ir-m and slightly shorter than cross-vein cu-a; petiole of anal cell in hind wing as long as cross-vein cu- a. Lancet narrow and long, with 24 serrulae (Figure 2G), serrulae slightly protruding and oblique, middle serrulae with 2 proximal and 7-10 distal teeth, subbasal teeth dis¬ tinct and small, annular spine bands somewhat narrow, the 8*-10* serrulae as shown in Figure 2H. Male. Unknown. Type material. Holotype, $, China: Yunnan Province: Liuku County, Pianma, Yakou, 25°58.21'N, 98°41.06'E, 3138 m, 19.vii. 2008, leg. Shuaiguo Nie, ethylacetate. Host plants. Unknown. Distribution. China (Yunnan). Etymology. The specific name '"nieshuaiguor is derived from the name of Mr. Shuaiguo Nie for collecting the ho¬ lotype of this new species. Macrophya zejiani Liu & Wei, sp. nov. http://zoobank.org/1528F16E-8F84-4FE3-9682-60D4733E2A20 Figure 3 Diagnosis. The new species is morphologically similar to M. weni Wei, 1998 in body and legs mainly black; an¬ tennae rather robust, middle antennomeres not inflated; ovipositor sheath much longer than middle tibia; but dif¬ fers from the latter in having postocellar area twice as broad as long; pronotum entirely black; distance between cenchri twice breadth of a cenchrus; middle serrulae with 2 or 3 proximal and 9 or 10 distal teeth; fore wing below pterostigma with slightly smoky and ill-defined maculae. The new species is also morphologically similar to M. omeialpina Li, Jiang & Wei, 2018 in body and legs main¬ ly black; antennae rather robust, middle antennomeres zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 417-427 425 Figure 3. Macrophya zejiani sp. nov., holotype. A. Female adult, dorsal view; B. Head of female, dorsal view; C. Head of female, frontal view; D. Antenna of female, lateral view; E. Mesopleuron and metapleuron of female; F. Ovipositor sheath, lateral view; G. Lancet; H. The 8*''-10* serrulae. Scale bars: 2 mm (A); 100 pm (G); 50 pm (H). not inflated; pronotum entirely black; dorsal surface of hind tibia with a subapical white macula smaller than M. weni; but differs from the latter in having postocellar area 2.5x broader than long; POL: OOL: OCL = 4.5: 10: 5.5; antennomere 3 approximately 1.4x as long as antennom- ere 4 (13: 9); distance between cenchri twice breadth of a cenchrus; middle serrulae with 2 or 3 proximal and 9 or 10 distal teeth; cell 2Rs as long as cell IRj, petiole of anal cell in hind wing 0.6x as long as cross-vein cu-a. Description. Holotype: female. Body length 7.5 mm. Body and legs black; a small triangular macula in anteri¬ or margin of clypeus pale brown; following parts white: basal half of mandibles, narrow band on posterior margin of abdominal tergum 1 submedially, apical margins of fore and middle coxae, apical half in anterior surface of fore femur, base mostly in anterior surface of fore tibia, hind trochanter entirely, hind tibia with small dorsal mac¬ ula subapically. Body hairs short and dense, silvery; se¬ tae on ovipositor sheath slightly curved, blackish brown. Wings hyaline, below pterostigma with pale smoky mac¬ ula, boundary ill-deflned, pterostigma and veins mostly blackish brown (Figure 3A). Vertex less shiny; frontal area coarsely and densely punctured, interspaces smooth but weak; postocellar area mostly with sparse large punctures, interspaces narrow (Figure 3B); labrum and clypeus less shiny, punctures on labrum and clypeus sparse shallow and microsculpture fine (Figure 3C). Mesonotum less shiny, punctures small¬ er than those on head, interspaces smooth but indistinct and without microsculpture; center of mesoscutellum with sparse large punctures and fine microsculpture; basal half of mesoscutellar appendage rugose, apical half smooth, without distinct puncture; metascutellum somewhat shiny, punctures indistinct and microsculpture weak. Mesopleu¬ ron less shiny, mesepisternum with dense and coarse punc¬ tures, interspaces smooth but indistinct; anepimeron dull, with coarse wrinkles; anterior margin of katepimeron very zse.pensoft.net 426 Mengmeng Liu et al.: Three new species of Macrophya imitator group Figure 4. Geographical distribution map of M imitator group in China. smooth and shiny, without punctures or microsculpture, otherwise with sparse large shallow punctures, dorsal half with sparse coarse punctures; metepisternum dull, with minute punctures; metepimeron less shiny, most parts with sparse punctures and weak microsculpture; metepimeronal appendage platform-shaped, with some minute punctures (Figure 3E). All abdominal terga somewhat shiny, abdom¬ inal tergum 1 with sparse shallow punctures, with fine but distinct microsculpture submedially; other abdominal terga with minute and shallow punctures, microsculpture weak. Outer surface of hind coxa with somewhat dense and coarse punctures, ventral surface of hind coxa and out¬ er surface of hind femur with sparse shallow punctures and fine microsculpture. Surface of ovipositor sheath coria¬ ceous, with indistinct punctures and fine microsculpture. Labrum elevated medially, anterior margin slightly truncate; clypeus weakly elevated, base slightly broader than distance between lower inner orbits of eyes, later¬ al sides distinctly convergent apically, anterior margin deeply incised to approximately 1/5 its length, lateral cor¬ ners short and broad, lobe margin roundly subtriangular (Figure 3C); malar space linear, approximately 0.5x as broad as diameter of middle ocellus; frontal area and face fiat, slightly higher than top of eyes in lateral view; mid¬ dle fovea weak, lateral foveae shallow, short furrow-like; interocellar furrow shallow, postocellar furrow weak; POL: OOL: OCL = 9: 20: 11; postocellar area weakly el¬ evated, 2.5x broader than long; lateral furrow somewhat narrow, divergent posteriorly; head narrowed behind eyes in dorsal view, occipital carina complete. Antenna rather robust, approximately 1.2x longer than head and thorax together (17:14), approximately Fix longer than abdo¬ men (17: 15); antennomere 2 approximately 1.2x as long as breadth; antennomere 3 approximately 1.4x as long as antennomere 4 (39: 27), approximately 0.8x as long as antennomeres 4 and 5 together (39: 51), middle antenno- meres not infiated, subapical antennomeres weakly com¬ pressed (Figure 3D). Mesoscutellum roundly elevated, with weak middle ridge or carina, as high as top of me- sonotum in lateral view; mesoscutellar appendage with lower middle longitudinal carina; metascutellum with short and lower carina; posterodorsal platform of mese- pimeron as broad as diameter of middle ocellus; metepi¬ meronal appendage small platform-shaped; distance be- zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 417-427 427 tween cenchri twice breadth of a cenchrus; mesopleuron and metapleuron as shown in Figure 3E. Inner tibial spur of hind leg 0.6x length of metabasitarsus (5: 8); metaba- sitarsus slender, about l.lx longer than following four tarsomeres together (8: 7); claw with inner tooth slightly shorter than outer tooth. Ovipositor sheath clearly longer than metabasitarsus (5: 4), apical sheath as long as basal sheath, setae on ovipositor sheath slightly curved in dor¬ sal view, apical margin round in lateral view (Figure 3F). Fore wing with cross-vein cu-a joining cell IM in basal 1/4, cross-vein 2r Joining cell 2Rs in apical 1/4, cell 2Rs as long as cell IRj, petiole of anal cell slightly shorter than cross-vein Ir-m and petiole of anal cell in hind wing 0.6x longer than cross-vein cu-a. Lancet narrow and long, with 21 serrulae (Figure 3G), serrulae clearly protruding and oblique platform-shaped, middle serrulae with 2 or 3 proximal and 9 or 10 distal teeth, subbasal teeth distinct and small, annular spine bands narrow, the 8*-10* serru¬ lae as shown in Figure 3H. Male. Unknown. Type material. Holotype, China: Yunnan Prov¬ ince: Deqin County, Mountt Meri, 28°425'N, 98°805'E, 2700 m, 20.vi.2009, leg. Yihai Zhong, ethylacetate. Host plants. Unknown. Distribution. China (Yunnan). Etymology. The specific name '"zejianr is derived from the name of Dr. Zejian Li who has made great contribu¬ tions to the study of Macrophya systematics. Discussion The Macrophya imitator group orginally proposed by Liu et al. (2015) is the second largest of defined species group of Macrophya Dahlbom, 1835 in terms of known species. At present, there are 20 species belonging to this group in China, including the three new species described above: M bui Wei & Li, 2012, M changbaina Li, Liu & Heng, 2015, M circulotibialis Li, Liu & Heng, 2015, M curva- tisaeta Wei & Li, 2010, M curvatitheca Li, Liu & Heng, 2015, M flactoserrula Chen & Wei, 2002, M. funiushana Wei, 1998, M. imitatoidesWQi, 2007, M. imitator Takeu- chi, 1937, M. jiaozhaoae Wei & Zhao, 2010, M. kang- dingensis Wei & Li, 2012, M. longlingensis Li, Liu & Wei, sp. nov., M.nieshuaiguoi Li, Liu & Wei, sp. nov., M. nigromaculata Wei & Li, 2010, M omeialpina Li, Jiang & Wei, 2018, M parimitator Wei, 1998, M. postscutellar- is Malaise, 1945, M. semipunctata Li, Liu & Wei, 2018, M weni Wei, 1998 and M. zejiani Liu & Wei, sp. nov. The host plants of this group are unknown. The included key and geographical distribution map of the M. imitator group should facilitate the recognition and identification of the Chinese species. Acknowledgements The authors are deeply grateful to Dr. Kees van Achter- berg and Spencer K. Monckton for valuable comments and suggestions. This research was partly supported by the Natural Science Foundation of Zhejiang Province (No. LY18C040001) and the National Natural Science Foundation of China (No. 31672344). References Benson RB (1954) Some sawflies of the European Alps and the Medi¬ terranean region (Hymenoptera: Symphyta). Bulletin of the British Museum (Natural History). Entomology 3(7): 267-295. https://doi. org/10.5962/bhl.part. 1054 Ei ZJ, Wei MC, Eiu MM, Chen ME (2018) Macrophya Dahlbom in Chi¬ na. China Agricultural Science and Technology Press, Beijing, 456 pp. Ei Z J, Eiu MM, Wei MC (2019a) A new species of Macrophya Dahlbom (Hymenoptera: Tenthredinidae) with a key to species of the Mac¬ rophya coxalis group from China. Entomological Research 49(2): 105-109. https://doi.org/10.1111/1748-5967.12335 Ei ZJ, Eiu MM, Wei MC (2019b) Three new species of Macrophya maculitibia group (Hymenoptera: Tenthredinidae) with a key to known species from China. Zoosystematics and Evolution 95(1): 37-48. https://doi.org/10.3897/zse.95.28804 Eiu MM, Heng XM, Eiang XM, Zhong YH, Ei ZJ (2015) Three new spe¬ cies of imitator-group of the genus Macrophya (Hymenoptera: Ten¬ thredinidae) from China. Zoological Systematics 40(2): 212-222. Eiu MM, Ei ZJ, Wei MC (2019a) Review of the Macrophya formosa- na group (Hymenoptera: Tenthredinidae) from China with descrip¬ tions of two new species. Entomological Research 49(5): 203-213. https://doi.org/10.1111/1748-5967.12347 Eiu MM, Hong Z, Zhong YH, Ei ZJ, Wei MC (2019b) Two new species of Macrophya flavomaculata group (Hymenoptera, Tenthredinidae) from China. Entomotaxonomia 41(1): 8-18. Malaise R (1945) Tenthredinoidea of South-Eastern Asia with a general zoogeographical review. Opuscula Entomologica, Eund, Suppl. 4: 1-288. https://doi.Org/10.1080/11035894509446460 Ross HH (1945) Sawfly genitalia: terminology and study techniques. Entomological News 61(10): 261-268. Takeuchi K (1937) A study on the Japanese species of the genus Macro¬ phya (Hymenoptera Tenthredinidae). Tenthredo. Acta Entomologica 1(4): 376-454. Takeuchi K (1952) A Generic Classification of the Japanese Tenthredin¬ idae (Hymenoptera: Symphyta). Kyoto, 90 pp. Viitasaari M (2002) The Suborder Symphyta of the Hymenoptera. In: Viitasaari M (Ed.) Sawfiies (Hymenoptera, Symphyta) I. A Review of the Suborder, the Western Palaearctic Taxa of Xyeloidea and Pamphilioidea. Tremex, Helsinki, 11-174. zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 429-452 | DOI 10.3897/zse.95.35435 4>yEnsPFr. BERLIN The paleoichthyofauna housed in the Coleecion Naeional de Paleontologia of Universidad Naeional Autonoma de Mexico Kleyton Magno Cantalice\ Alejandra Martinez-Melo^’^, Violeta Amparo Romero-Mayen^ 1 Departamento de Paleontologia, Instituto de Geologia, Universidad Naeional Autonoma de Mexico, Circuito de la Investigacion S/N, Ciudad Universitaria, Coyoacdn, Ciudad de Mexico, 04510, Mexico 2 Coleecion Naeional de Paleontologia, Instituto de Ceologia, Universidad Naeional Autonoma de Mexico, Circuito de la Investigacion S/N, Ciudad Universitaria, Coyoacdn, Ciudad de Mexico, 04510, Mexico 3 UMR 7207, Centre de recherche en Paleontologie - Paris. 8 rue Bujfon, CP 38, 75005 Paris, France http://zoobank.org/514DEB4F-BD40-4EDl-898B-27A5F9013FB5 Corresponding author; Kleyton Magno Cantalice (kleytonbio@yahoo.com.br) Academic editor: Mco/a5♦ Received 14 April 2019 ♦ Accepted 3 August 2019 ♦ Published 30 August 2019 Abstract Fishes are a paraphyletic group composed by craniates except for the four-limbed clade Tetrapoda. This group was the only vertebrate representative until the Devonian but now comprises almost half of the vertebrate species, dominating nearly all aquatic environ¬ ments. The fossil record is the key to understand the ancient paleobiodiversity and the patterns that lead the modem fish fauna, and paleontological collections play a fundamental role in providing accommodation, maintenance, and access to the specimens and their respective metadata. Here we present a systematic checklist of fossil fishes housed in the type collection of the Coleecion Naeional de Paleontologia which is located at the Instituto de Geologia of Universidad Naeional Autonoma de Mexico. Currently housed in the type collection are 14 chondrichthyan specimens, belonging to two superorders, five orders, seven families, 10 genera, and five nom¬ inal species, and 361 osteichthyan specimens, belonging to eight orders, nine families, nine genera, and 26 nominal species. These fossils come from 32 localities and 15 geological units, which range temporally from the Jurassic to the Pleistocene. The paleoich¬ thyofauna housed in the type collection of the Coleecion Naeional de Paleontologia is remarkable for its singularity and reveals new insights about the origin and diversification of many groups of fishes. The recovery and curation of this fossil material indicates that knowledge of Mexican fossil fish diversity and its role in understanding lower vertebrate evolution are just emerging and reaffirms the importance of the biological and paleontological collections to the future biodiversity research. Key Words Collection, diversity, fishes, paleontology, taxonomy, Mexico Introduction Fishes are craniate animals that have gill arches and use fins for locomotion in aquatic environments (Berra 2007; Nelson et al. 2016; Clarke and Friedman 2018). Among vertebrates, fishes exhibit incomparable diversity in mor¬ phology, behavior, physiology, and distribution (Nelson et al. 2016). Currently, the extant fishes are classified in four distinct classes: Myxini (hagfishes), Petromyzontida (lampreys), Chondrichthyes (cartilaginous fishes, such as sharks and stingrays), and Osteichthyes. The last class is divided into ray-finned (Actinopterygii) and lobe-finned fishes. Dipnoi and Actinistia, respectively (Helfman et al. 1997). Nevertheless, if we consider the extinct ichthyo¬ fauna, the number of classes at least doubles (see Nelson et al. 2016). In the current understanding of vertebrate systematics, fishes constitute a paraphyletic group (Gill and Mooi 2002; Berra 2007) because it excludes the four-limbed osteich¬ thyan clade Tetrapoda, which shares a common ancestor Copyright Kleyton Magno Cantalice etal. This is an open access articie distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the originai author and source are credited. 430 Kleyton Magno Cantalice et al.: Fossil fishes in the Mexican National Collection of Paleontology with lobe-finned fishes (Zhu and Yu 2002). Nevertheless, with at least 32,000 species, fishes represent one-half of the world’s living vertebrates and the study of fishes contrib¬ utes to many subjects of scientific natural history knowl¬ edge, such as ontogeny, distribution, speciation, and di¬ versification through space and time (Kornfield and Smith 2000; Penaz 2001, Cavin et al. 2008; Nelson et al. 2016). Fossils are physical evidence that helps in the recogni¬ tion and interpretation of biological patterns and process¬ es on Earth through time (Cavin et al. 2008). Neverthe¬ less, by its composition, fossil material is so fragile that specific care is required for good long-term preservation, and museums and academic institutions are responsible for protecting and conserve this material (Allmon 1994, 2005; Llorente-Bousquets et al. 1994). Paleontological collections not only have the respon¬ sibility of accommodating and storing the fossil record but also of providing the care and good conditions to use these materials for scientific research purposes and en¬ tertainment (Llorente-Bousquets et al. 1994; Cristin and Perrilliat 2011). Good practices in museum curation, such as the collection of specimens in appropriate spaces and conditions, the creation of systematic catalogues, and regulations for the proper use of fossils, are essential to generate and validate the information on which the ad¬ vance of science is possible (Allmon 2005). Furthermore, paleontological collections have an important social and teaching role in spreading the scientific discoveries ac¬ cessible to the public (Suarez and Tsutsui 2004). The Coleccion Nacional de Paleontologia of Univer- sidad Nacional Autonoma de Mexico (CNP-UNAM) is remarkable by its number of specimens collected in many regions of the country from various geological ages (e.g. Perrilliat 1993; Cristin and Perrilliat 2011; Perrilliat and Castaneda-Posadas 2013; Rojas-Zuniga and Gio-Argaez 2016). Although the birth of CNP-UNAM was at the end of the 19* century, its formal consolidation inside the Institute of Geology occurred between 1978 and 1986. Only in 2004 did this collection become recognized as the National Collection of Paleontology and given the des¬ ignation as the “Museum Maria del Carmen Perrilliaf’. All fossils housed at the CNP-UNAM have the acronym (IGM) and began to be incorporated in 1978. Previous fossil records are in other institutions or are lost. Today, the CNP-UNAM has five sections: geograph¬ ic reference, foreign materials. Recent materials, molds, and the collection of types (Perrilliat et al. 1986; Carreno and Montellano-Ballesteros 2005). The type collection includes: 1) specimens belonging to type series and 2) voucher specimens, which are recorded in this collec¬ tion as hypotypes. This section comprises about 10,000 specimens of microfossils, plants, invertebrates, and ver¬ tebrates, ranging from the end Precambrian to the Quater¬ nary period of the Cenozoic (e.g. Perrilliat 1993; Perrilliat and Castaneda-Posadas 2013). After 25 years since the last report about the fossil vertebrates housed in the type collection of CNP-UNAM (Perrilliat 1993), fossil fish specimens have been incre¬ mentally added and many changes in fish taxonomy and classification have occurred. Therefore, following Article 72 of the International Code of Zoological Nomenclature, we present the systematic list of fishes currently housed in this collection and their respective localities. Further¬ more, we provide information about the implications of these discoveries to understanding the taxonomy, bioge¬ ography, and early evolutionary history of some taxa and highlight the importance of biological collections to fu¬ ture research on paleodiversity in Mexico. Methods All fish fossils housed in the CNP-UNAM type collec¬ tion were reviewed. Information on the taxonomy, age, and distribution are from both the collection database and the literature. For each species, we include the catalogue number (IGM), taxonomic classification, and respective distribution and age. Nomenclature on extinct Chon- drichthyes follows Nelson et al. (2016) and Van de Laan (2018), while nomenclature on Recent Osteichthyes fol¬ lows Betancur-R. et al. (2017) and Fricke et al. (2019). The nomenclature of extinct bony fishes follows Nelson et al. (2016), Van der Laan (2018), and original referenc¬ es. The maps were created with QGIS software version 2.18.19 (QGIS Development Team 2018) and the fossil fish localities were plotted using the software Arc View version 3.3 (Environmental Systems Research Institute, Inc., Redlands, California). Most of the taxonomic data¬ base used is available under open access at Unidad In- formatica para la Paleontologia of UNAM (UNIPALEO; http ://www. unipaleo. unam. mx). Results 1. Systematic checklist list of CNP-UNAM fossil fishes Subphylum Vertebrata Cuvier, 1812 Infraphylum Gnathostomata Zittel, 1879 Class Chondrichthyes Huxley, 1880 Subclass Elasmobranchii Bonaparte, 1838 Cohort Euselachii Hay, 1902 Order tHybodontiformes Maisey, 1975 Superfamily tHybodontoidea Owen, 1846 Family tHybodontidae Agassiz, 1843 Genus "fPlanohybodus Rees & Underwood, 2008 ^Planohybodus indet. Referred specimen. IGM 9316, IGM 9317 (Alvara- do-Ortega et al. 2014). Locality and age. Llano Yosobe, Sabinal Formation, Tlaxiaco, Oaxaca; Jurassic (Kimmeridgian-Tithonian). zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 429-452 431 Subcohort Neoselachii Compagno, 1977 Superorder Galeomorphii Compagno, 1973 Order Carcharhiniformes Compagno, 1973 Carcharhiniformes indet. Referred specimen. IGM 6990 (Ferrusquia-Villafranca etal. 1999). Locality and age. Rancho el Jobo, San Juan Formation, Tuxtla Gutierrez, Chiapas; Middle Eocene. Family Carcharhinidae Jordan & Evermann, 1896 Genus Galeocerdo Muller & Henle, 1837 fGaleocerdo rosaliensis Applegate, 1978 Referred specimen. IGM 5854 (holotype). Locality and age. Tirabuzon Formation, Santa Rosalia, Baja California Sur; Pliocene. Galeocerdo indet. Referred specimen. IGM 6989 (Ferrusquia-Villafranca etal. 1999). Locality and age. Rancho el Jobo, San Juan Formation, Tuxtla Gutierrez, Chiapas; Middle Eocene. Family Hemigaleidae Hasse, 1878 Genus Hemipristis Agassiz, 1843 Hemipristis indet. Referred specimen. IGM 6988 (Ferrusquia-Villafranca etal. 1999). Locality and age. Rancho el Jobo, San Juan Formation, Tuxtla Gutierrez, Chiapas; Middle Eocene. Order Lamniformes Berg, 1958 Family Lamnidae Bonaparte, 1835 Genus Carcharodon Smith, 1938 fCarcharodon auriculatus Jordan, 1923 Referred specimen. IGM 6986 (Ferrusquia-Villafranca etal. 1999). Locality and age. Rancho el Jobo, San Juan Formation, Tuxtla Gutierrez, Chiapas; Middle Eocene. Genus hums Rafinesque, 1810 ^Isurus cf. praecursor Leriche, 1902 Referred specimen. IGM 6985 (Ferrusquia-Villafranca etal. 1999). Locality and age. Rancho el Jobo, San Juan Formation, Tuxtla Gutierrez, Chiapas; Middle Eocene. Family Odontaspididae Muller & Henle, 1839 Genus Carcharias Rafinesque, 1810 Carch arias indet. Referred specimen. IGM 6983 (Ferrusquia-Villafranca etal. 1999). Locality and age. Rancho el Jobo, San Juan Formation, Tuxtla Gutierrez, Chiapas; Middle Eocene. Genus Odontaspis Agdi^siz, 1838 Odontaspis indet. Referred specimen. IGM 6984 (Ferrusquia-Villafranca etal. 1999). Locality and age. Rancho el Jobo, San Juan Formation, Tuxtla Gutierrez, Chiapas; Middle Eocene. Genus "fStriatolamia Glikman, 1964 ■\Striatolamia macrota (Agassiz, 1843) Referred specimen. IGM 6982 (Ferrusquia-Villafranca etal. 1999). Locality and age. Rancho el Jobo, San Juan Formation, Tuxtla Gutierrez, Chiapas; Middle Eocene. Family tOtodontidae Glikman, 1964 fOtodontidae indet. Referred specimen. IGM 6987 (Ferrusquia-Villafranca etal. 1999). Locality and age. Rancho el Jobo, San Juan Formation, Tuxtla Gutierrez, Chiapas; Middle Eocene. zse.pensoft.net 432 Kleyton Magno Cantalice et al.: Fossil fishes in the Mexican National Collection of Paleontology Order Orectolobiformes Compagno, 1973 Suborder Oreetoloboidei Regan, 1908 Family Ginglymostomatidae Gill, 1862 Genus Nebrius Riippel, 1837 Nebrius indet. Referred specimen. IGM 6981 (Ferrusqula-Villafranca etal. 1999). Locality and age. Rancho el Jobo, San Juan Formation, Tuxtla Gutierrez, Chiapas; Middle Eocene. Superorder Batoidea Compagno, 1973 Order Rhinopristiformes Last, Seret & Naylor, 2016 Family incertae sedis Genus ’\Tlalocbatos Brito, Villalobos-Segura & Alvarado-Ortega, 2019 ^Tlalocbatos applegatei Brito, Villalobos-Segura & Alvarado-Ortega, 2019 Referred specimens. IGM 5853 (holotype). Locality and age. Tlayua quarry, Tlayua Formation, Te- pexi de Rodriguez, Puebla; Cretaceous (Albian). Megaelass Osteiehthyes Huxley, 1880 Class Aetinopterygii Woodward, 1891 Subelass Neopterygii Regan, 1923 Order incertae sedis Family incertae sedis Genus "fCipactlichthys Brito & Alvarado- Ortega, 2013 ^Cipactlichthys scutatus Brito & Alvarado-Ortega, 2013 Referred specimens. IGM 6605 (holotype), IGM 6606 (paratype). Locality and age. Tlayua quarry, Tlayua Formation, Te- pexi de Rodriguez, Puebla; Cretaceous (Albian). Order fAspidorhynehiformes Bleeker, 1859 Family fAspidorhynehidae Bleeker, 1859 Genus "fYinctifer Jordan, 1919 ^Vinctiferferrusquiai Cantalice, Alvarado-Ortega & Brito, 2018 Referred specimen. IGM 8873 (holotype). Locality and age. Llano Yosobe, Sabinal Formation, Tlaxiaco, Oaxaca; Jurassic (Kimmeridgian-Tithonian). Order tPyenodontiformes Berg, 1937 Suborder tPyenodontoidei Nursall, 1966 Family tPyenodontidae Agassiz, 1833 fPycnodontidae indet. Referred specimen. IGM 3143 (Carranza-Castaneda and Applegate 1994). Locality and age. Cerro los Mendoza, El Doctor For¬ mation, Zimapan, Hidalgo; Cretaceous (Albian-Ceno- manian). Subfamily tPyenodontinae (Agassiz, 1833) Genus '^Pycnodus Agassiz, 1833 ^Pycnodus indet. Referred specimen. IGM 4551 (Alvarado-Ortega et al. 2015). Locality and age. Belisario Dominguez quarry, Teneja- pa-Eacandon geological unity; Salto de Agua, Chiapas; Paleocene (Danian). Genus "fTepexichthys Applegate, 1992 fTepexichthys aranguthyorum 1992 Referred specimens. IGM 3286 (holotype), IGM 3288- IGM 3289, IGM 3291-IGM 3300, IGM 3455, IGM 3513, IGM 3587, IGM 3689, IGM 3690, IGM 4052-IGM 4122 (paratypes). Locality and age. Tlayua quarry, Tlayua Formation, Te- pexi de Rodriguez, Puebla; Cretaceous (Albian). Suborder tGyrodontoidei Nursall, 1996 Family tGyrodontidae Berg, 1940 Genus "^Gyrodus Agassiz, 1833 •fGyrodus indet. Referred specimens. IGM 9318, IGM 9319 (Alvara¬ do-Ortega et al. 2014). Locality and age. Llano Yosobe, Sabinal Formation, Tlaxiaco, Oaxaca; Jurassic (Kimmeridgian-Tithonian). zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 429-452 433 Infraclass Holostei Miiller, 1845 Division Ginglymodi Cope, 1871 Order tSemionotiformes Arambourg & Bertini, 1958 (sensu Lopez-Arbarello 2012) Family tSemionotidae Woodward, 1890 Genus "fTlayuamichin Lopez-Arbarello & Alvarado-Ortega, 2011 'fTlayuamichin itztli Lopez-Arbarello & Alvarado- Ortega, 2011 Family Lepidotidae Owen, 1860 Genus Scheenstia Lopez-Arbarello & Sferco, 2011 Scheenstia indet. Referred specimen. IGM 9320 (Alvarado-Ortega et al. 2014). Locality and age. La Lobera, “Caliza con Cidaris'\ Tlax- iaco, Oaxaca; Jurassic (Oxfordian-Early Kimmeridgian). Referred specimens. IGM 6716 (holotype), IGM 6717- IGM 6720 (paratypes). Locality and age. Tlayua quarry, Tlayua Formation, Te- pexi de Rodriguez, Puebla; Cretaceous (Albian). Family tMacrosemiidae Wagner, 1860 Genus "fNotagogus Agassiz, 1843 ^Notagogus novomundi Gonzalez-Rodriguez & Reynoso, 2004 Referred specimen. IGM 8172 (holotype), IGM 8173- IGM 8181 (paratypes). Locality and age. Tlayua quarry, Tlayua Formation, Te- pexi de Rodriguez, Puebla; Cretaceous (Albian). Genus "fMacrosemiocotzus Gonzalez- Rodriguez, Applegate & Espinosa- Arrubarrena, 2004 ■fMacrosemiocotzus americanus Gonzalez-Rodriguez, Applegate & Espinosa-Arrubarrena, 2004 Superfamily Lepisosteoidea Lopez- Arbarello, 2012 Family Lepisosteidae Agassiz, 1832 Lepisosteidae indet. Referred specimens. IGM 7657-IGM 7662 (Rodriguez De la Rosa and Cevallos-Ferriz 1998). Locality and age. El Pelillal, Cerro del Pueblo Forma¬ tion, Coahuila; Cretaceous (Campanian). Genus "fAhanulepisosteus Brito, Alvarado- Ortega & Meunier, 2017 ^Nhanulepisosteus mexicanus Brito, Alvarado-Ortega & Meunier, 2017 Referred specimens. IGM 4898 (holotype), IGM 4899- IGM 4902 (paratypes). Locality and age. Llano Yosobe, Sabinal Formation, Tlaxiaco, Oaxaca; Jurassic (Kimmeridgian-Tithonian). Referred specimens. IGM 8163 (holotype), IGM 8164- IGM 8171 (paratypes). Locality and age. Tlayua quarry, Tlayua Formation, Te- pexi de Rodriguez, Puebla; Cretaceous (Albian). Order Lepisosteiformes Hay, 1929 Lepisosteiformes indet. Division Haleeomorphi Cope, 1872 Order flonoseopiformes Grande & Bemis, 1998 Family flonoseopidae Lehman, 1966 Genus '\Quetzalichthys Alvarado-Ortega & Espinosa-Arrubarrena, 2008 fQuetzalichthys Alvarado-Ortega & Espinosa-Arrubarrena, 2008 Referred specimens. IGM 9321, IGM 9322 (Alvara- Referred specimen. IGM 8592 (holotype), IGM 8593- do-Ortega et al. 2014). IGM 8596 (paratypes). Locality and age. Llano Yosobe, Sabinal Formation, Locality and age. Tlayua quarry, Tlayua Formation, Te- Tlaxiaco, Oaxaca; Jurassic (Kimmeridgian-Tithonian). pexi de Rodriguez, Puebla; Cretaceous (Albian). zse.pensoft.net 434 Kleyton Magno Cantalice et al.: Fossil fishes in the Mexican National Collection of Paleontology Family fOphiopsidae Bartram, 1975 Genus "fTeoichthys Applegate, 1988 fTeoichthys kallistos Applegate^ 1988 Referred specimen. IGM 3460 (holotype), IGM 4126 (paratype). Locality and age. Tlayua quarry, Tlayua Formation, Te- pexi de Rodriguez, Puebla; Cretaceous (Albian). ^Teoichthys brevipina Machado, Alvarado-Ortega, Machado & Brito, 2013 Referred specimens. IGM 6741 (holotype), IGM 6742 and IGM 6744 (paratypes), IGM 6604, IGM 6743, IGM 6745-IGM 6747 (Machado et al. 2013). Locality and age. Tlayua quarry, Tlayua Formation, Te- pexi de Rodriguez, Puebla; Cretaceous (Albian). Order Amiiformes Hay, 1929 Family Amiidae Bonaparte, 1837 Subfamily tVidalamiinae Grande & Bemis, 1998 Genus "fPachyamia Chalifa & Tehernov, 1982 ^Pachyamia mexicana Grande & Bemis, 1998 Referred specimens. IGM 7379 (holotype), IGM 7380- IGM 7387 (paratypes). Locality and age. Tlayua quarry, Tlayua Formation, Te- pexi de Rodriguez, Puebla; Cretaceous (Albian). Genus ^Melvius Bryant, 1987 ^Melvius indet. Referred specimens. IGM 7663, IGM 7664 (Rodriguez De la Rosa and Cevallos-Ferriz 1998). Locality and age. El Pelillal, Cerro del Pueblo Forma¬ tion, Coahuila; Cretaceous (Campanian). Infraelass Teleostei Muller, 1845 Order tPholidophoriformes Wagner, 1860 Family tPleuropholidae Saint-Seine, 1949 Genus "fPleuropholis Egerton, 1858 ^Pleuropholis Alvarado-Ortega & Brito, 2016 Referred specimens. IGM 4733 (holotype), IGM 4734, IGM 4735, IGM 9323 (paratypes). Locality and age. Llano Yosobe, Sabinal Formation, Tlaxiaco, Oaxaca; Jurassic (Kimmeridgian-Tithonian). Order tiehthyodeetiformes Bardaek & Sprinkle, 1969 flchthyodectiformes indet. Referred specimen. IGM 9048 (Alvarado-Ortega et al. 2007). Locality and age. Tlayua quarry, Tlayua Formation, Te- pexi de Rodriguez, Puebla; Cretaceous (Albian). Suborder tiehthyodeetoidei Maisey, 1991 Family flehthyodeetidae Crook, 1892 Genus "fUnamichthys Alvarado-Ortega, 2004 'fUnamichthys espinosai Alvarado-Ortega, 2004 Referred specimens. IGM 8373 (holotype), IGM 8374- IGM 8376 (paratypes). Locality and age. Tlayua quarry, Tlayua Formation, Te- pexi de Rodriguez, Puebla; Cretaceous (Albian). Megaeohort Elopoeephalai Arratia, 1999 Cohort Elopomorpha Greenwood, Rosen, Weitzman & Myers, 1966 Order Anguilliformes Goodrieh, 1909 Anguilliformes indet. Referred specimen. IGM 4547 (Alvarado-Ortega et al. 2015). Locality and age. Belisario Dominguez quarry, Teneja- pa-Lacandon geological unity, Salto de Agua, Chiapas; Paleocene (Danian). Megaeoehort Osteoglossoeephalai Betaneur-R., Broughton, Wiley, Carpenter, Lopez, Holeroft, Areila, Saneiangeo, Cureton, Zhang, Borden, Rowley, Reneau, Hough, Lu, Grande, Arratia & Orti, 2013 (=Osteoglossoeephala xewxw Arratia 1999) Supereorhort Osteoglossomorpha Greenwood, Rosen, Weitzman & Myers, 1966 Order Osteoglossiformes Berg, 1940 Suborder Osteoglossoidei Regan, 1909 Eamily Osteoglossidae Bonaparte, 1832 Subfamily Osteoglossinae Nelson, 1968 Genus '\Phaerodus Leidy, 1873 'fPhaerodus indet. Referred specimen. IGM 4549 (Alvarado-Ortega et al. 2015). zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 429-452 435 Locality and age. Division del Norte quarry, Teneja- pa-Lacandon geological unity, Palenque, Chiapas; Paleo- cene (Danian). Supercohort Clupeocephala Patterson & Rosen, 1977 Order tCrossognathiformes Taverne, 1989 Suborder tPachyrhizodontoidei Forey, 1977 fPachyrhizodontoidei indet. Referred specimen. IGM 9049 (Alvarado-Ortega et al. 2007). Locality and age. Tlayua quarry, Tlayua Formation, Te- pexi de Rodriguez, Puebla; Cretaceous (Albian). Family incertae sedis Genus "^Michin Alvarado-Ortega, Mayrinck & Brito, 2008 •fMichin csernai Alvarado-Ortega, Mayrinck & Brito, 2008 Referred specimens. IGM 9028 (holotype), IGM 9029- IGM 9033 (paratypes). Locality and age. Tlayua quarry, Tlayua Formation, Te- pexi de Rodriguez, Puebla; Cretaceous (Albian). Cohort Otomorpha Wiley & Johnson, 2010 Subcohort Clupei Wiley & Johnson, 2010 Order tEllimmichthyiformes Grande, 1982 Family tParaclupeidae Chang & Chou, 1974 fParadupea seilacheri Alvarado-Ortega & Melgarejo-Damian, 2017 Referred specimens. IGM 4717 (holotype), IGM 4718- IGM 4723 (paratypes). Locality and age. Tlayua quarry, Tlayua Formation, Te- pexi de Rodriguez, Puebla; Cretaceous (Albian). Order Clupeiformes Rahnesque, 1810 Suborder Clupeoidei Rahnesque, 1810 Family incertae sedis ^Ranulfoichthys Alvarado-Ortega, 2014 Referred specimens. IGM 9034 (holotype), IGM 9467, IGM 9468 (paratypes); IGM 9035-IGM 9047 (Alvara¬ do-Ortega, 2014). Locality and age. Tlayua quarry, Tlayua Formation, Te- pexi de Rodriguez, Puebla; Cretaceous (Albian). Family Clupeidae Cuvier, 1817 Clnpeidae indet. Referred specimen. IGM 4548 (Alvarado-Ortega et al. 2015). Locality and age. Division del Norte quarry, Teneja- pa-Lacandon geological unity, Palenque, Chiapas; Paleo- cene (Danian). Subcohort Ostariophysi Lord, 1922 Section Otophysa (=Series Otophysi sensu Rosen & Greenwood, 1970) Superorder Cypriniphysae Fink & Fink, 1981 Order Cypriniformes Rahnesque, 1810 Superfamily Cobitoidea Swainson, 1839 Family Catostomidae Agassiz, 1850 Subfamily Ictiobinae Smith, 1992 Genus Ictiobus Rahnesque, 1820 ffictiobus aguilerai Alvarado-Ortega, Carranza- r Castaneda & Alvarez-Reyes, 2006 Referred specimens. IGM 8444 (holotype), IGM 8445- IGM 8591 (paratypes). Locality and age. La Cementera, La Viga, Tecalco, and El Hoyo, Tarango Formation, Tula de Allende, Hi¬ dalgo; Pliocene. Order Siluriformes Rahnesque, 1810 Suborder Siluroidei Rahnesque, 1810 Superfamily B agroidea Bleeker, 1858 Family Ariidae Bleeker, 1858 Ariidae indet. Referred specimen. IGM 5318, IGM 5319 (Hernan- dez-Junquera 1977). Locality and age. Laguna de la Media Luna, Rio Verde, San Luis Potosi; Pleistocene. Cohort Euteleosteomorpha Greenwood, Rosen, Weitzman & Myers, 1966 Subcohort Neoteleostei Nelson, 1969 Infracohort Eurypterygia Rosen, 1973 Section Ctenosquamata Rosen, 1973 Subsection Acanthomorphata Rosen, 1973 Division Acanthopterygii Rosen & Patterson, 1969 Subdivision Percomorphaceae Betancur-R., Broughton, Wiley, Carpenter, Lopez, Holcroft, zse.pensoft.net 436 Kleyton Magno Cantalice et al.: Fossil fishes in the Mexican National Collection of Paleontology Arcila, Sanciangco, Cureton, Zhang, Borden, Rowley, Reneau, Hough, Lu, Grande, Arratia & Orti, 2013 (=Pereomorphaeea sensu Wiley & Johnson, 2010) Percomorphaceae indet. Referred specimen. IGM 7968 (Cantalice and Alvara- do-Ortega in press). Locality and age. Ixtapa locality, Ixtapa Formation, Ixta- pa, Chiapas; Miocene. Genus "fKelemeJtubus Cantaliee & Alvarado- Ortega, 2017 ^Kelemejtubus castroi Cantalice & Alvarado- Ortega, 2017 Referred specimens. IGM 4864 (holotype), IGM 4865- IGM 4867, IGM 4908, IGM 4909 (paratypes). Locality and age. Belisario Dominguez and Division del Norte quarries, Tenejapa-Lacandon geological unity, Salto de Agua and Palenque, Chiapas; Paleocene (Dani- an). Series Syngnatharia Betaneur-R., Wiley, Bailly, Miya, Leeointre et ah, 2014 Order Syngnathiformes Berg, 1940 Suborder Syngnathoidei Regan, 1909 Superfamily Aulostomoidea Greenwood, Rosen, Weitzman & Myers, 1966 Family fEekaulostomidae Cantaliee & Alvarado-Ortega, 2016 Genus '\Eekaulostomus Cantaliee & Alvarado- Ortega, 2016 ^Eekaulostomus cuevasae Cantalice & Alvarado- Ortega, 2016 Referred specimen. IGM 4716 (holotype). Locality and age. Belisario Dominguez quarry, Teneja¬ pa-Lacandon geological unity, Salto de Agua, Chiapas; Paleocene (Danian). Series Carangaria Betaneur-R., Broughton, Wiley, Carpenter, Lopez, Holeroft, Areila, Saneiangeo, Cureton, Zhang, Borden, Rowley, Reneau, Hough, Lu, Grande, Arratia & Orti, 2013(=Carangimorpha sensu Li et al. 2009) Order Istiophoriformes Betaneur-R., Broughton, Wiley, Carpenter, Lopez, Holeroft, Areila, Saneiangeo, Cureton, Zhang, Borden, Rowley, Reneau, Hough, Lu, Grande, Arratia & Orti, 2013 Family Istiophoridae Rahnesque, 1815 Istiophoridae indet. Referred specimens. IGM 7885-IGM 7887, IGM 7890- IGM 7892, IGM 7894 (Fierstine et al. 2001). Locality and age. La Angostura and Rancho Algodones, Trinidad Formation, Baja California Sur; Upper Miocene. Genus LaeepMe, 1802 Makaira nigricans Lacepede, 1802 Referred specimens. IGM 7882-IGM 7884, IGM 7888, IGM 7889, IGM 7893 (Fierstine et al. 2001). Locality and age. La Angostura, Los Dientes Grandes, Canada de En medio, and Rancho Algodones, Trinidad Formation, Baja California Sur; Upper Miocene. Series Ovalentaria Wainwright, Smith, Priee, Tang, Sparks, Ferry, Kuhn, Eytan & Near, 2012 Superorder Atherinomorphae Betaneur-R., Wiley, Arratia, Aeero, Baily, Miya, Leeointre & Orti, 2017 (=Atherinomorpha sensu Greenwood et al. 1996) Order Cyprinodontiformes Berg, 1940 Cyprinodontiformes indet. (sensu Espinosa-Perez et al. 1991) Referred specimen. IGM 7967 (Cantalice and Alvara¬ do-Ortega in press) Locality and age. Los Ahuehuetes, Pie de vaca Forma¬ tion, Tepexi de Rodriguez, Puebla; Oligocene. Suborder Cyprinodontoidei Dyer & Chernoff, 1996 Family Goodeidae Jordan & Gilbert, 1883 _ r Genus "fTapatia Alvarez & Arriola-Longoria, 1972 r ■\Tapatia occidentalis Alvarez & Arriola-Longoria, 1972 Referred specimen. IGM 7966 (Cantalice and Alvara¬ do-Ortega in press). Locality and age. Barranea de Santa Rosa, Amatitan, Jalisco; Pliocene. zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 429-452 437 Series Euperearia Saneiangeo, Carpenter & Betaneur-R., 2016 Order Pereiformes Rafinesque, 1810 Suborder Serranoidei Imamura & Yabe, 2002 Family Serranidae Swainson, 1839 Genus Pale os err anus Cantaliee, Alvarado- Ortega & Alaniz-Galvan, 2018 ^Paleoserranus lakamhae Cantaliee, Alvarado-Ortega & Alaniz-Galvan, 2018 Referred specimens. IGM 4550 (holotype), IGM 9469- IGM 9477 (paratypes). Locality and age. Belisario Dominguez and Division del Norte quarries, Tenejapa-Lacandon geological unity, Salto de Agua and Palenque, Chiapas; Paleocene (Dani- an). 2. CNP-UNAM fossil fish loealities The fossil fishes catalogued into the Type Collection of CNP-UNAM are from 32 paleontological localities be¬ longing to four undefined geological units (Fig. 1; Suppl. material 1), seven marine formations, and five freshwa¬ ter formations. The oldest strata found is from Oxfordian “Caliza con Cidaris” geological unit, Oaxaca, while the youngest fish fossil beds are from the Pleistocene of La¬ guna de Media Luna (San Luis Potosi), the Tarango For¬ mation, near Tula (Hidalgo), and Pie de Vaca, near Tepexi de Rodriguez (Puebla) (Fig. 2). This range represents ap¬ proximately the last 150 million years of Earth’s history. Below are the main features of these Mexican lithostrati- graphic units. The Tirabuzon Formation This geological unit was first known as the Gloria For¬ mation and based on an outcrop exposed few kilometers away from Santa Rosalia town, Baja California Sur (Wil¬ son 1948). Given that this last name was pre-occupied for a unit of Jurassic rocks from Coahuila (Imlay 1936), Applegate and Espinosa-Arrubarrena (1981) suggested the name of Canada Gloria Formation for the rocks of Baja California Sur; however, the name of this geologi¬ cal unit changed to Tirabuzon Formation (Carreno 1981), based on Wilson (1948) and on nomenclatural incongru¬ ences present in the previous suggestions. The stratotype of the Tirabuzon Formation is inside the Bole Basin, at Santa Rosalia, Baja California Sur (Wilson 1948; Car¬ reno 1981) and is a discontinuous basal conglomerate covered with potentially fossiliferous marine sandstone sediments that are overlaid with conglomerate strata containing a gradual lateral transition to littoral, deltaic, and nonmarine facies (Carreno 1981; Ortlieb and Colleta 1984; Carreno and Smith 2007). The abundant fossil con¬ tent in this formation includes ichnofossils, foraminifers. 30 25 20 15 1 IGM-loc 81 (Fm Tirabuzon) 2 IGM-loc 95 (Fm Trinidad) 3 IGM-loc 93 (Fm Trinidad) 4 IGM-loc 3954 (Fm ND) 5 IGM-loc 3404 (Fm Cerro del Pueblo) 6 IGM-loc 1431 (Fm ND) 7 IGM-loc 3530-3533 (Fm Tarango) 8 IGM-loc 370, 1970-1971,1995, 2280, 2432, 2513, 2774, 2776-2777, 2781,2828 (Fm Tlayua) 9 IGM-loc 2435 (Fm Pie de Vaca) 10 IGM-loc 3907 (Fm El Doctor) 11 IGM-loc 3871 (Fm Sabinal) 12 IGM-loc 3872 (’’Calizas con Cidaris") 13 IGM-loc 3302 (Fm San Juan) 14 IGM-loc 3981 (Fm Ixtapa) 15 IGM-loc 3869-3870 (Fm Tenejapa-Lacandon) -115 -110 -105 -100 -95 -90 Figure 1. Map of Mexico, showing the localities currently catalogued at CNP-UNAM. zse.pensoft.net 438 Kleyton Magno Cantalice et al.: Fossil fishes in the Mexican National Collection of Paleontology o Quaternary Holocene Upper Middle Pleistocene Calabrian Gelasian Neogene Pliocene Piacenzian Zanclean Miocene Messinian Tortonian Serravallian Langhian o N O c (D o Burdigalian Aquitanian Paleogene Oligocene Chattian Rupelian Eocene Priabonian Bartonian Lutetian Ypresian Paleocene Thanetian Selandian Danian Cretaceous Upper Maastrichtian Campanian Santonian Coniacian Turonian Cenomanian Lower Albian Aptian Barremiian o Hauterivian Valanginian o N O W 0 Berriasian Upper Tithonian Kimmeridgian Oxfordian Calinvian ■w Jurassic Middle Bathonian Bajocian Aalenian Lower Toarcian Pliensbachian Sinemurian Hettanaian ! Fm ND Fm Trinidad Fm Tirabuzon, Fm Tarango Fm ND Fm Ixtapa Fm Pie de Vaca Fm San Juan Fm Tenejapa-Lacandon Fm Cerro del Pueblo Fm El Doctor Fm Tlayua Fm Sabinal “Calizas con Cidaris” Figure 2. Partial chronostratigraphic chart correlating each for¬ mation catalogued in CNP-UNAM that contains fossil fishes and its respective geological age. mollusks, echinoderms and cetaceans (Wilson 1948; Ap¬ plegate 1978; Carreno 1981; Ortlieb and Colleta 1984; Quiroz-Barroso and Perrilliat 1989; Barnes 1998; Car¬ reno and Smith 2007; Shroat-Lewis 2007). The age of the unit is Late Miocene-Early Pliocene based on planktonic foraminifers (Carreno 1981), nannofossils (Ortlieb and Colleta 1984), and the shark fauna (Applegate 1978). however, these outcrops are from both San Jose del Cabo and Los Barriles Basins (Pantoja-Alor and Carrillo-Bravo 1966; Martlnez-Gutierrez and Sethi 1997; Schwennicke et al. 2017). The unit mainly has a fine-grained sandstone, silt- stone, and mudstone but is also composed of gray-green¬ ish, laminated, fine to medium marine sandstone, shale and siltstone, and some diatomite laminate toward the center of the San Jose del Cabo Basin (Martlnez-Gutierrez and Sethi 1997; Schwennicke et al. 2017). The age of the Trin¬ idad formation range from the middle Miocene to upper Pliocene and its depositional environment has three dis¬ tinct types of strata: 1) a basal nearshore-lagoon deposit; 2) a deep marine environment; 3) a high-energy, shallow marine waters, which they relate to inner shelf shoals and bars (Martinez-Gutierrez and Sethi 1997; McCloy 1984; Carreno 1992; Fierstine et al. 2001; Schwennicke et al. 2017). The fossil record of the unit comprises nanoplank¬ tons (Schwennicke et al. 2017), foraminifers and diatomite (McCloy 1984; Carreno 1992), mollusks (Martinez-Guti¬ errez and Sethi 1997) fishes (Fierstine et al. 2001), and trace fossils (Schwennicke et al. 2017). The Cerro del Pueblo Formation Located in Coahuila state, this formation was described at the beginning 20* century (Imlay 1936); however, its formal description as Cerro del Pueblo Formation was lat¬ er (Murray et al. 1962). Belonging to the Difunta Group (Eberth et al. 2004), its stratotype is northeast to Saltillo City, and its outcrops are in several localities in the Parras Basin (Murray et al. 1962). The lithology of the unit is mainly composed of mudstone and sandstone but also of lesser amounts of limonite, conglomerates, and limestone (McBride et al. 1974; Kirkland et al. 2000). The unit contains a vast fossil record, comprising plants (fruiting structures, palm fronds, conifer cones), rudists, bivalves, gastropods, cephalopods, elasmobranchs, bony fishes, dinosaurs, crocodiles, turtles (Kirkland et al. 2000), and insects (Cifuentes-Ruiz et al. 2006). There are seven dis¬ tinct facies, and sediments were laid down in a low coastal plain and shallow marine conditions which were strongly influenced by frequent changes in the relative sea-level or coastal physical processes (Eberth et al. 2004). Hence, the paleoenvironment is a cyclic alternation of marine, estu¬ arine, and freshwater environments (Cifuentes-Ruiz et al. 2006). The age of the formation ranges from uppermost late Campanian to Maastrichtian (Kirkland et al. 2000). The Tarango Formation The Trinidad Formation Pantoja-Alor and Carrillo-Bravo (1966) first described this unit which is also located in Baja California Sur. Its stra¬ totype is near the intersection of the Coyote and La Trini¬ dad Streams, in the western margin of the Coyote Stream; This formation is in the Valley of Mexico (which includes the states of Ciudad de Mexico, Estado de Mexico, and Hi¬ dalgo) and was first proposed based on sediments exposed about 4 km southwest of Mixcoac, Mexico City (Bryan 1948; Ferrusquia-Villafranca et al. 2017). This geological unit is composed of sandstone and poorly cemented con- zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 429-452 439 glomerate, poorly cemented sandstone and interleaved clay, layers of clay, some layers of basalt interspersed with detri- tal units, lightly compacted conglomerate lenses, insulated limestone lenses, thin lenses of volcanic ash and tuff, and caliche nodules tuff, tuff-breccia, fluvial volcanic gravel, and thin pumice layers (Bryan 1948; Cervantes-Medel and Armienta2004; Ferrusquia-Villafrancaetal. 2017). The fos¬ sil record includes ostracods and diatoms (Ferrusquia-Vil- lafranca et al. 2017), bony Ashes (Alvarado-Ortega et al. 2006), and mammals of the genera Equus Linnaeus, 1758, Cnvieronius Osborn, 1923, Mammuthus Brookes, 1828, Sylvilagus Gray, 1867, Canis Linnaeus, 1758, and Bison Smith, 1827 (Castillo-Ceron et al. 1996). The deposition- al environment is a series of fluvial/lacustrine conditions with sandstone crossbedding which indicates deltaic condi¬ tions (Segerstrom 1962; Alvarado-Ortega et al. 2006; Fer- rusquia-Villafranca et al. 2017). Based on the paleofauna, geochronology, and fault systems, the Tarango Formation is currently considered to present a Pliocene-Quaternary age (Castillo-Ceron et al. 1996; Suter et al. 2001). The El Doctor Formation The El Doctor Formation outcrops from the eastern portion of Queretaro state to the western edge of Hidalgo (Wilson et al. 1955). Its type locality is in the north flank, near El Doctor village in the Sierra Gorda, Queretaro, northeast¬ ern Pena de Bernal (Wilson et al. 1955; Segerstrom 1961; Aguirre-Diaz et al. 2013). This unit has a large limestone bank of varied textures with some chert lenses, dolomite interbeds, and shale partings (Segerstrom 1961). Based on lithology, there are four subunits: 1) EaNegra, deposited in the deep of the neritic zone; 2) San Joaquin, with the same sediments of Ea Negra but deposited at a depth and un¬ der storm wave action; 3) Cerro Eadron, a calcareous bank formed in shallow waters; and 4) Socavon, with clastic sediments deposited not far from the input origin (Wilson et al. 1955). The fossil record found in the El Doctor For¬ mation includes rudists (Wilson et al. 1955), miliolids, cor¬ als, oysters, gastropods, ammonites, echinoids (Segerstrom 1962), planktic and benthic foraminifers, radiolarians (Bravo-Cuevas et al. 2009), crustaceans (Feldmann et al. 2007), and Ashes (Carranza-Castaneda and Applegate 1994; Bravo-Cuevas et al. 2009). Its deposits originate on a platform shelf followed by a transitional marine system, including open sea to deep shelf margin, with alternation of neritic and open oceanic waters and, occasionally, influx of near-shore waters, probably during storms (Wilson et al. 1955; Carrillo-Martinez 1981; Bravo-Cuevas et al. 2009). The age of this formation is Albian-Cenomanian (Wilson et al. 1955; Bravo-Cuevas et al. 2009). The Tlayua Formation The stratotype is a few kilometers north of Tepexi de Ro¬ driguez town, Puebla state, eastern Mexico (Pantoja-Alor 1992). There is a subdivision of the Tlayua Formation: 1) the Tower Member, formed by micritic limestones (mudstone to wackestone) with silica concretions and chert lenses; 2) the Middle Member, formed by red litho¬ graphic laminar limestone (mudstone) and chert lenses; and 3) the Upper Member, with layers of dolomite and dolomitic limestone (Pantoja-Alor 1992). The fossil record is preserved in the Middle Member and, by its abundance and exceptional quality of preservation, the Tlayua Formation is currently considered the first fos- si\-Lagerstdtte site found in Mexico (Alvarado-Ortega et al. 2007). The paleontological record are both marine and terrestrial fauna and flora, including rudists (Alencaster 1973), foraminifera, sponges, gorgonians, gastropods, ammonoids (Cantu-Chapa 1987), belemnoids (Seibertz and Buitron 1987), bivalves, arthropods (Feldmann et al. 1998), asteroids (Buitron-Sanchez et al. 2015), holothu- rians (Applegate et al. 1996), ophiuroids. Ashes (Apple- gate 1987; Alvarado-Ortega 2004), lizards, crocodiles, turtles (Reynoso 1997, 2000), pterosaurs (Cabral-Perdo- mo and Applegate 1994), algae, and gymnosperms (Es¬ pinosa-Arrubarrena et al. 1996). The paleoenvironment is a double-enclosed shallow lagoon behind a barrier reef with stagnant, anaerobic, and hypersaline conditions and bounded by semi-arid land, on the other side, by a barrier bordering a deeper, well-oxygenated lagoon (Applegate 1987; Pantoja-Alor 1992; Espinosa-Arrubarrena and Ap¬ plegate 1996). Nevertheless, some influences of an open sea have been proposed (Kashiyama et al. 2004). The age of Tlayua quarry strata ranges between the Aptian and Albian stages in the Early Cretaceous between 125 and 100 Ma (Cantu-Chapa 1987; Seibertz and Buitron 1987; Kashiyama et al. 2004). The Pie de Vaea Formation The outcrops of the Pie de Vaea Formation are in the southern portion of Puebla, a few kilometers northeast of Tepexi de Rodriguez town very close to the Tlayua For¬ mation (Pantoja-Alor 1992). The lithology of the forma¬ tion consists of continental deposits of fluvial-lacustrine and alluvial environments formed by conglomerates, gravel, silt, clay, marl, limestone, travertine, and volca¬ nic rocks (Pantoja-Alor 1992). These are followed by micritic sandstone with siliciclastic bands and intraclasts of flint, limestone, and volcanic rocks (Cabral-Perdomo et al. 2018). The fossil records of the unit are ichnites belonging to birds, camelids, felids, proboscideans, small artiodactyls (Cabral-Perdomo 1995, 2013), and bony Ashes (Gonzalez-Rodriguez et al. 2013; Guzman 2015). Fungus, leaves, leaflets, wood, and fruits of angiosperms are also well conserved in this unit, as well as ostracods and stromatolites (Beraldi-Campesi et al. 2006). The pa- leoenvironmental condition is a tropical paleolake which evolved from a basin with alluvial conditions to shal¬ low, alkaline, and evaporitic lacustrine circumstances, indicating the gradual desertification of the environment zse.pensoft.net 440 Kleyton Magno Cantalice et al.: Fossil fishes in the Mexican National Collection of Paleontology (Beraldi-Campesi et al. 2006). Although the palynolog- ical record indicates an Eocene-Oligocene age for this formation (Martinez-Hernandez and Ramirez-Arriaga 1999), the paleobotanical and ichnological record with associated mammals indicates with more robustness an Oligocene-Pleistocene age (Cabral-Perdomo 1995, 2013; Cabral-Perdomo et al. 2018; Ramirez and Cevallos-Fer- riz 2002). The Sabinal Formation The Sabinal Formation is in northeastern Oaxaca, with its outcrops (Yosobe and Fa Fobera) in the southern portion of Tlaxiaco Basin (Fopez-Ticha 1985; Meneses-Rocha et al. 1994; Alvarado-Ortega et al. 2014). The lithology of the formation consists of a sequence of mudstone and wackestone clay, marl, and dark gray to black bitumi¬ nous shale strata with abundant calcareous concretions arranged in thin laminar layers and showing abundant light oil impregnations (Fopez-Ticha 1985; Felix 1891; Meneses-Rocha et al. 1994; Alvarado-Ortega et al. 2014). The fossil contents of the unit are microfossils (ostra- cods), plants, invertebrates (mostly ammonites), marine reptiles, and bony fishes (Alvarado-Ortega et al. 2014; Barrientos-Fara et al. 2015). The paleoenvironment is as a transitional environment under the marine infiuence (Alvarado-Ortega et al. 2014). A Kimmeridgian-Titho- nian age is attributed to the Sabinal Formation based on the ammonite assemblage (Alvarado-Ortega et al. 2014). The “Caliza con Cidaris” geological unit This Jurassic geological unit was informally named be¬ cause it carries numerous remains of urchins belonging to the genus Cidaris Feske, 1778. Main outcrops of this unit are present between Tlaxiaco and Mixtepec in Oax¬ aca State (Buitron 1970). This includes the outcrops in “Fa Titana” hills, near Tlaxiaco, firstly reported by Felix (1891) (also see Alvarado-Ortega et al. 2014). The nu¬ merous marine invertebrates recovered in the gray marls and limestones interbedded with shales present in this site suggest that its age could extend from the Fate Callovian to the Early Kimmeridgian (Buitron 1970). Besides the first fossil fishes from the Fobera site reported by Alvara¬ do-Ortega et al. (2014), where “Caliza con Cidaris’’’’ sed¬ iments outcrops; the fossils already documented in this geological unit include three bivalves, echinoids, calcare¬ ous sponges, gastropods, annelids, crinoids, a brachiopod (Buitron 1970; Felix 1891). The San Juan Formation Focated to the northwest of Tuxtla Gutierrez in Chiapas state (Ficari 1960; Allison 1967; Ferrusquia-Villafranca 1996), the San Juan Formation have light-brown shales and yellowish-brown fine-grained calcarenite, composed of conglomerates, sandstone, siltstone, limestone, marl, and coquina (Ferrusquia-Villafranca 1996; Perrilliat et al. 2003). The fossil record in the unit contains foraminifers (Ferrusquia-Villafranca 1996; Perrilliat et al. 2003), calcar¬ eous algae, wood, bivalves, corals, annelids, gastropods, nautiloids, bivalves, echinoderms (Perrilliat et al. 2003), crustaceans, bony fishes (Vega et al. 2001), and sharks (Fer¬ rusquia-Villafranca et al. 1999). The paleoenvironments are episodes of shallow, marine waters with high organic productivity and low terrigenous influence, combined with well-oxygenated shallow waters influenced by continental sedimentation and, probably, with marsh conditions from a deltaic lagoon system (Ferrusquia-Villafranca 1996; Perril¬ liat et al. 2003). Based on the foraminifer fossil record, the age assigned to the San Juan Formation is middle Eocene (Ferrusquia-Villafranca 1996; Perrilliat et al. 2003). The Ixtapa Formation Also located in Chiapas, the Ixtapa Formation is 28 km east from Tuxtla Gutierrez City and its outcrops mainly at the east side of the Soyalo-Ixtapa highway (State Road 195) next to the bridge that crosses the Rio Hondo 1 km north of Ixtapa Municipality (Fangenheim and Frost 1963; Fer¬ rusquia-Villafranca 1996). This unit has a sequence of py¬ roclastic materials interbedded with calcitic pebbly grav¬ els and tuffs, which become more frequent towards the base of the unit forming part of interbedded layers of con¬ glomerates, sandstones, and clays where crystalline and calcareous conglomerates are sporadically present (Fan¬ genheim and Frost 1963; Martinez-Hernandez 1992). The fossil record in Ixtapa Formation is diverse and includes charophytes, foraminifers, mollusks (Daily and Durham 1966), palynomorph assemblages (e.g. dinoflagellates and mangrove pollen), proboscides, horses, rhinoceros (Fan¬ genheim and Frost 1963; Daily and Durham 1966), fresh¬ water turtles (Ferrusquia-Villafranca 1996), and only one record of a bony fish (Cantalice and Alvarado-Ortega in press). The Ixtapa Formation was formed under a low-en¬ ergy fluvial-lacustrine conditions over the Middle Mio¬ cene continental sandstones of the Coyolar Formation and is below the Pliocene-Pleistocene volcanic deposits of the Punta de Flano Formation (Ferrusquia-Villafranca 1996; Martinez-Amador et al. 2004; Hernandez-Villalva et al. 2013). The paleoenvironment is a continental lacustrine or a brackish transitional environment near the coast (Daily and Durham 1966; Martinez-Hernandez 1992). The age assigned to this Formation ranges from the Middle to Fate Miocene (Ferrusquia-Villafranca 1996). The Tenejapa-Lacandon geological unity The Tenejapa-Facadon geological unity was first men¬ tioned by Islas-Tenorio et al. (2005). This layer represents the union of two contemporary and laterally continu¬ ous formations: Tenejapa, first described from outcrops of San Cristobal de las Casas City (Quezada-Muneton zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 429-452 441 1987), and Lacandon, primary known from Peten, Gua¬ temala (Vinson 1962; Fourcade et al. 1999). Because it is not possible to determine the boundaries between the two Formations and the proper identity of each unit is poorly understood, here we interpret both formations as a single element, the Tenejapa-Lacandon geological uni¬ ty (see Alvarado-Ortega et al. 2015). This layer outcrops in diverse portions of Chiapas state (Islas-Tenorio et al. 2004, 2005); however, the fossils housed in the CNP are from the Division del Norte and Belisario Dominguez quarries. The first one is approximately 2 km southeast of the archeological site of Palenque City, while the last is 9.5 km from Palenque City. Both strata are limestone marls deposited in laminated and parallel strata, which show yellowish-creamy colors with some dark-grey silicified bands and nodules (Alvarado-Ortega et al. 2015; Cantalice et al. 2018a). The fossil specimens are many poor-preserved impressions of plant remains and a singu¬ lar paleoicthyofauna (Alvarado-Ortega et al. 2015; Can¬ talice and Alvarado-Ortega 2016, 2017; Cantalice et al. 2018a). The number of well-preserved specimens (most¬ ly on massive mortality) makes this geological unity a fossiX-Lagerstdtte site. The paleoenvironment of Teneja¬ pa-Lacandon geological unity is a marine platform with infiuences of external conditions to the west, originating the Tenejapa Formation, and to the east, shallow waters infiuenced by primarily internal conditions composes the Lacandon Formation (Quezada-Muneton 1987; Alvara¬ do-Ortega et al. 2015). Studies based on stable strontium isotopes indicate a Paleocene (Danian) age for this geo¬ logical unity (Alvarado-Ortega et al. 2015). Discussion Since the last systematic review of the vertebrates housed on the CNP-UNAM (Perrilliat 1993), the number of spec¬ imens of fossil fishes has increased from five to 375, and the number of valid species raised from two to 27, in¬ cluding 20 new to science. This means that the number of fossil fishes species housed in the collection is currently 11 times greater than previously, not considering, howev¬ er, the fossils that were are not determinable to species, which represent almost one-half of the palaeoichthyo- logical material currently housed in the type collection. These numbers show a great increment in the knowledge of Mexican fossil fishes over the last two decades. Many of the species housed in CNP-UNAM are the oldest record of its respective group and the first report of the taxon in North America (Table 1). These repre¬ sent not only an increase of the knowledge about the fish diversity but also constitute valuable tools to a new understanding of the historical biogeography of fishes. This is the case of '\Vinctifer ferrusquiai (Fig. 3A), from Kimmeridgian-Tithonian marine deposits from Oaxaca, which is the oldest fossil record and the first report of a member of the genus "^Vinctifer outside the Cretaceous Period (Cantalice et al. 2018b). Its age and distribution suggest that the family Aspidorhynchidae under went a Table 1. Remarks of outstanding Mexican species housed at the CNP-UNAM. Abbreviation; A. America; L. last occurrence; N. North America; O. oldest occurrence. The asterisk means the oldest generic occurrence. Taxon Oldest First Apparent or Last report endemism _ occurrence Class CHONDRICHTHYES Order fHYBODONTIFORMES Family tHYBODONTIDAE ■[Planohybodus indet. Order CARCHARHINIFORMES Family CARCHARHINIDAE ■[Galeocerdo rosaliensis Order RHINOPRISTIFORMES Family incertae sedis ■[TIalocbatos applegatei Class ACTINOPTERYGII Order incertae sedis Family incertae sedis ■[Cipactiichthys scutatus Order tASPIDORHYNCHIFORMES Family fASPIDORHYNCHIDAE t Vinctifer ferrusquiai Order fRYCNODONTIFORMES Family tPYCNODONTIDAE ■[Pycnodus sp. ■[Tepexichthys aranguthyorum Division GINGLYMODI Order fSEMIONOTIFORMES Family fSEMIONOTIDAE ■[Tiayuamichin itztii Family fMACROSEMIIDAE ■[Notagogus novomundi ■[Macrosenniocotzus americanus Order LEPISOSTEIFORMES Family LEPIDOTIDAE Scheenstia sp. Family LEPISOSTEIDAE ■[Nhanuiepisosteus mexicanus Division HALECOMORPHI Order flONOSCOPIFORMES Family flONOSCOPIDAE ■[Quetzaiichthys perriiiiatae Family fOPHIOPSIDAE ■[Teoichthys kaiiistos ■[Teoichthys brevipina Order AM 11 FORMES Family AM 11 DAE ■[Pachyamia mexicana Order tPHOLIDOPHORIFORMES Family tPLEUROPHOLIDAE ■[Pieurophoiis cinerosorum Order flCHTHYODECTIFORMES Family flchthyodectyidae fUnannichthys espinosai Order OSTEOGLOSSIFORMES Family OSTEOGLOSSIDAE ■[Phaerodus indet. Order fCROSSOGNATHIFORMES Family incertae sedis ■\Michin csernai Order fELLIMMICHTHYlFORMES Family tPARACLUPEIDAE ■[Paraciupea seiiacheri Order CLUPEIFORMES Family incertae sedis ■[Ranuifoichthys dorsonudum Order CYPRINIFORMES Family CATOSTOMI DAE ■[Ictiobus aguiierai Division ACANTHOPTERYGII Order incertae sedis Family incertae sedis ■[Keiennejtubus castroi Order SYNGNATHIFORMES Family fEEKAULOSTOMIDAE ■[Eekauiostonnus cuevasae Order CYPRINODONTIFORMES Family GOODEIDAE ■[Tapatia occidentaiis Order PERCIFORMES Family SERRANIDAE ■[Paieoserranus iakamhae 0 * 0 X X 0 A A 0 0 * 0 " 0 0 0 A A X X 0 zse.pensoft.net 442 Kleyton Magno Cantalice et al.: Fossil fishes in the Mexican National Collection of Paleontology Figure 3. Some species housed in the CNP-UNAM type collection. A. 'fVinctifer ferrusquiai Cantalice, Alvarado-Ortega & Brito, 2018; B. ‘\Nhanulepisosteus mexicanus Brito, Alvarado-Ortega & Meunier, 2017; both being the most ancient species of their fam¬ ilies to date; C. "fMacrosemiocotzus americanus Gonzalez-Rodriguez, Applegate & Espinosa-Arrubarrena, 2004, the first report of Macrosemiidae in North America. Scale bars: 10 mm. rapid diversification and had a wide distribution during the Late Jurassic (Cantalice et al. 2018b). Another exam¬ ple is the earliest known lepisosteoid, '\Nhanulepisosteus mexicanus (Fig. 3B), which raises the origin of modern gars to the Late Jurassic (Brito et al. 2017). Furthermore, the Paleocene fossil fishes from Chiapas (Fig. 4) push back the absolute age of origin of many acanthomorph groups (e.g. seabasses and flutemouth fishes) to the early Cenozoic, just after the K/Pg boundary (Late Paleocene, 63 Ma). These finds reveal that the Caribbean Region is an important place for the origin and diversification of some modem ray-finned fish lineages (Alvarado-Ortega et al. 2015; Cantalice and Alvarado-Ortega 2016, 2017; Cantalice et al. 2018a). zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 429-452 443 Figure 4. Paleocene fossil fishes found in marine outcrops of Chiapas housed in CNP-UNAM A. ‘\Eekaulostomus cuevasae Can- talice & Alvarado-Ortega, 2016; B. ‘\Kelemejtubus castroi Cantalice & Alvarado-Ortega, 2017; C. ^Paleoserranus lakamhae Can- talice, Alvarado-Ortega & Alaniz-Galvan, 2018. Scale bars: 10 mm. When included in a phylogenetic context, some spe¬ cies housed in CNP-UNAM offer valuable details that help understanding the morphological and ecological changes that occurred in some groups of fishes through time. One example is the aulostomoid '\Eekaulostomus cuevasae (Fig. 4A), which is considered the stem group of Recent flutemouth fishes and reveals that reduction of body scutes size and enlargement of the snout, trunk, and fin rays are evolutionary trends of aulostomoids (Can¬ talice and Alvarado-Ortega 2017). These morphological modifications are possibly related to improvements in predation in extant species (Cantalice and Alvarado-Or¬ tega 2017). Moreover, including CNP-UNAM species in phylogenetic analyses also provide support to solve some incongruences on fish classification, such as "fQuetza- lichthys and "fTeoichthys (Fig. 5), two genera collected in Tlayua quarry (Puebla), which when included in the phy- logeny of the order lonoscopiformes proved the mono- phyly of the families Ophiopsidae and lonoscopidae (Al¬ varado-Ortega and Espinosa-Arrubarrena 2008). Mexico has been in the tropical region since the Ju¬ rassic, the period of the oldest fossil fish records report- zse.pensoft.net Kleyton Magno Cantalice et al.: Fossil fishes in the Mexican National Collection of Paleontology 444 Figure 5. Cretaceous fossil fishes from Tlaua quarry, Puebla, housed in CNP-UNAM A. ‘\Quetzalichthys perrilliatae Alvarado-Or- tega & Espinosa-Arrubarrena, 2008; B. '\Teoichthys kallistos Applegate, 1988; C. ^Teoichthys brevipina Machado, Alvarado-Orte- ga, Machado & Brito, 2013. Scale bar: 10 mm. ed at CNP to date (Scotese 2014). The 150 million years since the Jurassic to present day could explain the diver¬ sification of several groups of fishes and the richness of possible Mexican endemic fauna (Table 1). Therefore, the continual collection and increases in knowledge of the paleoichthyofauna housed at CNP is fundamental to understand the patterns of fish diversity through the geological ages and highlights the Mexican fossil records as essential to understanding biogeographic patterns and current global fish diversity. zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 429-452 445 Conclusions After 25 years since the last vertebrate catalogue (Per- rilliat 1993), we present the first fossil fish catalogue of the species housed in the type collection of the Colec- cion Nacional de Paleontologia of Universidad Nacional Autonoma de Mexico (CNP-UNAM). The increase of knowledge of fossil fish diversity, as its biological and biogeographical implications, are evidence that main¬ taining a proper collection of the Mexican fossil record is necessary for understanding the complex evolution¬ ary history of fishes. The knowledge of the Mexican pa- leoichthyofauna is emerging. The formal description of many specimens housed in the geographic reference sec¬ tion is still necessary and increasing with periodic field¬ work. Palaeoichthyology is a promising research area in Mexican paleontology. Acknowledgements Our sincere thanks to J. Alvarado-Ortega, the curator of CNP-UNAM. We give thanks to all researchers and members of the Institute de Geologia that helped add to the number of specimens in the CNP-UNAM type col¬ lection over time. We also thank D. Ruiz-Ramoni for the help with preparing the map, L.P Crivano Machado, G.R. de Paula Machado, and J. Miguel Contreras for the photographs, R. Forsyth, L. Cavin, M.R. de Britto, M.E. Bichuette, N. Yonow, and P. Pankov for the review of the manuscript, and to R.O. Roney for his English revision of the manuscript. This research is supported by DGA- PA-PAPIIT project IN209017, UNAM. K.M. Cantalice was supported by the DGAPA postdoctoral fellowship; A. Martinez-Melo was supported by the EPE-CONACYT postdoctoral fellowship. References Agassiz L (1832) Untersuchungen uber die fossilen Fische aus der Li¬ as-Formation. Jahrbuch ftlr Mineralogie, Geognosie, Geologic und Petrefaktenkunde 3: 139-149. Agassiz JLR (1833-1843) Recherches sur les poisons fossils, tome II, contenat I’histoire de I’ordre des ganoides. Neuchatel, Swiss, 336 pp. https;//doi.org/10.5962/bhl .title.4275 Agassiz JLR (1833-1843) Recherches sur les poisons fossils, tome hi, contenant I’histoire de I’ordre des placoides. 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Supplementary material 1 Supplementary Information Authors: Kleyton Magno Cantalice, Alejandra Martinez-Melo, Violeta Amparo Romero-Mayen Data type: locality data Explanation note: Table with essential information about the distinct geological localities containing fishes cata¬ logued in the CNP-UNAM. Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons. org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow us¬ ers to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Link: https://doi.org/10.3897/zse.95.35435.suppll zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 453-463 | DOI 10.3897/zse.95.38259 4>yEnsPFr. BERLIN Taxonomic revision of the genus Hyperaulax Pilsbry, 1897 (Gastropoda, Stylommatophora, Odontostomidae) Rodrigo B. Salvador^ Daniel C. Cavallari^ 1 Museum of New Zealand Te Papa Tongarewa. 169 Tory Street, 6011, Wellington, New Zealand 2 Departamento de Biologia, Faculdade de Filosofia, Ciencias e Letras de Ribeirdo Preto, Universidade de Sdo Paulo. Avenida Bandeirantes 3900, 14040-900, Ribeirdo Preto, SP, Brazil http://zoobank.org/EE9B3C6B-D4AE-4508-BF0E-5CCEEB7F12EA Corresponding author; Rodrigo B. Salvador (salvador.rodrigo.b@gmail.com) Academic editor; FrawA: .^o/z/er ♦ Received 15 July 2019 ♦ Accepted 27 August 2019 ♦ Published 11 September 2019 Abstract The genus Hyperaulax Pilsbry, 1897 comprises two living species endemic to the oceanic Fernando de Noronha Archipelago, off north-eastern Brazil. They are currently allocated in two subgenera, Hyperaulax s. str. and Bonnanius Jousseaume, 1900, belonging to the family Odontostomidae. Herein we present a taxonomic revision of these species, assessing their familiar allocation within Orthalicoidea, offering updated diagnoses and descriptions, figuring the type materials and further relevant specimens, and providing barcoding DNA sequences. We conclude that Bonnanius is a junior synonym of Hyperaulax, which is classified in Odontostomidae. The genus contains two valid species, H. ridleyi and H. ramagei, both endemic to Fernando de Noronha. Key Words Bonnanius, endemic species, Fernando de Noronha, island speciation, Orthalicoidea, Pulmonata Introduction The genus Hyperaulax Pilsbry, 1897 had a unique com¬ position. It comprised two living species endemic to the oceanic Fernando de Noronha Archipelago off Brazil, H. ridleyi (Smith, 1890) and H. ramagei (Smith, 1890), and eight species from Tampa Silex beds (Oligocene) of Flor¬ ida, USA: H. americanus (Heilprin, 1887), H. ballistae (Dali, 1915), H. floridanus (Conrad, 1846), H. heilprin- ianus (Dali, 1890), H. remolinus (Dali, 1915), H. stearnsii (Dali, 1890), 77. tampae (Dali, 1915), and 77. tortilla (DdM, 1915). Suspecting this could not reflect an actual relation¬ ship, the fossil species previously assigned to Hyperaulax were revised by Auffenberg et al. (2015). Those authors concluded that the fossils bore only a superficial similar¬ ity to Hyperaulax and erected the new genus Tocobaga Auffenberg et al, 2015 to house the North American fossil species, which they classified in the Bulimulidae rather than in Odontostomidae. Auffenberg et al. (2015) consid¬ ered most of the fossil forms to be synonymous and rec¬ ognized only three valid species: Tocobaga americanus, T floridanus and T. wakullae Mansfield, 1937 (previously considered a subspecies of T. americanus). The two Recent species are also not without prob¬ lems, as two names are currently considered synonymous with 77. ramagei, moreover, this species is included in the subgenus Bonnanius Jousseaume, 1900, which can sometimes be recognized as a valid genus (e.g., Simone 2006). The peculiar morphological features of Hyper¬ aulax also present some challenges, as it is reminiscent of several lineages of Orthalicoidea. The genus is usually placed in Odontostomidae, but it also bears resemblance to insular Bulimulidae from the Galapagos; furthermore, Hyperaulax had also been assigned to the extinct fam¬ ily Grangerellidae by Henderson (1935), although this author suggested this based on Oligocene material from Florida (now in Tocobaga), not the Recent Brazilian spe¬ cies (for a full discussion see Auffenberg et al. 2015). Therefore, herein we conducted a taxonomic review of the two Recent species of Hyperaulax, assessing their Copyright Rodrigo B. Salvador, Daniel C. Cavallari. This is an open access articie distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the originai author and source are credited. 454 Salvador, R.B. & Cavallari, D.C.: Revision of Hyperaulax familiar allocation within Orthalicoidea and the validity of the subgenus Bonnanius and of the two species syn- onymized with H. ramagei. Furthermore, an updated di¬ agnosis and description, alongside images of the type and additional materials, is provided here for eaeh speeies. Methods Fernando de Noronha is an archipelago located ea 350 km off northeastern Brazil (3°50'-3°52'S, 32°24'-32°28'W) originating from extinct volcanic structures estimated to be 1.8-12.4 Ma old. It comprises the main island of Fer¬ nando de Noronha (17 km^) and over 20 smaller islands and rocks (Fig. 1). The local climate is dry tropical with a mean annual rainfall of 1,300 mm and a mean annual temperature of 25.4 °C. It has strong oeeanie influence, with two well-defined seasons: dry from August to Janu¬ ary and rainy from February to July (Favaro et al. 2006; Marques et al. 2007). The vegetation on the islands has elose aflinities with the Atlantic Rainforest, though only 5% of the original eover remains (Claudino-Sales 2019). Even so, the islands are rieh in endemic species of plants and animals but have a low overall diversity compared to the mainland, which is especially true for terrestrial snails, of which there are three known endemic spe¬ cies (Lopes and Alvarenga 1955). The entire area totals 112.7 km^ ineluding land and sea; it was designated as an environmental protection area by the Brazil in 1989 and became a World Heritage Site in 2001. Nevertheless, the islands are populated and suffer the negative impacts of overtourism and pollution (Claudino-Sales 2019). All the type specimens were analyzed and the main mu¬ seum collections worldwide that could contain material of Hyperaulax were visited or contacted for loans or photo¬ graphs and information of their speeimens. The type ma¬ terials are ill lustrated herein, alongside additional speei- mens to thoroughly show conehological variation. Shell measurements were taken with a digital caliper. SEM images of the protoconchs were obtained at the Staatli- ches Museum fur Naturkunde Stuttgart (SMNS; Stuttgart, Germany). The material studied in the present work is housed in the following collections: ANSP, Academy of Natu¬ ral Sciences of Drexel University (Philadelphia, USA); MNHN, Museum national d’Histoire naturelle (Paris, France); MNZ, Museum of New Zealand Te Papa Ton- garewa (Wellington, New Zealand); MZSP, Museu de Zoologia da Universidade de Sao Paulo (Sao Paulo, Bra¬ zil); NHMUK, Natural History Museum (London, UK); USNM, Smithsonian Institution National Museum of Natural History (Washington, DC, USA); ZMB, Museum fur Naturkunde, Leibniz Institute for Evolution and Bio- zse.pensoft.net Zoosyst. Evol. 95(2) 2019, 453-463 455 diversity Science (Berlin, Germany); ZSM, Zoologische Staatssammlung Miinchen (Munich, Germany). The following abbreviations are used herein for shell dimensions: H, shell height (parallel to coiling axis); D, greatest shell width (perpendicular to H); h, aperture height (maximum length parallel to aperture plane); d, aperture width (maximum width parallel to aperture plane); W, number of whorls of shell (approximated to closest quarter); w = number of whorls of protoconch (ap¬ proximated to closest quarter). Two adult specimens of H. ridleyi from lot MZSP 89940 had a fraction of their foot clipped for molecular study. No specimen of H. ramagei with preserved soft parts is known. Given our suspected systematic affinity of Hyperaulax, we also sequenced a specimen of Tomigerus corrugatus Iher- ing, 1905 (lot MZSP 43077). DNA extraction was carried out with QIAGEN DNeasy Blood & Tissue Kit, standard protocol. We targeted the barcoding fragment of the mito¬ chondrial COI gene (primers ECO and HCO of Folmer et al. 1994), with circa 650 bp. The PCR protocol was set as: (1) initial denaturation at 96 °C (2 minutes); (2) denatura- tion at 94 °C (30 seconds); (3) annealing at 48 °C (1 min¬ ute); (4) extension at 72 °C (2 minutes); (5) repeat steps (2) to (4) 34 times, for a total of 35 cycles; (6) final extension at 72 °C (5 minutes). PCR products were quantified via agarose gel electrophoresis, cleaned with ExoSAP-IT^m (Afiymetrix Inc.), and Sanger sequenced. The sequences were assembled and quality-checked in Geneious Prime (version 2019.0.3, Biomatters Etd), and uploaded to NCBI GenBank under the accession numbers MNl75954 and MNl75955 {H. ridleyi) and MNl75956 (T. corrugatus). Additional orthalicoid COI sequences were obtained from GenBank, originating from the work of Breure and Romero (2012; see Appendix 1 for accession numbers). We excluded some species from those authors’ dataset, namely those with uncertain identification, incomplete sequences, and the basal-most taxa (e.g., Bothriembryontidae, Me- gaspiridae), which could bring too much noise to the anal¬ ysis. We used one Planorbidae as outgroup (sequence from GenBank, see Appendix 1). A total of 35 species were used from that work; sequences were 654 bp long, with the ex¬ ception of the outgroup, with 669 bp due to an indel. All sequences were aligned in Geneious Prime with the MUSCEE plugin (Edgar 2004; default settings, accu¬ racy-optimized) and further proofed manually. A tree was built in Geneious Prime by Bayesian Inference (MrBayes plugin; Huelsenbeck and Ronquist 2001; Ronquist and Huelsenbeck 2003); settings: HKY85 substitution model, 200,000 burn-in length, 1,100,000 iterations. Systematics Superfamily Orthalicoidea Family Odontostomidae Pilsbry & Vanatta, 1898 Genus Hyperaulax Pilsbry, 1897 Bulimulus {Hyperaulax) Pilsbry 1897a: 10; Pilsbry 1897b: 82. Bonnanius 1900: 39; Schileyko 1999: 337. Hyperaulax-. Pilsbry 1901: 102; Wenz 1923: 729; Thiele 1931: 660; Henderson 1935: 145; Morretes 1949: 153; Oliveiraetal. 1981: 350; Parkinson et al. 1987: 29; Schileyko 1999: 320; Salgado and Coelho 2003: 165; Simone 2006: 178; Salvador 2019: 87. Hyperaulax {Bonnanius)’. Pilsbry 1901: 103; Thiele 1931: 661; Mor¬ retes 1949: 153; Zilch 1960: 505; Breure 1974: 52; Oliveira et al. 1981: 350. Hyperaulax {Hyperaulax)-. Thiele 1931: 661; Morretes 1949: 153; Zilch 1960: 505; Breure 1974: 109. Tomigerus {Bonnanius)-. Parodiz 1962: 453. Bonnarius [sic]: Simone 2006: 178. Type species. Bulimus (Bulimulus) ridleyi Smith, 1890, by original designation. Included species. Hyperaulax ridleyi (Smith, 1890) and H. ramagei (Smith, 1890). Diagnosis. Shell bulimoid. Protoconch sculptured by sinuous axial riblets, which can anastomose and fade on abapical region. Umbilicus surrounded by a periumbilical spiral angulation. Description. Shell small to medium-sized, bulimoid, with ca 4-5 convex whorls; ground color cream, ochre or brown, with \-A lighter-colored spiral bands on lat¬ eral portion of whorls; periumbilical region completely or marginally discolored, whitish; apex (especially proto¬ conch) usually of lighter color. Suture well-marked. Pro¬ toconch sculptured by numerous fine sinuous axial rib- lets, transition unclear. Teleoconch overall smooth except for axial growth lines. Aperture ovoid; peristome white, reflected, thickened, and continuous, with 0^ apertural teeth. Umbilicus perforate, well marked. Remarks. After Auffenberg et al. (2015) removed all fos¬ sil taxa from Hyperaulax, the genus was left only with two living species, Hyperaulax ridleyi (Fig. 2) and 77. ramagei (Figs 3,4), with the latter classified in the subgenus or full genus Bonnanius. Both species are known only from Fer¬ nando de Noronha Archipelago off north-eastern Brazil. The genus Bonnanius is considered here synony¬ mous with Hyperaulax as there are no diagnostic char¬ acters allowing its clear separation other than its larger shell size. The presence of teeth in the aperture of 77. ramagei (previously classified in Bonnanius) could be used as a diagnostic genus-level character; however. zse.pensoft.net 456 Salvador, R.B. & Cavallari, D.C.: Revision of Hyperaulax it is well known that odontostomid genera show great inter- and intraspecific variation in the presence and strength of apertural teeth. Moreover, some specimens of H. ridleyi do show a palatal tooth (Fig. 2A) similar in position and length to that of H. ramagei. Further¬ more, for a genus with only two species, keeping them separated into two distinct subgenera is excessively zealous taxonomy. As for the other conchological char¬ acters, Bonnanius share all of them with Hyperaulax, as discussed below. The protoconch sculpture has always been deemed a good character to define genera in Orthalicoidea and has more recently received large support from molecular studies (Breure and Romero 2012). The protoconchs of Hyperaulax and Bonnanius are very similar and indicate a close affinity between the two forms: same number of whorls (ca PA); same sculpture pattern (sinuous axial riblets, more clearly separate on adapical area of whorl, but anastomosing on abapical area and sometimes fad¬ ing into scattered dots). There are also some differences on the protoconch, but nothing that would suggest two distinct genera (many genera of Orthalicoidea bear some minor differences in their protoconchs, which helps with species diagnosis; e.g., Salvador and Cavallari 2013; Sal¬ vador and Simone 2016). The main difference is that the protoconch of H. ramagei is more flattened and rounded, which makes the riblets a little more spaced; this can be attributed to its shell being larger and wider overall. Fur¬ thermore, the protoconch of H. ridleyi has a raised ridge on its middle region. Finally, the protoconch sculpture of museum specimens of H. ramagei is often faded or erod¬ ed, which has led to claims of a smooth protoconch in the literature (e.g., Parodiz 1962; Abbott 1989). Other conchological characters are very similar in both H. ridleyi and H. ramagei. the roughly pentagonal shape of the aperture and its positioning in relation to the body whorl, the long palatal tooth (absent in most H. ridleyi specimens), the shape of the umbilicus and the perium¬ bilical spiral angulation, the unsculptured teleoconch (except for growth striations), and the periostracum color (brown with at least one white spiral band). The classification of Hyperaulax in Odontostomidae has been well supported in the literature, with just a few different classification schemes. For instance, Schileyko (1999) argued in favor of Bulimulidae because H. rid¬ leyi has no teeth; however, not all odontostomids ac¬ tually have teeth and some specimens of H. ridleyi do show a faint palatal tooth, as discussed above and already remarked by Pilsbry (1901). Moreover, H. ridleyi has the typical channel-like structure on the junction of the parietal and palatal regions of the peristome. Schileyko (1999), however, maintained Bonnanius (and hence H. ramagei) in Odontostomidae. In any event, there are other conchological characters favoring an allocation within Odontostomidae, such as the elevated embryonic whorls (Auffenberg et al. 2015) and the protoconch sculpture (wavy riblets, similar to Plagiodon- tes Doering, 1876; Piza and Cazzaniga 2016). The overall shell shape of Hyperaulax is very similar to Tomigerus Spix, 1827 and Biotocus Salgado & Feme, 1990, but with a different position of the aperture in relation to the body whorl and a different structure of the umbilical region; also, the protoconch of Tomigerus is smooth. The periostracum color is also similar to what is seen in Tomigerus (e.g., T clausus Spix, 1827, T matthewsi Salgado & Feme, 1991, and T rochai Ihering, 1905), but a striped pattern can also be found in species of Moricandia Pilsbry & Vanatta, 1898 and even Anostoma Waldheim, 1807. The dentition of Hyperaulax (mainly of H ramagei) is similar to that of Burringtonia Parodiz, 1944 and also Anctus angiostomus (Wagner, 1827). Finally, the channel-like structure on the junction of the parietal and palatal regions of the peristome is virtually identical to what is observed in some species of Cyclodontina Beck, 1837, Spixia Pilsbry & Vanata, 1898, and AWaldheim, 1807. Our analysis of interspecific affinities using the bar¬ coding region of the COI marker has grouped Hyperaulax (H ridleyi only) with Tomigerus, as expected by our mor¬ phological analysis, with a posterior probability of 0.997. However, these two species were grouped with Simpulop- sis Beck, 1837 (family Simpulopsidae) in our Bayesian tree, instead of being grouped with other odontostomids. This is likely due to the fact that COI alone is not sufficient to solve family-level relationships among stylommato- phoran snails (Breure and Romero 2012), despite being sufficient to capture the relationship of close species-level taxa. Based on morphological data, we retain Hyperaulax (and Tomigerus) in the family Odontostomidae. It is curious that another orthalicoid lineage, from the other side of South America, evolved an uncannily similar shell shape to Hyperaulax: Naesiotus wolfi (Reibisch, 1892), from the Galapagos (lectotype ZMB 47.950, paralectotype NHMUK 1894.6.8.7). Furthermore, N. wolfi is within the size range of Hyperaulax, and has a similar color pattern to H ridleyi, including the median white spiral band. The protoconch, naturally, is different {Naesiotus Albers, 1850 has very fine and well-defined axial striae), alongside other more general shell features: higher spire, different propor¬ tion of body whorl to spire, and a larger number of whorls (ca 614). In any event, this is a remarkable case of conver¬ gent evolution on islands and deserves further investigation. Hyperaulax ridleyi (Smith, 1890) Fig. 2 Bulimus {Bulimulus) Ridleyi E.A. Smith 1890; 501, pi. 30, fig. 9; Dali 1896:415. Bulimulus {Hyperaulax) ridleyi. Pilsbry 1897a: 10; Pilsbry 1897b: 82, pi. 14, figs 11-13. Buliminus ridleyi: Mbllendorlf 1901: 126. Hyperaulax ridleyi: Pilsbry 1901: 103; Ihering 1923: 191; Wenz, 1923: 729; Schileyko 1999: 321, fig. 396; Salgado and Coelho 2003: 165; Simone 2006: 178, fig. 637; Breure and Ablett 2012: 36, figs 20C, D, 20ii; Salvador 2019; 87. zse.pensoft.net Zoosyst. Evol. 95(2) 2019, 453-463 457 Figure 2. Hyperaulax ridleyi. A-E. Lectotype, NHMUK 1888.6.27.106. F-L Paralectotypes #1 to #4 (in order), NHMUK 1888.6.27.107-110. J. Extreme form, large and with thick callus, NHMUK 1888.6.27.88-94. K. Weathered specimen, ZMS unnum¬ bered (ex Heimburg colln.). L. Protoconch detail (same specimen from K). zse.pensoft.net 458 Salvador, R.B. & Cavallari, D.C.: Revision of Hyperaulax Hyperaulax (Hyperaulax) ridleyi: Thiele 1931; 661; Morretes 1949: 153; Zilch 1960: 505, fig. 1771; Breure 1974: 51; Breure and Schouten 1985: 4. Hyperaulax (s. str.) ridleyi'. Lopes and Alvarenga 1955: 181. Bulimus (Bulimulus) ridleyi. Oliveira and Oliveira 1984; 19. Type locality. Fernando de Noronha Archipelago, Fer¬ nando de Noronha Island and Rata Island. Original (Smith 1890: 501): “Living under bark of Mango-trees in the garden and on north side of island; also at base of the Peak, north side, under stones, and on Rat Island.” Distribution. Known only from Fernando de Noronha Archipelago. Type material. Lectotype NHMUK 1888.6.27.106 (des¬ ignation by Breure and Ablett 2012). Paralectotypes: NHMUK 1888.6.27.107-110, 4 shells. Material examined. Types. BRAZIL: Fernando de Noronha: ANSP 71271, 7 shells, H.H. Smith leg., 1896; MNZ 205835, 4 shells, ex Suter coll. 5637; ANSP 81426, 1 shell; ANSP 100530, 4 shells, H.v. Ihering leg., 1910; ANSP 220399, 2 shells, ex B.R. Bales coll., IS. Schwengel leg., 1958; MNZ 205835,4 shells, ex Suter coll. 5637; MZSP 501, 14 shells, 1900; MZSP 7752, 20 shells. MZSP 30134, 10 shells, dunes of Praia das Caiei- ras, E.F. Nonato leg., 25/vii/1955; MZSP 30135, 8 shells, de Fiore leg., 1983; MZSP 31064, 1 shell, Praia do Meio, L.R.L. Simone & Souza leg. 22/vii/1999; MZSP 31305, 30 specimens, between Baia dos Porcos and Baia do Sancho, L.R.L. Simone leg., 21/vii/1999; MZSP 31676, 14 shells, Praia das Caieiras, L.R.L. Simone et al. leg., 23/vii/1999; MZSP 31681, 15 specimens, mangrove on Praia do Sudeste, C.M. Martins leg., 20/vii/1999; MZSP 31686, 50 specimens, Praia do Meio, C.M. Mar¬ tins leg., 17-23/vii/1999; MZSP 48824, 1 shell, Praia do Porto, 3°50'05"S, 32°24'04"W, L.R.L. Simone leg., 30/iv/2005; MZSP 48990, 9 shells, Praia das Caieiras, 3°50'19"S, 32°24'00"W, L.R.L. Simone leg., 3/v/2006; MZSP 49001, 11 shells, mangroove on Praia do Sudeste, 3°5F58"S, 32°25'35"W, L.R.L. Simone leg., 4/v/1005; MZSP 49089, 39 shells, in front of Morro Dois Irmaos and Cacimba do Padre, L.R.L. Simone leg., 3/v/2005; MZSP 86542, 13 shells; Praia do Porto, 3°50'11"S, 32°24'04"W, L.R.L. Simone et al. leg., 28/x/2007; MZSP 89929, >50 shells, 3°50'00''S, 32°24'05"W, L.R.L. Sim¬ one & C.M. Cunha leg., iii/2009; MZSP 89933, 17 shells, L.R.L. Simone & C.M. Cunha leg., ll/iii/2009; MZSP 89939, 2 specimens; 3°50'21"S, 32°24'10"W, L.R.L. Simone & C.M. Cunha leg., 12/iii/2009; MZSP 89940, 5 specimens, Mirante, L.R.L. Simone & C.M. Cunha leg., ll/iii/2009; MZSP 89993, 1 shell, Praia do Sudeste, 3°52'06"S, 32°25'32"W, L.R.L. Simone & C.M. Cunha leg., 9/iii/2009; MZSP 97854, 3 shells, ex J. Vaz coll., A. Niissenbaum leg., viii/1973; MZSP 97878, 3 shells, ex J. Vaz coll., Praia das Caieiras, C. Bardelli leg., vi/1994; MZSP 119089, 23 shells, Cacimba do Pa¬ dre, 3°50'36"S, 32°25T4"W, L.R.L. Simone et al. leg., 8/v/2013; MZSP 119090, 29 shells, Cacimba do Pa¬ dre, 3°50'36"S, 32°25T4"W, L.R.L. Simone et al. leg., 7/V/2013; NHMUK 1888.6.27.88-94, 7 shells; NHMUK 1888.6.27.95-100, 6 shells. Rata Island; NHMUK 1888.6.27.101-105, 5 shells; NHMUK 20170270, 4 shells, H.E.J. Biggs coll, H. Fiedrick leg.; USNM 134849, 1 shell, H.A. Pilsbry leg.; USNM 214401, 1 shell, H.A. Pilsbry leg.; USNM 307580, 2 shells, Hen¬ derson coll, H. Clapp leg.; USNM 518214, 3 shells, W. Williamson leg.; ZSM no nr., 2 shells, Blume coll. 4003; ZSM no nr., 4 shells, H.H. Smith coll. Diagnosis. The shell is smaller overall and has a more elongated and slender profile. The protoconch has a more raised ridge and its sculpture consists largely of more anas¬ tomosed riblets. Typically, there is no apertural dentition. Description. Shell small, bulimoid, slender; spire tall; W ~ 4%-5. Shell color ochre to brown; body whorl some¬ times darker than rest; fine white spiral band on middle portion of whorl; periumbilical spiral angulation dis¬ colored, whitish; peristome white. Protoconch (w ~PA) rounded, with raised ridge that becomes a faint subsu- tural ridge on teleoconch; sculptured by fine sinuous ax¬ ial riblets, which sometimes anastomose (especially on abapical area of whorl); transition to teleoconch unclear. Teleoconch smooth (except for growth lines). Suture well marked, but not deep. Aperture ovoid, elongated. Peri¬ stome refiected and slightly thickened; presence of small channel-like structure on division between parietal and palatal regions of aperture; parietal callus might be pres¬ ent in older specimens. Apertural teeth usually absent, but faint elongated tooth on middle portion of palatal region may be present (Fig. 2A). Umbilicus narrow, deep, sur¬ rounded by a periumbilical spiral angulation. Dimensions. Eectotype: H = 11.4 mm, D = 5.9 mm, h = 4.8 mm, d = 3.0, W = 5%, w = VA. Paralectotype #1: H = 9.5 mm, D = 5.3 mm, h = 4.4 mm, d = 3.0 mm, W = 414, w = 114. Paralectotype #2: H = 9.1 mm, D = 5.3 mm, h = 4.3 mm, d = 2.9 mm, W = 414, w = PA. Paralectotype #3: H = 8.9 mm, D = 5.2 mm, h = 4.3 mm, d = 2.9 mm, W = 414, w = PA. Paralectotype #4: H = 8.9 mm, D = 5.2 mm, h = 4.2 mm, d = 2.8 mm, W = 414, w = PA. Average (n = 63, except for w, where n = 10): H = 10.2 ± 1.00 mm (min = 7.8 mm, max = 12.7 mm), D = 5.7 ± 0.51 mm, h = 4.9 ± 0.45 mm, d = 3.5 ± 0.34 mm, W = 4% to 5 (min = 4%, max = 514), w = PA (occasionally 114). Remarks. Other than showing a reasonable variation in shell size, the species displays little conchological varia¬ tion (Fig. 2). Rare specimens, however, do deviate from the typical form, for instance by having broader shells with shorter spires (Fig. 2F-I) or by having a palatal tooth (Fig. 2A, lectotype). The color might vary from more ochre tones to more reddish-brown ones, but the single white spiral band on the mid-section of the whorl is always pres- zse.pensoft.net Zoosyst. Evol. 95(2) 2019, 453-463 459 ent. Hyperaulax ridelyi can be easily distinguished from its only congener, H. ramagei, by its smaller shell (with a sin¬ gle spiral white band) and more elongated and slender shell profile. Moreover, its protoconch has a more raised ridge and the riblets of its sculpture are much more anastomosed. Finally, H. ridleyi typically bears no apertural dentition (al¬ though a weak palatal tooth might be present; Fig. 2A). Unfortunately, not much can be found in the literature about this species’ habitat or habits, but the museum la¬ bels point to a variety of collection locales, albeit more usually referring to dead shells only. In any event, this species has been reported alive from forested areas, man¬ grove, beaches, dunes, and gardens. Hyperaulax ramagei (Smith, 1890) Figs 3, 4 “Turbine, in cui la prima voluta e (...)” Buonanni 1681: 185, fig. Tur¬ bine #44. Bulimus {Tomigerus) Ramagei E.A. Smith 1890: 500, pi. 30, fig. 8. Bulimus (Tomigerusl) Ramagei: Dali 1896: 415. Bonnanius bouvieri Jousseaume 1900: 39, pi. 1, fig. 19. Bonnanius bonnanius Jousseaume 1900: 41. Hyperaulax (Bonnanius) ramagei: Pilsbry 1901: 103, pi. 11, figs 60-62; Thiele 1931: 611; Morretes 1949: 153; Lopes and Alvarenga 1955: 181; Zilch 1960: 505, fig. 1772; Breure 1974: 52; Breure 1975: 1158; Oliveira et al. 1981: 350; Parkinson et al. 1987: 29; Abbott 1989: 106, text fig. Tomigerus (Bonnanius) ramagei: Parodiz 1962: 453. Bulimus (Tomigerus) ramagei: Oliveira and Oliveira 1984: 19. Bonnanius ramagei: Schileyko 1999: 339, fig. 419; Breure and Ablett 2012: 34, figs 20A, B, 20i. Hyperaulax ramagei: Salgado and Coelho 2003: 165; Salvador 2019: 87. Bonnarius [sic] ramagei: Simone 2006: 178, fig. 638. Type locality. Fernando de Noronha Archipelago, Fer¬ nando de Noronha Island, Ponta do Tabaco. Original (Smith 1890: 500): “imbedded in sandy mud on a raised reef at Tobacco Point (G.A. Ramage leg.)”. Distribution. Known only from Fernando de Noronha Archipelago. Type material. Lectotype NHMUK 1888.6.27.163 (des¬ ignation by Breure and Ablett 2012). Paralectotypes: NHMUK 1888.6.27.164-170, 7 shells. Material examined. Types. BRAZIL: Fernando de Noronha: ANSP 100531,4 shells, H.v. Ihering leg., 1910; MNZ 205835, 4 shells, ex Suter coll. 5637; MNHN- IM-2000-28020 syntype of Bonnanius bouvieri, Jous¬ seaume coll; MNZ 205822, 5 shells, ex Suter coll. 5639; MZSP 7738, 14 shells; MZSP 97933, 3 shells, ex J. Vaz coll., A. Niissenbaum leg., viii/1973; NHMUK 1902.10.16.4, 1 shell; MZSP 131996, 3 shells, Ponta das Caracas, 3°52'28''S, 32°25'24"W, F. Schunck leg., 27/ix/2013; NHMUK 20170271, 13 shells, from sand on north end of island, 16/vi/1887; USNM 518215, >30 shells, W. Williamson leg.; USNM 709805, >30 shells, dunes in Porto Santo Antonio, L. Storrs et al. leg., vii- viii/1973; USNM 709806, >30 shells, Porto Santo Anto¬ nio, L. Storrs et al. leg.; ZSM 7861, 1 shell, 1940; ZSM no nr., 3 shells. Diagnosis. The shell is larger overall and has a broader profile. The riblets on the second part of the protoconch are more defined. The peristome is strongly thickened and displays marked apertural teeth. Description. Shell medium-sized, bulimoid, rounded; W ~ 414. Shell color chestnut brown; spire apex light brown to cream-colored; up to four equidistant white spi¬ ral bands might be present on lateral portion of whorls (but entirely brown morphs also occur); periumbilical re¬ gion usually discolored, whitish; peristome and apertural teeth white. Protoconch (w -PA) rounded; first 14 whorl presenting undefined anastomosing sculpture; remain¬ der sculptured by fine sinuous axial usually well-defined riblets (but sometimes anastomosed in some areas) that become less pronounced towards teleoconch; transition to teleoconch unclear (but sometimes with thickening of the last riblet). Teleoconch smooth (except for growth lines, which become more marked towards aperture). Su¬ ture well-marked, but not deep. Aperture roughly ovoid, but angulate. Peristome refiected and strongly thickened; some older specimens show continuous thickening of the peristome; parietal callus might be present in older spec¬ imens. Apertural teeth present: two knob-like parietal teeth positioned slightly towards the interior of shell (not always present); long palatal tooth in the middle portion of palatal region (its surface goes from smooth to ser¬ rated, with up to three distinct points); columellar tooth elongated, with smooth surface. Both columellar and pal¬ atal tooth produce a marked depression on outer wall of shell. Umbilicus slit-like. Dimensions. Lectotype: H = 17.3 mm, D = 12.3 mm, h = 8.8 mm, d = 6.6, W = 4, w = 2. Paralectotype #1: H = 22.3, D = 15.6 mm, h = 10.9 mm, d = 7.7 mm, W = 4%. Paralectotype #2: H = 23.5 mm, D = 16.0 mm, h = 11.2 mm, d = 8.7 mm, W = 5. Paralectotype #3: H = 19.5 mm, D = 14.6 mm, h = 10 mm, d = 7.2 mm, W = 414, w = PA. Paralectotype #4: H = 19.6 mm, D = 13.7 mm, h = 8.8 mm, d = 6.7 mm, W = 414. Paralectotype #5: H = 20.3 mm, D = 13.2 mm, h = 9.8 mm, d = 7.4 mm, W = 4%, w = PA. Syntype of Bonnanius bouvieri: H = 22.5 mm, D = 15.4 mm (Breure, 1975). Average (n = 34, except for w, where n= 10): H= 17.9 ± 1.56 mm (min = 16.1 mm, max = 22.0 mm), D = 12.9 ± 0.96 mm, h = 9.7 ± 0.71 mm, d = 7.6 ± 0.63 mm, W = 414 (min = 4%, max = 5), w = PA (occasionally 2). Remarks. The names H. bouvieri and H. bonnanius were synonymized with H. ramagei by Pilsbry (1901); this de¬ cision is followed here. The syntype of H. bouvieri (Fig. zse.pensoft.net 460 Salvador, R.B. & Cavallari, D.C.: Revision of Hyperaulax Figure 3. Hyperaulax ramagei. A-E. Lectotype, NHMUK 1988.6.24.163. F-J. Paralectotypes #1 to #5 (in order), including large forms in apparent sub-fossil state, NHMUK 1988.6.24.164-170. K. Specimen without the white spiral bands, NHMUK 1902.10.16.4. zse.pensoft.net Zoosyst. Evol. 95(2) 2019, 453-463 461 Figure 4. Hyperaulax ramagei. A-C. Syntype of Bonnanius bouvieri, MNHN-IM-2000-28020 (MNHN). D. Reproduction of “Tur¬ bine #44” of Buonanni (1861), the holotype of Bonnanius bonnanius. E. Protoconch detail, NHMUK 1902.10.16.4. 4A-C) is indistinguishable from H. ramagei, but the dis¬ cussion regarding H. bonnanius is slightly more colorful and it is worthwhile to recapitulate it here. Its original description (Jousseaume 1900) was based upon the work of the Jesuit scholar Filippo Buonanni (1638-1723), who compiled the first conchology manual (Buonanni 1681) and is thus considered the Father of Conchology (Leon¬ hard 2007). As Pilsbry (1901) argued, Buonanni’s (1681) description of his Turbine #44 and its illustration (allowing for some distortion in the drawing) are vastly consistent with H. ramagei. Despite later authors such as Linnaeus having relied on Buonanni’s work, this particular species was overlooked until Jousseaume (1900) published it as Bonnanius Bonnanius, misspelling the Jesuit’s name and likely without knowing the work of Smith (1890). The shell features of H. ramagei display some mor¬ phological variation: (1) shell size, from some rather small specimens to very large ones (H^^^ =16 mm and ^max ^ mm); (2) shell color can go from entirely brown to marked with four white spiral bands; (3) aperture size, relative to remainder of the shell; (4) shell shape, with some specimens having a much shorter spire (Fig. 3A-C, lectotype); (5) two parietal teeth might be absent (Fig. 31); (6) the surface of the palatal tooth goes from nearly smooth (Fig. 31, J) to serrated (Fig. 3F, G), with up to three distinct points (Fig. 3A), reminiscent of the carnas- sial tooth of Carnivora (apparently this is not related to the age of the individual or to the freshness of the speci¬ men when collected). The syntype of H. bouvieri show a four-pronged palatal tooth, which is also unusually large, and a three-pronged parietal tooth (Fig. 4A-C); this could be seen as morphological variation, but, as this specimen bears a mark of breakage near the aperture and further growth (Fig. 4C), it could be simply post-trauma anom- zse.pensoft.net 462 Salvador, R.B. & Cavallari, D.C.: Revision of Hyperaulax alous growth. For a comparison with its single congener, H. ridleyi, see the Discussion section of that species. Some of the specimens available (including some para- lectotypes) appear to be of a sub-fossil state, as already not¬ ed by Smith (1890). These appear to be larger than the fresh specimens, but this could be due to collection bias towards larger specimens; at present, there is not enough sub-fossil material for a statistically meaningful assessment. Conclusions Hyperaulax is here classified in the Odontostomidae and presently contains two species, H. ridleyi and H. ramagei, both endemic to Fernando de Noronha Archipelago. This genus seems to be more closely related to Tomigerus than to other odontostomids. According to recent collection events on Fernando de Noronha, H. ramagei cannot be found alive in spite of meaningful search efforts (L.R.L. Simone personal com¬ munication). In fact, museum specimens of H. ramagei still bearing the periostracum typically date back to the first half of the 20* century. It is a troubling possibility that presently this species has a much-reduced range or is altogether extinct. Future collection efforts should focus on this species to properly define its status according to current guidelines for conservation (lUCN 2012). Finally, another curious aspect of the fauna of Fernando de Noronha deserving further study is Amphisbaena ridleyi Boulenger, 1890, an amphisbaenid (worm lizard) endemic to the archipelago that possesses adaptations for a duropha- gous diet including a large proportion of land snails (Pre¬ gill 1984). In the original report, the author could not indi¬ cate which species of terrestrial gastropods were part of the lizard’s diet and a study involving this species and possible defensive adaptations of the snails would be very welcome. Acknowledgements We are very grateful to Luiz R.L. Simone (MZSP), Jon Ablett (NHMUK), and Enrico Schwabe (ZSM) for grant¬ ing access to the material under their care; to Ellen Wild- ner (ANSP) and Eisa Comer and Ellen Strong (USNM) for photos and information about the material housed in the collections under their care; to Barbara M. Tomotani (MNZ) for help with lab work; to Christina G. Martin (SMNS) for the SEM images; and to Bram Breure and Frank Kohler for the helpful comments on the manu¬ script. RBS received support from: SYNTHESYS Proj¬ ect (proposal GB-TAF-6613), financed by the European Community Research Infrastructure Action under the FP7 Integrating Activities Programme; Malacological Society of Eondon (Early Career Research Grant 2017); Staatliches Museum fur Naturkunde Stuttgart (SEM im¬ aging); Bruce Fraser Hazelwood fund and MNZ. References Abbott RT (1989) Compendium of land shells. American Malacolo- gists, Melbourne, 240 pp. Aulfenberg K, Slapcinsky J, Portell RW (2015) A revision of the fossil taxa assigned to Hyperaulax (Gastropoda: Odontostomidae), with the description of a new genus (Gastropoda: Bulimulidae). The Nau¬ tilus 129(2): 54-62. Breure ASH (1974) Catalogue of Bulimulidae (Gastropoda, Euthyneu- ra), II. Odontostominae. Basteria 38(5-6): 109-127. 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Appendix 1 Below are listed the COI sequenees of other Orthalieoidea species, obtained from the work of Breure and Romero (2012), used to ascertain the classification of the species studied herein. The list is organized by family and species (in alphabetical order), followed by the respective Gen- Bank accession number. Bulimulidae: Bostryx agueroi JF514623; Bostryx bilineatus JF514637; Bostryx edmundi JF514622; Bostryx longispira JF514624; Bostryx strobe- li JF514636; Bostryx superbus JF514621; BuUmulus di- aphanus JF514633; BuUmulus guadalupensis JF514630; BuUmulus hummelincki JF514629; BuUmulus sporadicus JF514632; BuUmulus tenuissimus JF514631; Drymaeus inusitatus JF514648; Drymaeus laticinctus JF514646; Drymaeus multifasciatus JF514647; Drymaeus serra- tus JF514649; Drymaeus vexillum JF514625; Naesiotus quitensis JF514635; Naesiotus stenogyroides JF514650; Neopetraeus tessellatus JF514627; Rabdotus alternatus JF514638; Scutalus chiletensis JF514628. Odontosto¬ midae: Clessinia cordovana cordovana JF514618; Cles- sinia cordovana stelzneri JF514617; Clessinia pagoda JF514613; Cyclodontina guarani JF514619; Plagiodon¬ tes multiplicatus JF514620; Spixiapervarians JF514614; Spixia philippii JF514612; Spixia popana JF514616; Spix¬ ia tucumanensis JF514615. Simpulopsidae: Leiostracus perlucidus JF514640; Simpulopsis decussata JF514639. The sequence of the outgroup taxon was obtained from GenBank: Planorbis planorbis EF012175 (Planorbidae). zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 465-505 | DOI 10.3897/zse.95.36762 4>yEnsPFr. BERLIN Contributions to the taxonomy of the long-jawed orb-weaving spider genus Tetragnatha (Araneae, Tetragnathidae) in the Neotropical region, with comments on the morphology of the chelicerae Pedro de Souza Castanheira\ Renner Luiz Cerqueira Baptista\ Daniela Dos Passos Pizzetti^, Renato Augusto Teixeira^ 1 Laboratdrio de Diversidade de Aracnideos, Universidade do Brasil/ Universidade Federal do Rio de Janeiro. Av. Carlos Chagas Filho 373, 21941-902, Ilha do Fimddo, Rio de Janeiro, Brazil 2 Laboratdrio de Arachnida e Myriapoda, Museu de Ciencias e Tecnologia - PUCRS, Av. Ipiranga, 6681, predio 40, Sala 125, 90619-900, Porto Alegre, RS, Brazil http://zoobank. org/6lA44D72-5E9B-40C6-9440-2 7E39511 ODES Corresponding author; Pedro de Souza Castanheira (pedrocastanheira.bio@gmail.com) Academic editor; Z)ara7o♦ Received 4 June 2019 ♦ Accepted 18 September 2019 ♦ Published 22 October 2019 Abstract We newly diagnose, illustrate, and clarify the distribution ranges of six of the most common and broadly distributed species of Tetragnatha Latreille, 1804 found in the Neotropical region. Twenty new junior synonyms from around the world are included, nine for T. bogotensis Keyserling, 1865, four for T. mandibulataWalckmaer, 1841, three for T. keyserlingi Simon, 1890, three for T. nitens (Audouin, 1826), and one for T. e/owgato Walckenaer, 1841. Tetragnatha vermiformis Emerton, 1884 is newly recorded from South America. The Argentine T. major Holmberg, 1876 and T. riparia Holmberg, 1876 are considered nomina dubia. Finally, we discuss the terminology of the structures of the chelicerae to establish a coherent nomenclature for teeth and fang cusps. Key Words Biodiversity, Araneoidea, Systematics, Tetragnathinae Introduction The long-jawed spider genus Tetragnatha Latreille, 1804 (family Tetragnathidae) comprises 349 species, 67 of which restricted to the Neotropics (World Spider Cata¬ log 2019). All species share spiny and elongate chelicer¬ ae, elongate and dorsally flattened carapace, parallel eye rows, and female genital openings located at the poste¬ rior end of the procurved epigastric furrow (Levi 1981; Gillespie 1992a, b; Barrion et al. 2011). Nonetheless, many species are poorly diagnosed at the taxonomic level and their geographic ranges are imperfectly known. Although no complete revision is available for Tetrag¬ natha, species redescriptions and local revisions are fre¬ quent. The first revisionary papers on the genus addressed species from North America (Seeley 1928) and Europe (Lendl 1886; Wiehle 1939, 1963). Chickering (1957a) reviewed the Central American and Mexican species, fol¬ lowed by those from Jamaica (Chickering 1957b); Pana¬ ma (Chickering 1957c); United States and Canada (Levi 1981); Australasia (Okuma 1987), Asia (Okuma 1988a, b), Mexico, and part of the Neotropical region (Okuma 1992), Hawaii (Gillespie (1992a, b, 2003a); Marquesas Islands (Gillespie 2003b); and Society Islands (Gillespie (2003c)). In this paper, we analyse most of the widespread spe¬ cies of Tetragnatha in the Neotropical region. We pro¬ pose 20 new synonymies for five of them and include new records from South America for one species previously known from Asia and introduced in North and Central America. We also provide new diagnoses, illustrations, and many new distribution records. Furthermore, we consider two species from Argentina as nomina dubia. Copyright Pedro de Souza Castanheira etal. This is an open access articie distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the originai author and source are credited. 466 Pedro de Souza Castanheira et al..: Taxonomy of Neotropical Tetragnatha Methods The taxonomic summary for all species is abbreviated to save space, including only the information we consider most relevant. See the World Spider Catalog (2019) for a complete list of synonymies. Terminology for chelicer- ae follows Okuma (1987, 1992) and Gillespie (1992a, 1992b). For the male palp, see Levi (1981), and forthe fe¬ male genitalia, see Alvarez-Padilla and Hormiga (2011). Colour patterns were described based on specimens pre¬ served in 75% ethanol. Structures were cleaned using a Cristofoli Ultrason¬ ic Cleaner and positioned in 70% ethanol gel or glass spheres for automontage photographs and measurements. Images were taken with a Leica DFC450 camera mount¬ ed on a Leica M205C stereoscope microscope (Leica Camera AG, Wetzlar, Germany) at the Laboratorio de En- tomologia, Universidade do Brasil/Universidade Federal do Rio de Janeiro. All images were edited with Adobe Photoshop CS5.1 and figures were prepared using Adobe Illustrator CSS. 1 (Adobe Inc., San Jose, California, USA). Measurements are given in millimeters. The position of teeth and fang cusps (upward, downward, distalward, and basalward) was noted when the chelicerae were attached to the body. The genital fold length was measured from the inner angle of book-lung plates to the posterior rim of the fold. The genital fold proportion is the comparison of its length versus the span between the outer angle of posterior rim of one book-lung plate to the outer angle of the other. Males were matched with conspecific females by cheliceral morphology and collection sites. For scanning electron microscopy (SEM), samples were critically point dried, mounted on adhesive cop¬ per tape (Electron Microscopy Sciences, EMS 77802), affixed to a stub and sputter-coated with Au-Pd for ex¬ amination under high vacuum with a JEOE JSM-6510 microscope at Eaboratorio de Imagens, Institute de Bio- logia, Universidade Federal do Rio de Janeiro; a Philips XE 30 Field Emission ESEM at the Centro de Microsco- pia e Microanalises, Pontificia Universidade Catolica do Rio Grande do Sul, and a JEOE JSM-6390EV at Centro de Microscopia, Funda^ao Oswaldo CRuz. To clear female genitalia, we used a borax solution (Alvarez-Padilla and Hormiga 2008) and digestive en¬ zyme tablets of “Orthoplex D.E.F” (Bioconcepts Pty Etd, Banyo, Queensland, Australia). We noted that sometimes the internal genitalia may appear darker or lighter, after clearing, usually related to the time spent in the solution. For example, the spermathecae and central membranous sac may appear dark, roundish, and well defined (Fig. 2H) or pale and partially collapsed (Fig. 21). Maps were produced using QGIS v. 2.14 software and geographic coordinates were extracted from original labels. When no coordinates was available, the closest nearby area coordinates were obtained from Global Gaz¬ etteer V. 2.3 (http://www.fallingrain.com/world/index. html) or Google Earth v. 9.1.39.1 (https://earth.google. com/web/). Cited institutions and their acronyms are: AMNH American Museum of Natural History, USA (curator: E. Prendini); CAS California Academy of Sciences, USA (E. Esposito); IBSP Institute Butantan, Brazil (A. Brescovit); MACN Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Argentina (M. Ramirez); MCTP Museu de Ciencia e Tecnologia da Pontifi¬ cia Universidade Catolica do Rio Grande do Sul, Brazil (R. Teixeira); MCZ Museum of Comparative Zoology, Harvard University, USA (G. Giribet); MIZ Muzeum i Instytut Zoologii Polskiej Aka- demii Nauk, Poland (W. Wawer); MLP Museo de Ea Plata, Argentina (E. Pereira); MLPC Mello-Eeitao’s private collection, now at Museu Nacional, Universidade do Brasil/ Universidade Federal do Rio de Janeiro, Brazil (A. Kury); MNHN Musee National d’Histoire Naturelle, France (C. Rollard); MNRJ Museu Nacional, Universidade do Brasil/ Universidade Federal do Rio de Janeiro, Brazil (A. Kury); MPEG Museu Paraense Emilio Goeldi, Brazil (A. Bonaldo); MRAC Royal Museum for Central Africa, Belgium (D. Van den Spiegel); MZUF Universita di Firenze, Museo Zoologico “Ea Specola”, Italy (E. Bartolozzi); MZUSP Museu de Zoologia da Universidade de Sao Paulo, Brazil (R. Pinto-da-Rocha); NHM Natural History Museum, United Kingdom (J. Beccaloni); NHMW Naturhistorisches Museum Wien, Austria (C. Horweg); NHRS Naturhistoriska Riksmuseet (Swedish Muse¬ um of Natural History), Sweden (J. Stigen- berg); NMB Naturhistorisches Museum Basel, Switzer¬ land (A. Haenggi); NMV Museums Victoria, Australia (K. Walker); OUMNH Oxford University Museum of Natural His¬ tory, United Kingdom (Z. Simmons); SMF Senckenberg Museum Frankfurt, Germany (P. Jager); UFRJ Eaboratorio de Diversidade de Aracnideos, Universidade Federal do Rio de Janeiro, Brazil (R. Baptista); USNM Smithsonian National Museum of Natural History, USA (J. Coddington); ZMB Museum fur Naturkunde, Humboldt-Univer- sitat, Germany (J. Dunlop); ZMH Zoologisches Museum Hamburg, Germany (D. Harms). zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 465-505 467 Abbreviations used below for the chelicera: a male dorsal apophysis, used to lock the fangs of females during copulation. AXl auxiliary guide tooth of the lower row, present in some species. AXu auxiliary guide tooth of the upper row, above Gu, present in some species. BC basal cusp on the cheliceral fang of females (new terminology). Okuma applied different names to this structure: excrescence (1987) and posterior cusp (1992). CB cheliceral bulge, a protruding area between the two rows of teeth, near the base of the fang (new terminology). CRu cheliceral crest, a protruding marked area on the upper row of teeth (new terminology). CRl cheliceral crest, a protruding marked area on the lower row of teeth (new terminology). Gu guide tooth of the upper (or dorsal) row (follows Okuma 1983, 1987, 1992). G1 guide tooth of the lower (or ventral) row (follows Okuma (1983, 1987, 1992). IC inner cusp of fang (follows Okuma 1987, 1992) (new abbreviation). L2-n teeth on the lower row numbered from the distal end after Gl. OC outer cusp of fang (present in some species) (new terminology). rsu remaining proximal teeth on the upper row of males after T (Okuma 1987; Gillespie 1992a, 1992b). rsl remaining proximal teeth on the lower row of males and females after the last specialized tooth (Okuma 1987). si first maj or tooth after Gu in the upper row of males (absent in some species). T elongated tooth in the upper row of some males (Okuma 1983, 1987, 1992). t a tooth or prominence found in males of some species. U2-n teeth on the upper row numbered from the distal end after Gu. Abbreviations for male and female genitalia, male palps: E embolus; C conductor; Y cymbium; P paracymbium K knob at the ectal side of paracymbium (new terminol¬ ogy), L translucent lobe at the mesal side of paracymbium, N notch at the apex of paracymbium. Female genitalia: GF genital fold; Sp spermatheca; CS central membranous sac. Results Order Araneae Clerck, 1757 Family Tetragnathidae Menge, 1866 Genus Tetragnatha Latreille, 1804 Tetragnatha bogotensis Keyserling, 1865 Figs 1-4, 20A, 21A-G, 22A Tetragnatha bogotensis Keyserling 1865: 854, pi. 21, fig. 5 ($). Tetragnatha awc/ZwaTaczanowski 1878: 144, pi. 1, fig. 2 (;^); Levi 1981: 291 (removed from syn. with T. nitens) syn. nov. Tetragnatha boydi O. Pickard-Cambridge 1898: 389, pi. 31, fig. 4 syn. nov. Tetragnathapeninsulana Banks 1898: 246, pi. 15, fig. \2 {S $); Levi (1981): 291 (removed from syn. with T. nitens) syn nov. Tetragnathapraedator TnWgXQW 1910: 147, pi. 3, fig. 69 ((3) Tetragnatha mandibulata bidentata Gravely 1921: 442, fig. 3c, f ((3 $). Tetragnatha eremita Chamberlin 1924: 645, figs 89, 90 ((3); Levi 1981: 292 (removed from syn. with T. nitens) syn. nov. Tetragnatha nitens Lawrence 1927: 27, pi. 3, fig. 61, pi. 4, fig. 77 {S $ misidentified). Tetragnatha bemalcuei Mello-Leitao 1939: 67, figs 42-44 ($) syn. nov. Tetragnatha ramboi Mello-Leitao 1943: 193, fig. 24,24a, b ((3) syn. nov. Tetragnatha haitiensis Bryant 1945: 408, fig. 37 (;^); Levi 1981: 292 (removed from syn. with T. nitens) syn. nov. Tetragnatha nitens kullmanni Wiehle 1962: 379, figs 1-5, 6b, 9-11, 14, 15 ((3 $); Wunderlich 1992: 365 (removed from syn. with T nitens) syn nov. Tetragnatha infuscata Benoit 1978: 667, fig. 2D, E ((3); Saaristo 2003: 23, figs 21A, B, 25 (syn. with T boydi rejected, see T. mandibulata). Tetragnatha boydi praedator Schmidt and Krause 1993: 6, fig. 5 ((3 $) syn. nov. Type material. Tetragnatha bogotensis'. COLOMBIA: syntypes, “Nova Granada” [surely Bogota], 1859- 1863? Alexander Lindig leg., not located, probably lost. Tetragnatha andina'. PERU: SS, $$ syntypes, “Amable Maria” [surely province of Tarma, region of Junin] (38 syntypes. Coll. K. Jelski, MIZ 225346-225446), exam¬ ined (photos). Tetragnatha boydi'. YEMEN: $ syntype, Socotra, not located. Tetragnatha peninsulana'. MEXI¬ CO: (7(7 (CAS), $ $ (MCZ 22587) syntypes: San Jose del Cabo, Baja California Sur, females examined (photos), males destroyed (Eevi 1981). Tetragnatha praedator. KENYA: 4(7, syntypes (NHRS JUST000000671-672), Kilimandjaro and Meru, examined (photos). Tetragnatha eremita'. MEXICO: S holotype, Baja California, Puerto Escondido, Arroyo de Escondido (Coll. J. C. Chamberlin, 14.vi.l921, male pedipalp in MCZ 15283, labeled RVC nil; whole specimen in CAS 1430), examined (photos of CAS 1430); $ paratype, same data as holotype (MCZ 25228), examined (photos). Tetragnatha bemalcuei'. PARAGUAY: $ holotype, Asuncion (Coll. C. Ternetz, 1895, NMB-ARAN 01092a), examined (photos). Tetrag¬ natha ramboi'. BRAZIE: S, lectotype, Rio Grande do Sul (Coll. Father Balduino Rambo, MNRJ 42467), examined. Tetragnatha haitiensis'. HAITI: $ holotype, “Hispanio- zse.pensoft.net w ^ 468 Pedro de Souza Castanheira et al..: Taxonomy of Neotropical Tetragnatha Figure 1. Tetragnatha hogotensis Keyserling, 1865, male (UFRJ 0035). A. Dorsal habitus; B. Ventral habitus; C. Chelicerae upper row and eyes; D. Chelicerae lower row and maxilla; E-H. Left chelicera: E. Upper view; F. Inner view; G. Lower view; H. Outer view; I-K. Left male palp; L Mesal view; J. Dorsal view; K Ventral view (paracymbium). Scale bars; 2 mm (A, B); 1 mm (C, D, E, F, G, H); 0.5 mm (I, J, K). zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 465-505 469 la” [Ennery] (Coll. R J. Darlington Jr., 07.vii.l934, MCZ 21516), examined (photos). Tetragnatha nitens kullman- nr. ITALY: S, holotype (SMF 12741), 2$, 1 immature, paratypes (SMF 12742), Sardinia, Oliena (E. Kullmann leg.), not examined. Extended diagnosis. Males of T. bogotensis are similar to T. nitens (Audouin, 1826) and share chelicerae with an elongated ‘a’, and AXu and ‘t’ extremely elongated and distally bent. AXu and ‘E differ as follows: thick and trian¬ gular in T. bogotensis (Figs 1C, E, F, 3B), but thinner and ‘F sickle-like in T. nitens (Figs 14C, D, F, 16A). Gu lon¬ ger, sharper, straight and larger than U2 in T. bogotensis (Figs 1C, E, F, 3B) while it is shorter than U2 and slightly curved downward in T. nitens (Figs 14C, D, 16A). Palps share medium-sized conductors without pleats (Figs II, 3A, 14G, 15D, E). Tetragnatha bogotensis palps differ by shorter tibias (ca 3x longer than wide) (Figs II-K, 3A), conductors not twisted at their distal halves, with com¬ pletely folded apexes enclosing the emboli tips (Figs II, J; 3A, E; 20A) and longer paracymbia, overreaching the upper border of teguli, each bearing deeper notch, result¬ ing in an elongated tip of the paracymbium proper, and thinner and relatively narrow translucent lobe (Figs II, K; 3D). In T. nitens, palps have longer and thinner tibias (al¬ most 4 x longer than wide) (Figs 14G-I, 16E), conductors twisted in distal half with their apexes rounded and ex¬ cavated, exposing the emboli tips (Figs 14G, H, 16D-F, 20E) and shorter paracymbia that do not reach the upper border of teguli, with expanded and very wide translucent lobes (Figs 141, J, 16F). The epiandrous field is straight, with 19 fusules, in T. bogotensis (Fig. 3F) but arched and smaller, with only 15 fusules, in T. nitens (Fig. 16C). Females are similar to T. nitens and also T. mandib- ulata Walckenaer, 1841, sharing: bulky body, wide and pointed at the terminal end (Figs 2A, B, 12A-C, 15A, B), and elongate genital fold (Figs 2G, 12H, 15J). Their chelicerae also bear an evident basal cusp (BC) in the fang (Figs 2D, E, 12D-G, 13C, D, 15D, F-I, 16B), Gu elongated and not contiguous to U2 (Figs 2C, D, 12D, E, 13C, 15C, E, F, 16B); G1 large and pointed distalward, followed by smaller L2 and L3 (Figs 2D, E, 3C, 12E, F, 13D, 15D, F, H, 16B). T. bogotensis and T. mandibulata differ from T. nitens by distinct bulky AXl, which are elongated, pointed and with large bases, clearly visible and overreaching the claws even in upper view (Figs 2C-F, 3C, 12D-G, 13C, D), while in T. nitens it is short and not pointed (Figs 15F-H, 16B). T. bogotensis and T. mandibulata also differ by the conspicuous bulge (CB) in the area between both rows (Figs 2C, E, 3C, 12D, F, 13C, D), which is absent in T. nitens. Tetragnatha bogotensis chelicerae can be distinguished from T mandibulata by the following characters: more robust basal cusp (BC), placed at the middle line of the lower side of the claw, compared to a smaller BC, displaced towards the outer face of the claw (Figs 2D, E, 3C-F, 12D-G, 13C, D); Gu straight with a large basis, being separated from U2 by a wide and deep furrow, versus both teeth not quite spaced (Figs 2C, D, 3C, 12D, E, 13C); AXl bulkier, with much larger basis, compared to a thinner and shorter tooth (Figs 2C-F, 3C, 12D-G, 13C, D); G1 shorter, straight and pointed, with wider basis, and much smaller than AXl, versus a longer and slanted Gl, regularly tapered and just a bit smaller than AXl (Figs 2D, E, 12E, F, 13D); and CB rounded and wide, extending from a bit above L2 to L3, and placed in the middle line between both rows of teeth, contrasting to a smaller and lower CB, extending from the basis of Gl to the middle of the gap between L2 and L3, and adjoined to the basis of L2 (Figs 2C, E, 3C, 12D, F, 13C, D). In T nitens, BC is placed at a similar position to T bogotensis, but is larger (Figs 15D, F-I, 16B) and Gu is connected to U2 by a thin and dark ridge, with a gap of similar size to T bogotensis (Figs 15C, E, F, 16B). The genital fold (Figs 2G, 12H, 15J) is similar in all three species but shorter in T nitens (genital fold length around 0.6x the width), intermediate in T bogotensis (0.8x) and longer in T mandibulata (l.lx). The internal genitalia of T bogotensis is more similar to T mandibu¬ lata, with medium-sized spermathecae and a sclerotized and rounded fundus (Figs 2H, 1,121), in contrast to wider spermathecae without a well-defined fundus in T nitens (Fig. 15K, L). The central membranous sac (CS) of T bogotensis is medium-sized, with almost the same size of the spermathecae, and placed at the same level of their bases (Fig. 2H, I), while in T mandibulata it is massive, longer, and placed at the same level as the spermathecae (Fig. 121). In T nitens, the CS is about the same size as in T bogotensis, but with a stalk of variable size, some¬ times longer than the spermathecae, placing the apical portion of the CS at the same level or anteriorly to the spermathecae (Fig. 15K, L). Synonymy and notes. Keyserling (1865) described this species based on female specimens from “Nova Granada”, which encompasses a large area ranging from Panama to Ecuador, and which were collected by Alexander Lindig, who gathered many animals and plants in Bogota (Colom¬ bia) from 1859 to 1863 (Meagher 2012). Therefore, the type specimens were most likely collected, at least partial¬ ly, in this city, as indicated by the name T bogotensis. The drawing of one female by Keyserling (1865: pi. 21, fig. 5) allows the recognition of the species by showing the long and robust AXl placed near to a smaller and traverse Gl of the right chelicera. In the original description, Keyserling wrote that he had “many copies in my collection”, without citation of any males (Keyserling 1865: 855). The first author of this paper visited three collections with type material by Keyserling: MIZ; NHM (Beccaloni 2012), and ZMB (Kretschmann 2006), and we also con¬ tacted curators of NHMW and USNM. Possible well-pre¬ served type specimens were found in two vials at only the NHM. In the first vial (Fig. 4A), there were three females labeled “type” and originated from “Taquara” (originally “Taquara do Mundo Novo”, state of Rio Grande do Sul, Brazil). In the second vial (Fig. 4B), there were two fe¬ males and one male labeled “Bogota”, but with no clear indication as type. All five females from both vials are T bogotensis but the male from Bogota belongs to T nitens. zse.pensoft.net 470 Pedro de Souza Castanheira et al..: Taxonomy of Neotropical Tetragnatha AVii A .Gil AXii A - AVI V -AVI — k AXii Figure 2. Tetragnatha bogotensis Keyserling, 1865, female. A. Dorsal habitus (UFRJ 1314); B. Lateral habitus (UFRJ 1314); C-F. Left chelicera (UFRJ 1314); C. Upper view; D. Inner view; E. Lower view; F. Outer view; G-I. Genital area; G. Genital fold, ventral view (MCTP 3381); H. Internal genitalia, cleared, ventral view (UFRJ 1314); 1. Internal genitalia, cleared, ventral view (MCTP 13581). Scale bars; 2 mm (A, B); 1 mm (C, D, E, F, G); 0.2 mm (H); 0.1 mm (I). zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 465-505 471 Figure 3. Tetragnatha bogotensis Keyserling, 1865, SEM photos. A. Left male palp, mesal view (MCTP 4299); B. Left male cheli- cera, upper view (MCTP 4299); C. Left female chelicera, upper view (MCTP 4299); D. Left male palp paracymbium, ventral view (ULRJ 0044); E. Left palp conductor detail, mesal view (MCTP 4299); F. Epiandrous field, ventral view (UFRJ 0044). Scale bars: 0.2 mm (A); 0.5 mm (B, C); 0.1 mm (D); 0.05 mm (E); 0.02 mm (F). zse.pensoft.net 472 Pedro de Souza Castanheira et al..: Taxonomy of Neotropical Tetragnatha We consider Keyserling’s syntypes to be lost, as there is no clear indication that the specimens above were used to de¬ scribe the species: females from Taquara are not syntypes because Keyserling had not cited any specimens from that Brazilian locality, and females from Bogota are also not syntypes as they were not labeled as type or “N. Grana¬ da” and are kept in the same vial as the additional male. Therefore, the confirmation of the identity of Keyserling’s species relies on the illustration in the original description and the fact that specimens collected in Bogota and oth¬ er cities from Colombia clearly display the key cheliceral teeth characters that allow the diagnosis of T. bogotensis. Tetragnatha boydi has been redescribed and illustrat¬ ed many times (see World Spider Catalog 2019). Pick- ard-Cambridge (1898) described this species based on a female from Socotra (Yemen), giving detailed illus¬ trations of the lower and inner views of the chelicerae (Pickard-Cambridge 1898: fig. 4a, b). Males were first de¬ scribed as T. praedator Tullgren, 1910 from Kenya, a spe¬ cies considered junior synonym afterwards (see below). Unfortunately, the female type material was not located at NHM or in OUMNH, institutions where O. Pickard-Cam¬ bridge normally deposited his specimens, and remains lost. Lawrence (1927) (sub T. nitens) and Okuma (1983) were the first authors to correctly match both sexes in their papers. Lawrence (1927) gave very detailed illustrations of the male left palp and chelicera, clearly showing the diagnostic characters of this species. On the other hand, the females were not illustrated, but his description points to the “inferior margin with a large apical tooth” (Law¬ rence 1927: 28), thereby rejecting the identification as T. nitens. Okuma (1983) also correctly matched the couples, gave very detailed drawings of chelicerae and genitalia, synonymised T. mandibulata bidentata and T. nitens kull- manni with T. boydi, and gave the first records for Bra¬ zil. Later, Okuma (1992) also provided new drawings of the species. After examining illustrations by Keyserling (1865: pi. 21, fig. 5), the NHM specimens of T. bogotensis from the type locality (Bogota), and making comparisons with O. Pickard-Cambridge’s (1898) illustrations and the later illustrations by Tullgren (1910), Lawrence (1927), and Okuma (1983, 1992), we conclude that all specimens belong to the same common species and establish that T. boydi is a junior synonym of T. bogotensis. Tetragnatha praedator Tullgren (1910: fig. 69a, b) was described from four male specimens from Kilimandjaro and Meru (Kenya). Lessert (1915) also cited males of T praedator, and the species was later synonymised with T boydi by Roewer (1942). Finally, it was treated as a sub¬ species of T boydi by Schmidt and Krause (1993), who also described females from Comoros Island, forming the combination T boydi praedator Tullgren, 1910. After comparing Tullgren’s (1910) illustrations and photos of the syntypes we received (Fig. 21C) with the specimens we identified as T bogotensis, we observed that they clearly match. Thus, T boydi praedator is synonymised here with T bogotensis. On the other hand, the females assigned to T boydi praedator by Schmidt and Krause (1993: fig. 5) probably belong to a different species judg¬ ing by their illustrations, which are, however, too poor to allow a proper evaluation. Tetragnatha bemalcuei was described by Mello-Leitao (1939: 68, figs 42-44), who mentioned on the original description “a robust conical frontward apophysis” (AXl) on the lower row of the chelicerae. We examined detailed images of the holotype (NMB) that show the characters of T bogotensis (Figs 4D, 2IE) and establish T bemalcuei as a junior synonym of T bogotensis Keyserling, 1865. Mello-Leitao (1943) described T ramboi based on males and females from Rio Grande do Sul, south Bra¬ zil. He indicated the vial MNRJ 42467 as “tipo” (type in Portuguese) in the description, but did not label it as “ty- pus”, in contrast to his common practice (Fig. 4C). We examined the type series (one male, two females, and one immature specimen) and agree with Silva-Moreira et al. (2010) that the whole series of specimens should be treat¬ ed as syntypes. In the original description, the male was cited first and its diagnostic chelicerae and palps were il¬ lustrated (Mello-Leitao 1943: fig. 24a, b), whereas only the habitus of the female was illustrated (Mello-Leitao 1943: fig. 24). Under the rule of the “First Reviser” (ICZN 1999, article 24), we consider the sequence of descriptions and the presence of diagnostic illustrations in establishing the male as the lectotype of the species. This male clearly belongs to T bogotensis according to the chelicerae and palp diagnostic characters (Figs 1C, E, J, 3A, B, E, 20A, 2IF; Mello-Eeitao 1943: fig. 24b) and must be newly synonymised with this species. Finally, we consider the female and juveniles of the type series of T ramboi as misidentified specimens of T argentinensis Mello-Eeitao, 1931 which were erroneously attributed to T ramboi. We also note that several species previously considered junior synonyms of T nitens should be newly synonymised with T bogotensis. For example, Eevi (1981: 291, 292) established 13 junior synonyms of T nitens. Indeed, most of those species are correctly junior synonyms of it, but at least four should now be regarded as junior synonyms of T bogotensis (see T nitens below). It also seems that Eevi (1981: figs 23-29) matched males of T nitens with at least some females of T bogotensis, as it is evident by the female illustrations he gave. These clearly depict the large AXl of T bogotensis (Eevi 1981: figs 23-25) and similar genitalia (Eevi 1981: figs 27-29), with a pattern very dif¬ ferent from T nitens (Fig. 15K, E, Zhu and Zhang 2011: fig. 125G). In the “Variation” and “Diagnosis” sections of his paper, Eevi (1981: 292) pointed out “On Panamanian specimens the diagnostic tooth at the posterior base of the fang is as long as the chelicerae are wide, and is sometimes smaller than illustrated on the most northern specimens” and “The female chelicerae have a large posterior lateral tooth at the insertion of the fang”. Unfortunately, he did not provide the collection site for the female specimen he illustrated, and although there is no formal record for T bogotensis or any of its synonyms from the United States, at least some of the females from the southern USA cited as T nitens may belong to that species instead. Eevi (1981: 291, 292) synonymised T andina Taczanowski, 1878 with T nitens based on multiple male zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 465-505 473 - if 7 ' z ,1 > 1 V M. ^||| yi *^r f| / 41 D ^SEUM BASEL /'O^Z CZ, _ ^(^zfra^nci/^cc. ^emcUcMi.^ '■ ^ ^ /f ~L -3er>^ . zy /> Figure 4. Tetragnatha hogotensis Keyserling, 1865 labels. A. T. hogotensis from Taquara, Brazil (NHM); B. T. hogotensis from Bogota, Colombia (NHM); C. T. ramboi syntype from Rio Grande do Sul, Brazil (MNRJ 42467); D. T. bemalcuei holotype from Paraguay (NMB). and female syntypes from Amable Maria, Peru, however, Taczanowski’s (1878: fig. 2) poor illustration of the fe¬ male chelicerae allows its recognition as T. hogotensis, as verified in the photos we received from MIZ (Fig. 21A). Thus, T. andina is here removed from the synonymy of T. nitens and newly synonymised with T. hogotensis. Banks (1898) described T. peninsulana from two males and “several females” from San Jose del Cabo, Baja Cal¬ ifornia Sur, Mexico. Photos of females provided by the MCZ (Fig. 2IB) include three specimens of T. hogotensis and one of T. nitens, which we consider a misidentification. Additionally, according to Levi (1981), the male syntypes were destroyed. Based on the elongated AXl (Fig. 2IB), this species must also be removed from the synonymy of T. nitens and be newly synonymised with T. hogotensis. Tetragnatha eremita Chamberlin, 1924 was based on a male holotype from Baja California, Mexico. Chamber¬ lin (1924: figs 89, 90) provided a short description and two good illustrations that clearly show the characteristic shape and teeth formula of T. hogotensis, besides citing a female paratype collected at the same time. The male holotype is represented by the right pedipalp in the MCZ (MCZ 15283, RVC 1111), and by the whole specimen in CAS 1430. We were able to examine the holotype’s che¬ licerae through photos (Fig. 2ID), and thus confirmed its identity, removing T. eremita from the synonymy with T. nitens and newly synonymising it with T. hogotensis. Furthermore, Levi (1981: 292) followed Chickering (1957b: 2) in the synonymization of T. haitiensis Bryant, 1945, which was based on a female from Haiti, with T. ni¬ tens. Bryant (1945: fig. 37) illustrated the huge AXl tooth typical of T. hogotensis, which we also observed in the photos we received from MCZ (Fig. 21G). Therefore, T. haitiensis Bryant, 1945 is also removed from the synony¬ my of T. nitens and newly synonymised with T. hogotensis. Wunderlich (1992: 365) also mistook T. hogotensis as T. nitens, removing T. nitens kullmanni from its proper synonymy with T. boydi (Okuma 1983: 70). We agree with Okuma (1983) that this species is “undoubtedly identical with T. boydf\ as both males and females of T. nitens kullmanni bear the same diagnostic characters of T. hogotensis in comparison with T. nitens: male chelicerae with Gu longer than U2 (Figs IE, 3B; Wiehle 1962: fig. 9) and female chelicerae with a long AXl (Figs 2D-F, 3C; Wiehle 1962: fig. 15). Therefore, the synonymy with T. nitens is rejected and T. nitens kullmanni is a junior syn¬ onym of T. hogotensis. Variation. Males {n = 18): total length, 7.36-11.60; fe¬ males (n = 17): total length, 7.52-11.76. The gap between Gu and U2 is variable in males of this species and can have almost the double of the length of the specimen illustrated. Distribution. This species is widespread in the Neotropics and Mexico, but potentially north into the southern United States; it also occurs in the Old World, with records from Africa, Yemen, India, Nepal, and China (Fig. 22A). Tetragnatha elongata Walckenaer, 1841 Figs 5-7, 20B, 22B Tetragnatha elongataWdAckenaQr 1841: 211 0 9). Tetragnatha tropica O. Pickard-Cambridge 1889: 11, pi. 2, fig. 3 (5); F. O. Pickard-Cambridge 1903: 431, pi. 40, figs 10, 11 (3' $) syn. nov. Type material. Tetragnatha elongata: GUADELOUPE: 6' $ syntypes, lost; UNITED STATES OF AMERICA: S neotype, Raleigh, North Carolina (Coll. C. S. Brimley, 21-3Lviii.l944, MCZ 21192), not examined. Tetragna- zse.pensoft.net 474 Pedro de Souza Castanheira et al..: Taxonomy of Neotropical Tetragnatha Figure 5. Tetragnatha elongata Walckenaer, 1841, male (MCTP 28045). A. Dorsal habitus; B. Lateral habitus; C. Ventral habitus; D-G. Left chelicera; D. Upper view; E. Inner view; F. Lower view; G. Outer view; H-K. Left male palp: H. Mesal view with tibia; L Mesal view detail; J. Dorsal view; K. Ventral view (paracymbium). Scale bars; 2 mm (A, B, C); 1 mm (D, E, F, G, H); 0.5 mm (I, J, K). zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 465-505 475 Figure 6. Tetragnatha elongata Walckenaer, 1841, female. A. Dorsal habitus (MCTP 28045); B. Lateral habitus (MCTP 28045); C-F. Left chelicera (MCTP 28045): C. Upper view; D. Inner view; E. Lower view; F. Outer view; G, H. Genital area: G. Genital fold, ventral view (MCTP 28045); H. Internal genitalia, cleared, ventral view (MCTP 28306). Scale bars: 5 mm (A, B); 1 mm (C, D, E, F); 0.5 mm (G); 0.2 mm (H). zse.pensoft.net 476 Pedro de Souza Castanheira et al..: Taxonomy of Neotropical Tetragnatha Figure 7. Tetragnatha elongata Walckenaer, 1841, SEM photos. A. Left male chelicera, upper and lower views (MCTP 0229); B. Left female chelicera, upper view (MCTP 43306 ex 0370); C. Left male palp, mesal view (MCTP 0229); D. Left male palp conduc¬ tor detail and pleats, mesal view (MCTP 0229); E. Tip of conductor detail, dorsal view (MCTP 43306 ex 0370); F. Left male palp paracymbium, ventral view (MCTP 0229); G. Epiandrous field, ventral view (MCTP 43306 ex 0370). Scale bars: 1 mm (A); 0.5 mm (B, C); 0.2 mm (D); 0.01 mm (E); 0.3 mm (F); 0.05 mm (G). tha tropica. GUATEMALA: $ holotype, Menche, God- man and Salvin, not located. Extended diagnosis. Females of T. elongata can be dis¬ tinguished from all other Neotropical species by their elongated body, abdomen anteriorly enlarged and much narrower posteriorly, large chelicerae with an outer cusp (OC), and a short genital fold (Figs 6A-G, 7B). The unique internal genitalia has large spermathecae with two thick tu¬ bular lobes connected mid-way, forming a kidney-shaped structure, with the median lobe more than twice as long as its width and parallel to each other and to the longitudinal axis of the abdomen (Fig. 6H). The lateral lobe is smaller and thinner than the median lobe and may vary in posi¬ tion, with the fundus directed dorsally (Fig. 6H) or later¬ ally (Levi 1981: fig. 76). Central membranous sac (CS) is small and poorly sclerotized (Fig. 6H; Levi 1981: fig. 76). Male chelicerae of T. elongata are similar to those of T. laboriosa Hentz, 1850, insofar as sharing elongated apophyses with excavated tips, elongated and robust‘T’, and long Gl, the last as the longest teeth of both lower rows (Figs 5D-G; Okuma 1992: fig. IIA-C). Tetragna- zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 465-505 477 tha elongata differs from T. laboriosa by its longer and narrower chelicerae (4.4x vs 3.5x longer than wide), ‘sf shorter, ‘T’ with larger basis, more elongated and distalward projected, a higher number of ‘rsu’ teeth (Figs 5D, 7A; Okuma 1992: 231, fig. 11 A), AXl stout¬ er, slender and distalward projected and G1 pointed and much more elongated, both arising from a common base and displaced to a lower position than the remaining lower teeth (Figs 5D-F, 7A; Okuma 1992: 231, fig. 11 A, B). Males can also be distinguished from congeners by the palps with conductors having triple pleats, enclosing the long filiform emboli, which bear small bird-head tips that are projected in small tails (Figs 5H-J, 7C-E, 20B), and by short and thick paracymbia that are perpendicu¬ lar to the palps axes and with undivided notches, narrow, thin translucent lobes, and thumb-like projecting knobs (Figs 5K, 7C, F). The epiandrous field has a large de¬ pressed lateral area at both sides of the spigots (Fig. 7G). Synonymy and notes. Tetragnatha tropica was described from a single female from Guatemala (Menche, currently Atxchilan, near the Mexican border), not located in NHM or OUMNH collections. Thus, we rely on the original il¬ lustrations and subsequent descriptions of both sexes to diagnose this species (O. Pickard-Cambridge 1889: pi. 2, fig. 3; F. O. Pickard-Cambridge 1903: pi. 40, figs 10, 11; Chickering 1957c: figs 97-102). We compared these il¬ lustrations with our specimens and the drawings of both sexes and SEM images of the conductor tip of T. elongata by Eevi (1981), who proposed a neotype for this species (MCZ 21192) and identified many specimens from north¬ ern Mexico, and with Okuma (1992), who identified spec¬ imens from Mexico, Costa Rica, and Panama. Based on our investigations, T. tropica can be diagnosed as a junior synonym of T. elongata because the morphology of male palps and the chelicerae of both sexes perfectly match. We also highlight that Mello-Eeitao (1943, 1945, 1947, 1949) repeatedly recorded T. elongata from Brazil and Argentina, but all his specimens we analysed belong to other common species, especially T. bogotensis and T. nitens. However, after analyzing many other specimens, we were able to confidently record this species for these two countries. Variation. Males (n = 8): total length, 7.20-13.45; fe¬ males (n = 15): total length, 8.88-13.90. Distribution. Tetragnatha elongata is a very common species in the Nearctic and Neotropical regions, recorded from Canada (Dondale et al. 2003) to Misiones, north¬ eastern Argentina (Fig. 22B). Tetragnatha keyserlingi Simon, 1890 Figs 8-10, 20C, 21H, I, L, M, O, 22C Tetragnatha mandibulata: Keyserling 1865: 848, pi. 21, figs 6-9 ((3 $ misidentified). Tetragnatha mandibulata?'. L. Koch 1872; 194, pi. 17, figs 2a, b, 3a, b (3' $ misidentified). Tetragnatha keyserlingi Simon 1890; 134 {S ?)• Tetragnatha mandibulata'. Thorell 1890: 221 misidentified) Tetragnatha maxillosa Thorell 1895: 139 ((3 $) syn. nov. Tetragnatha kochi Thorell 1895: 140 (3' $) syn. nov. Tetragnatha japonica Bdsenberg and Strand 1906; 177, pi. 15, fig. 409a-d(6' ?). Tetragnatha conformans Chamberlin 1924; 9, pi. 2, figs 13-15 {S ?)• Tetragnathapropioides Schenkel 1936: 89, fig. 31 ((3 ?). Tetragnatha ethodon Chamberlin and Ivie 1936: 64, pi. 17, figs 144- 146 (3' $) syn. nov. Type material. Tetragnatha keyserlingi. COEOMBIA [Neu Granada]: S $ syntypes, not located. Tetragnatha maxillosa-. INDONESIA: $ syntype, Java, not located; SINGAPORE: S $ syntypes (Coll. Workman), not lo¬ cated. Tetragnatha kochi S $ syntypes, FIJI (Ovalau), not located; TONGA, not located; SAMOA (Upolu) [2$ sub T mandibulata, Mus. Godeffroy collection, 1869 (NHRS-GUEI000069809)], examined (photos). Tetragnatha japonica'. JAPAN: 10(5', 5$ syntypes. Saga (Yunohama Mountain) (Coll. W. Dbnitz, 25.V.1881, SMF 4212-121), not examined; 3$ syntypes, 2(5' (misiden¬ tified), Osaka (ZMH), examined (photos). Tetragnatha conformans'. CHINA: $ holotype, S paratype, Suzhou [Soochow] on the labels, Kuliang on the publication (Coll. N. Gist Gee, USNM 865), examined (photos). Tetragnatha propioides'. CHINA: S $ syntypes, Sichuan, not located. Tetragnatha ethodom PANAMA: f holo¬ type, Barro Colorado Island (AMNH), not examined. Extended diagnosis. Males of T keyserlingi are similar to T elongata. Both species have a long body, a very long paturon, AXl and G1 placed on a common base and dis¬ placed to a lower position than the remaining lower teeth, and conductor tips not extended in tail-like projections (Figs 5A-J, 7A, C-E, 8A-I, lOA, C, D, 20B, C; Eevi 1981: pis 5g, i, 6h, i; Okuma 1992 sub T maxillosa'. fig. 11A-D, F). Tetragnatha keyserlingi may be distinguished from T elongata by having pairs of black dots on the pos- tero-dorsal region of the abdomen, ‘a’ placed closer to the external border of the paturon and directed upwards and outwards, ‘F present, ‘sT absent, a crest filling the gap from Gu to ‘T’, a large gap between ‘T’ and ‘rsu’, lower teeth onwards from E3 placed in a shallow concave row (Figs 5A-E, 7A, 8A-E, lOA). Palps of both species are also similar, but those of T keyserlingi have shorter and wider tibias (2.3 x vs 4.7x), conductors with one pleat that only partially enfold the ribbon-like and twisted emboli, and paracymbia that are elongated, boomerang-shaped with slanted, basally projecting knobs and clearly visi¬ ble translucent lobes (Figs 8H-J, lOC-E, 20C). The epi¬ androus field has a spinning area which is not as high and without the depressed lateral areas found in T elongata (Figs 7G, lOF). Female chelicerae have elongated Gu, U2, and U3, where Gu is set apart from U2 by a very large gap and is lo- zse.pensoft.net 478 Pedro de Souza Castanheira et al..: Taxonomy of Neotropical Tetragnatha cated on an upper crest (CRu) (Figs 9D, E, 1 OB). U2 and re¬ maining upper side teeth form a row displaced to lower side of paturon, following the slanted fangs closing (Figs 9D, E, lOB). AXl very small and located near G1 base and lower row with first three teeth on a lower crest (CRl); G1 straight, elongated, bulky, pointed, its base large, and projected slightly upwards (Fig. 9E, F). Cheliceral fang enlarged in middle portion and apical third slanted and tapering to the acute tip, also harboring a narrow ridge (Figs 9D, E, lOB). Internal genitalia unique, with two rounded spermathecae linked by two thick tubes to a median, slender and very elongated stalk, which places CS at a far anterior position (Fig. 91; Zhu and Zhang 2011, sub T. mcaillosa: fig. 123G). Synonymy and notes. This species is widespread in the Old World tropics and has been cited and illustrated many times under T. maxillosa (World Spider Catalog 2019), especially by Okuma (1983, 1987) and Zhu and Zhang (2011). Tetragnatha keyserlingi was named by Simon (1890: 134) for the specimens described and illustrated as T. mandibulata by Keyserling (1865: 848, pi. 21, figs 6-9) from “Neu Granada” (currently Colombia) and T. man¬ dibulata? by E. Koch (1872: 194, pi. 17, figs 2, 3) from Fiji (Ovalau), Samoa (Upolu), and Tonga in the Pacif¬ ic. The illustrations by Keyserling (1865) and E. Koch (1872) already clearly show, for example, the large ‘T’ and small ‘F in male chelicerae and the characteristic Gu and G1 of the female chelicerae of T. keyserlingi (Figs 8D, E, G, 9D-F, lOA, B). Tetragnatha keyserlingi was also recorded from Java (Indonesia) (Thorell 1890: 221) and in his paper on Bur¬ mese spiders, Thorell (1895: 139) newly named this spe¬ cies as T maxillosa, based on the female from Java he described in 1890 and on males and females from Sin¬ gapore collected by Cel. Workman; he pointed out that he considered T maxillosa to be distinct from T keyser¬ lingi. One page later, he (Thorell 1895: 140) named the specimens reported by E. Koch (1872) from the Pacific islands as another new species: T kochi, heavily relying on geographical distributions to separate the specimens from South America, Southeast Asia, and the Pacific Is¬ lands. The only morphological differences mentioned in his paper (Thorell 1895: 139-140) are minor details in eye position and cheliceral teeth arrangement in females; no male characters were mentioned. In particular for T max¬ illosa and T keyserlingi, he compared the position of the second tooth on upper row of female chelicerae (U2 in our terminology) in relation to teeth of the lower row. Accord¬ ing to Thorell, U2 of T maxillosa would face the 6* or 7* tooth of the lower row, while in T keyserlingi it would face the 4* or 5* tooth. However, the apparent position of teeth of one row in relation to the teeth of the opposite row is not easy to evaluate, as small changes on chelicerae in¬ clination may change the apparent alignment. In addition, small changes in the relative position of teeth of the upper and lower rows are very common in this and other species. We agree with Simon (1890) that the detailed illustra¬ tions given by Keyserling (1865) from “Neu Granada”, and E. Koch (1872), from Pacific Islands (Fig. 21H), al¬ low the clear recognition of just one species, despite the great distances between localities. Moreover, the speci¬ mens from Southeast Asia that Thorell (1895) named T maxillosa also belong to the same widespread species. As Simon (1890) and Thorell (1895) had never seen the specimens they named as new species, no type specimens have ever been designated. No type or non-type material from the type localities of T keyserlingi or T maxillosa could be located in NHM, NMV, and ZMH. However, we received photos of two females from Upolu, collect¬ ed in 1869 and deposited at NHRS, both identified as T mandibulata (labeled Museum Godefifoy, NHRS-GU- EI000069809). These specimens are surely part of the material which E. Koch (1872) designated and illustrated as “E mandibulata?” (Fig. 21H) and Thorell (1895) after¬ wards named T kochi. Thus, these specimens represent part of the type series of T kochi. As T keyserlingi Simon, 1890 was proposed five years before both ThorelFs names (1895), it is senior synonym of T maxillosa and T kochi. Bosenberg and Strand (1906: 177, pi. 15, fig. 409a-d) provided comprehensive illustrations of male and female chelicerae and genitalia of T japonica. Based on photos of the chelicerae of syntypes of both sexes from Osaka (Japan) deposited at ZMH (Fig. 2H-K), we confirm that the female is T keyserlingi (Fig. 211) and that the male actually belongs to T nigrita Eendl, 1866 (Fig. 21J, K). Nonetheless, the illustrations of the male by Bosenberg and Strand (1906: fig. 409c, d) are like T keyserlingi, with a laterally directed apophysis, large Gu, absent or not noticeable ‘F, and apical portion of embolus and con¬ ductor curved in a gentle slope. Thus, we consider that at least the originally illustrated male syntype belongs to T keyserlingi and that it may be deposited in SMF instead. The female specimen of T japonica should be considered as name bearing, as it was described, measured, and il¬ lustrated first by Bosenberg and Strand (1906: fig. 409a, b). We agree here with Okuma’s synonymy (1983) of T japonica with T maxillosa (= T keyserlingi) and consider the male syntype from Hamburg and any other possible similar male syntype as misidentified specimens of T nigrita. Unfortunately, we were not able to examine the syntypes from Saga deposited in SMF. Chamberlin (1924: 12, pi. 3, figs 21-23) described T conformans (Fig. 2IE, M) and T cliens Chamberlin, 1924 (Fig. 21N-Q), each based on a couple from Su¬ zhou [Soochow on the labels (Fig. 2IP, Q), but Kuliang (Fuzhou) in the original paper], China. Eater, Schenkel (1936: 89-91, fig. 31) described T propioides based on a couple from Sichuan, also in China. All were separately synonymised with T keyserlingi (under T maxillosa or its junior synonyms): Zhu (1983) for T conformans (sub T japonica) and Okuma (1983: 72, 73) for T japonica, T cliens, and T propioides. Another mismatching of T keyserlingi and T nigrita occurred with the type series of T cliens, as observed by Song (1988: 127), who removed T cliens from the syn¬ onymy with T maxillosa. The male holotype of T cliens zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 465-505 479 Figure 8. Tetragnatha keyserlingi Simon, 1890, male (UFRJ 1553). A. Dorsal habitus; B. Lateral habitus; C. Ventral habitus; D-G. Left chelicera: D. Upper view; E. Inner view; F. Lower view; G. Outer view; H-J. Left male palp: H. Mesal view; 1. Dorsal view; J. Ventral view (paracymbium). Scale bars: 2 mm (A, B, C); 0.5 mm (D, E, F, G); 0.2 mm (H, I, J). zse.pensoft.net 480 Pedro de Souza Castanheira et al..: Taxonomy of Neotropical Tetragnatha Figure 9. Tetragnatha keyserlingi Simon, 1890, female. A. Dorsal habitus (UFRJ 1351); B. Lateral habitus (UFRJ 1351); C. Ventral habitus (UFRJ 1351); D-G. Left chelicera (MCTP 14749); D. Upper view; E. Inner view; F. Lower view; G. Outer view; H, 1. Gen¬ ital area: H. Genital fold, ventral view (MCTP 14749); 1. Internal genitalia, cleared, ventral view (UFRJ 1504). Scale bars: 2 mm (A, B, C); 1 mm (D, E, F, G); 0.5 mm (H); 0.1 mm (I). zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 465-505 481 Figure 10. Tetragnatha keyserlingi Simon, 1890, SEM photos. A. Left male chelicera, upper and lower views (MCTP 43319 ex 6944); B. Left female chelicera, upper and lower view (MCTP 14749); C. Left male palp, mesal view (MCTP 43319 ex 6944); D. Left palp conductor tip, mesal view (ULRJ 1552); E. Left male palp paracymbium, ventral view (ULRJ 1552); F. Epiandrous field, ventral view (MCTP 43319 ex 6944). Scale bars; 0.5 mm (A, B); 0.1 mm (C, E); 0.02 mm (D); 0.05 mm (F). is clearly T. nigrita and was wrongly coupled with the fe¬ male, which belongs to T. keyserlingi (Fig. 21N, O). Song (1988) noted that the male was labeled as “type” and the female as “paratype” in the original vials in USNM (Fig. 21P, Q). Like Song (1988), we were also able to analyse the type material and agree with the synonymy. Nonetheless, this mismatching was not noticed by Oku- ma (1983), who probably did not analyse the type materi¬ al and, years before Song, wrongly synonymised T. cliens with T. maxillosa, surely based on the order of description and illustrations of the original paper. Summing up, we agree with the synonymies for T. conformans (Fig. 21L, M), T. japonica (Fig. 211) and T. cliens (Fig. 2IN, P), according to original descriptions and illustrations and zse.pensoft.net 482 Pedro de Souza Castanheira et al..: Taxonomy of Neotropical Tetragnatha type photos we received. Additionaly, we also confirm the synonymy of T. propioides (see Schenkel 1936: fig. 31), but its syntypes were not located. Finally, T. ethodon was described by Chamberlin and Ivie (1936: pi. 17, figs 144-146) based on specimens from both sexes collected in Panama. This species was redescribed by Chickering (1957c: 316, figs 27-31) who transferred Chamberlin and Ivie’s females to T. tenuis- sima O. Pickard-Cambridge, 1889 and pointed out that the rather damaged male holotype was the only known specimen of the species from Panama. Okuma (1992: 228, fig. 7) also redescribed and illustrated T. ethodon, expanding its distribution to Puerto Rico and Barbados and adding a new description for females. She pointed out that this species was very similar to T. maxillosa (= T. keyserlingi) and separated both species by its wider gen¬ ital fold. Comparing our specimens with previous illus¬ trations under T. maxillosa, we consider the differences pointed out by Okuma (1992) to fall within the observed intraspecific variations, and hereby synonymise T. etho¬ don with T. keyserlingi. Variation. Males {n = 11): total length, 5.29-7.28; fe¬ males {n = 14): total length, 6.59-9.03. In Okuma (1968 sub T. japonica, 1983,1992 under T. ethodon) and Bosen- berg and Strand (1906 under T. japonica. fig. 409a), there is no visible upper crest (CRu) or lower crest (CRl). In fact, female syntypes of T. kochi (Fig. 2IK), the paratype of T. cliens and holotype of T. conformans (Fig. 21M, O) do not have crests on the paturon. In contrast, other publications (e.g. Okuma 1987, 1988b, Zhu and Zhang 2011), the female syntype of T. japonica from Osaka (Fig. 211), and all female specimens we examined from South America clearly possess CRu and CRl. So far, we cannot disregard the possibility that the absence of crests in some of the illustrated specimens is real or simply an artefact of poor illustration. Distribution. Pantropical, including Africa, Asia, Poly¬ nesia, Central America, and Brazil (Fig. 22C). Tetragnatha mandibulata Walckenaer, 1841 Figs 11-13, 20D, 21R-X, 22D Tetragnatha mandibulata Walcksnasr 1841; 211 $). Tetragnatha confraterna Banks 1909: 207, pi. 6, fig. 34 {S $) syn. nov. Tetragnatha necatoria Tullgren 1910; 149, pi. 3, fig. 72 ((3) syn. nov. Tetragnatha petrunkevitchi Caporiacco 1947: 24 (6' $) syn. nov. Tetragnatha petrunkevitchi: Caporiacco 1948; 646 Tetragnatha infuscata Benoit 1978: 667, fig. 2D, E {S)\ Saaristo 2003: 23, figs 21 A, B, 25 (removed from syn. with T. boydi) syn. nov. Type material. Tetragnatha mandibulata. GUAM: (5'$ syntypes, Mariana Archipelago, not located. Tetragnatha confraterna'. COSTARICA: syntypes, Machuca (Coll. P. Biolley); Escazii (Coll. J. F. Tristan); 3(5' 1 $ 1 immature, Tiribi (Coll. J. F. Tristan, MCZ 79139) and 2(5', Esparte (Coll. P. Biolley, MCZ 20875), examined (photos). Tetragnatha necatoria. TANZANIA: S holotype. Pare (Coll. Kimela, NHRS-GUEI0000069808), examined (photos). Tetragnatha petrunkevitchi'. GUYANA: 2(5,5$, 6j syntypes, Konawaruk (“Conwarook”), Potaro-Siparuni (18.iii.l937, MZUF 527), examined; 1(5 syntype, Hyde Park, (Teorgetown, Demer- ara-Mahaica (18.iv.l936, MZUF 528), examined; 1(5, 2$, Ij syntypes (1(5,1 3j in the museum’s catalog), Garraway Eanding, Potaro-Siparuni (30.vi.l936, MZUF 529), exam¬ ined. Tetragnatha infuscata. SEYCHEEEES: S, holotype, Mahe (Coll. P. E. G. Benoit & J. J.Van Mol, 24.vi.1972, MRAC 143319), examined (photos). Diagnosis. For females, see the extended diagnosis of T bogotensis. Males can be distinguished by the elongated body (Fig. IIA-C), chelicerae with pointed undivided apophyses (Figs IID-G, 13A), Gu extremely distinc¬ tive, thick, bulky and distally projected, followed by a tiny U2, contiguous to Gu (Figs IID, E, 13A), two very small and almost connected AXl and G1 (Figs HE, F, 13B) and palps with shorter tibias (ca 3x longer than large); conductor tips projected, large, flattened and winglet-shaped (Figs IIH-J, 13E, F, 20D) and paracym- bia with divided notches and large membranous translu¬ cent lobes that constitute the mesal halves of the notch¬ es, as in T bogotensis and T nitens (Figs IK, 3D, IIK, 13E, G, 141, J, 16D-F). Differing from T bogotensis, the translucent lobes of the other two species fill more than half of the total width of the paracymbia (Figs IK, 3D, IIK, 13G, 141, J, 16F). Finally, T mandibulata can be distinguished from T nitens by the narrower lobes (Figs IIK, 13G, 141, J, 16F). Synonymy and notes. We identified many specimens of T mandibulata from northern to southeastern Brazil. Unfortunately we were not able to study the type materi¬ al of this species from Guam. It was not available at the MNHN and NHM and is likely lost. However, this well- known and widely distributed species has been repeated¬ ly redescribed and illustrated in many papers (e.g. Simon 1900; Gravely 1921; Okuma 1983, 1987). After comparing our specimens with illustrations from redescriptions, we noticed close similarities be¬ tween T mandibulata and T confraterna (Fig. 21R-V; Chickering 1957c: 312, figs 19-26; Okuma 1992: 223, fig. 4), similar to Okuma (1983), who pointed out that both species might be synonyms. Banks (1909) de¬ scribed T confraterna from Costa Rica and published a simple illustration of the male chelicerae. Eater on, Chickering (1957c) and Okuma (1992) provided better illustrations of chelicerae and genitalia of specimens from Panama and Costa Rica, respectively. Okuma (1992) also redescribed T confraterna based on many specimens from various localities in Costa Rica and re¬ affirmed its relationship with T mandibulata, acknowl¬ edging that T confraterna “may be barely distinguished from the latter by the female cheliceral fang” (Okuma 1992: 225). We received photos of five males and one zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 465-505 483 Figure 11. Tetragnatha mandihulata Walckenaer, 1841, male (UFRJ 1356). A. Dorsal habitus; B. Lateral habitus; C. Ventral hab¬ itus; D-G. Left chelicera; D. Upper view; E. Inner view; F. Lower view; G. Outer view; H-K. Left male palp: H. Mesal view with tibia; 1. Mesal view detail; J. Dorsal view; K. Ventral view (paracymbium). Scale bars; 2 mm (A, B, C); 1 mm (D, E, F, G); 0.5 mm (H); 0.2 mm (I, J, K). zse.pensoft.net 484 Pedro de Souza Castanheira et al..: Taxonomy of Neotropical Tetragnatha Figure 12. Tetragnatha mandibulata Walckenaer, 1841, female. A. Dorsal habitus (UFRJ 1552); B. Lateral habitus (UFRJ 1552); C. Ventral habitus (UFPU 1552); D-G. Left chelicera (UFRJ 1552); D. Upper view; E. Inner view. F. Lower view; G. Outer view; H, I. Genital area: H. Genital fold, ventral view (UFRJ 1552); I. Internal genitalia, cleared, ventral view (UFRJ 1124). Scale bars: 2 mm (A, B, C); 1 mm (D, E, F, G, H); 0.1 mm (I). zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 465-505 485 Figure 13. TetragnathamandibuIataWsLlckQnsLQY, 1841, SEM photos. A. Left male chelicera, upper view (UFRJ 1356); B. Left male chelicera, lower view (UFRJ 1356); C. Left female chelicera, upper view (UFRJ 1552); D. Left female chelicera, lower view (UFRJ 1552); E. Left male palp, mesal view (UFRJ 1356); F. Left male palp conductor detail, mesal view (UFRJ 1356); G. Left male palp paracymbium, ventral view (UFRJ 1356); H. Epiandrous field, ventral view (UFRJ 1356). Scale bars; 0.5 mm (A, B, C, D, E); 0.05 mm (F); 0.1 mm (G); 0.02 mm (H). female syntypes from two different vials, all bearing the diagnostic characters of T. mandihulata (Fig. 21R-V). Based on the characteristic morphology of males and females of T. mandihulata, we consider the small differ¬ ences pointed out in Okuma (1992) as representing in¬ traspecific variation, and we propose that T. confraterna is a junior synonym of T. mandihulata. Tetragnatha necatoria Tullgren, 1910 was based on a male specimen from the Pare Mountains in Tan¬ zania. After studying one photo of the upper row of teeth of this male holotype and the original illustration of the distal part of the left chelicera of this species (Fig. 21W; Tullgren 1910: fig. 72), we noticed the large Gu had the U2 adjoined, which is typical of T. zse.pensoft.net 486 Pedro de Souza Castanheira et al..: Taxonomy of Neotropical Tetragnatha mandibulata. So, T. necatoria is a junior synonym of T. mandibulata. We received photos of one of the many specimens of T. petrunkevitchi Caporiacco, 1947 from Potaro and Georgetown, Guyana. This female (MZUF 529) clearly belongs to T. nitens, but there are also four specimens of T. mandibulata in the same vial. Additionally, all speci¬ mens in vials MZUF 527 and MZUF 528 are T. mandib¬ ulata, but the male and only adult specimen (amongst 25 immatures) in the vial MZUF 530 also belong to T. nitens. As the illustrations of chelicerae by Caporiacco (1948: figs 51, 52) of male and female specimens that he had preliminarly described in 1947 also match well with T. mandibulata, we conclude this species is another junior synonym of T. mandibulata. Finally, based on photos of the male holotype of T. infuscata from the Seychelles (MRAC 143319), we observed that it also has the diagnostic characters of T. mandibulata (Fig. 21X). Saaristo (2003: figs 21A, B, 25; 2010: figs 27, 31) illustrated the genitalia of one male and one female from the same locality and placed this spe¬ cies in the synonymy of T. boydi (= T. bogotensis), but without any evidence for his claim. Ironically, Saaristo (1978: 121, figs 224-231) was the first to correctly iden¬ tify T. mandibulata from the Seychelles and gave reliable illustrations of the typical female chelicerae and much elongated genital fold. However, Saaristo (2003, 2010) unexplicably considered his former identification as er¬ roneous and attributed it to T. boydi instead. Hence, we correct this misidentification, removing T. infuscata from the synonymy with T. boydi, and consider it to be a junior synonym of T. mandibulata. Variation. Males {n = 6): total length, 7.12-11.89; fe¬ males (n = 20): total length, 7.54-11.63. Distribution. Known from Africa, Asia, Australia, Cen¬ tral America, the Caribbean, and South America (Brazil and Guyana) (Fig. 22D). Tetragnatha nitens (Audouin, 1826) Figs 14-16, 20E, 22E Eugnatha nitens Audouin 1826: 118, pi. 2, fig. 2 (;^). Eugnathapelusia Audoum 1826; 119, pi. 2, fig. 3 ((3 $). Tetragnatha nitens Walckenaer 1841: 209. Tetragnatha peruviana Taczanowski 1878: 142, pi. 1, fig. la-c (S ?) syn. nov. Tetragnatha andinaTdLCZdLnowski 1878: 144, pi. 1, fig. 2 ($); Eevi 1981: 291 (syn. rejected, see T. bogotensis). Tetragnatha peninsulana Banks 1898: 246, pi. 15, fig. \2 {S Levi 1981: 291 (syn. rejected, see T. bogotensis). Tetragnatha eremita Chamberlin 1924: 645, fig. 89, 90 ((3); Eevi 1981: 292 (syn. rejected, see T. bogotensis). Tetragnatha decipiens Badcock 1932: 13, fig. 9 ($) syn. nov. Tetragnatha haitiensis Bryant 1945: 408, fig. 37 (^^j; Eevi 198): 292 (syn. rejected, see T. bogotensis). Tetragnatha tullgreni Caporiacco 1947: 24 ($ preoccupied by T. tull- greni Eessert, 1915) Tetragnatha caporiaccoi Platnick 1993: 381 (replacement name of T. tullgreni) syn. nov. Tetragnatha nitens kullmanni Wiehle 1962: 379, figs 1-5, 6b, 9-11, 14, 15 (c5' $); Wunderlich 1992: 365 (syn. rejected, see T. bogotensis). Type material. Tetragnatha nitens-. EGYPT: 6' $ syntype, Rosetta, Markaz Rasheed, lost (Levi 1981). Tetragnatha peruviana. PERU: SS, syntypes: Lima, El Callao and Pacasmayo (191 syntypes. Coll. K. Jelski and J. Sztoleman, MIZ 225658-225682), exam¬ ined (photos). Tetragnatha decipiens'. PARAGUAY: 1 $ holotype, “Nanahua”, probably Nanawa, Presidente Hayes, 05.ii. 1927 [Coll. G. S. Carter and L. C. Beadle] (NHM), examined. Tetragnatha caporiaccoi. GUY¬ ANA: $ holotype, Alto Demerara-Berbice, xii.1931 (MZUF 532), examined. Diagnosis. See the extended diagnosis under T bogoten¬ sis for the diagnostic characters of T nitens. Synonymy and notes. Tetragnatha nitens was first de¬ scribed from Egypt, but its syntypes are lost according to Levi (1981). This common species has been diagnosed and redescribed many times, with plentiful illustrations of its body, chelicerae and genital morphology (e.g. L. Koch 1872; O. Pickard-Cambridge 1872; Simon 1898; F. O. Pickard-Cambridge 1903; Chickering 1957c; Okuma 1983,1987,1988b, 1992). Furthermore, it is a senior syn¬ onym for 17 species or subspecies according to the World Spider Catalog (2019). As it is easily mistaken for other large-bodied species, misidentified specimens are com¬ monly found in museum collections. Levi (1981: 291, 292), for example, listed 13 syn¬ onyms of T nitens, of which 11 were new. We high¬ light that T festina Bryant, 1945 was listed as a new synonym by Levi (1981) but was previously synony- mised by Chickering (1957b: 2), T aptans Chamber¬ lin, 1920 was a new synonym but not listed as such, and T eremita Chamberlin, 1924 was not included in the synonymic list of the World Spider Catalog (2019). Indeed, Levi (1981: figs 23-29) clearly misidentified at least some of the females he ascribed to T nitens, whose illustrations belong to T bogotensis instead (see above Synonymy and notes for T bogotensis). Based on the original illustrations, we are able to confirm the synonymy of the following taxa: T pelusia Audouin, 1826, T antiliana Simon, 1897, T vicina Simon, 1897, T galapagoensis Banks, 1902, T aptans Chamberlin, 1920, T seminola Gertsch, 1936, T steckleri Gertsch & Ivie, 1936, T elmora Chamberlin & Ivie, 1942, and T festina Bryant, 1945. On the other hand, four species are in turn synonyms of T bogotensis (see Synonymy and notes for that species). Another lapsus occurred with T nitens kullmanni from Sardinia, Italy (Wiehle 1962: 379, figs 1-5, 6b, 9-11, 14, 15). It was first synonymised with T nitens by Wunder- zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 465-505 487 Figure 14. Tetragnatha nitens (Audouin, 1826), male (MCTP 1426). A. Dorsal habitus; B. Ventral habitus; C-F. Left chelicera; C. Upper view; D. Inner view; E. Lower view; F. Outer view; G-J. Left male palp; G. Mesal view. H. Dorsal view; I. Ventral view (paracymbium); J. Paracymbium detail, ventral view. Scale bars, 2 mm (A, B); 1 mm (C, D, E, F); 0.5 mm (G, H, I); 0.2 mm (J). zse.pensoft.net 488 Pedro de Souza Castanheira et al..: Taxonomy of Neotropical Tetragnatha Figure 15. Tetragnatha nitens (Audouin, 1826), female. A. Dorsal habitus (UFRJ 1528); B. lateral habitus (UFRJ 1528); C. Che- licerae upper row and eyes (UFRJ 1528); D. Chelicerae lower row and maxilla (UFRJ 1528); E-L Left chelicera (UFRJ 1528); E. Upper view. F. Inner view. G. Lower view. H. Outer view. 1. Basal cusp detail, distal view; J-L. Genital area; J. SEM of genital fold, ventral view (MCTP 11555); K. Internal genitalia, cleared, ventral view (MCTP 43323 ex 7313); L. Internal genitalia varia¬ tion, cleared, ventral view (UFRJ 1528). Scale bars; 2 mm (A, B); 1 mm (C, D, E, F, G, H, J); 0.2 mm (I, K, L). zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 465-505 489 Figure 16. Tetragnatha nitens (Audouin, 1826), SEM photos. A. Left male chelicera, upper and lower views (MCTP 1618). B. Left female chelicera, upper and lower views (MCTP 11555). C. Epiandrous field, ventral view (MCTP 1618). D. Left male palp, bulb detail, mesal view (MCTP 5985). E. Left male palp, mesal view (MCTP 5985); F. Left male palp paracymbium, ventral view (MCTP 5985). Scale bars; 1 mm (A, B); 0.1 mm (C); 0.5 mm (D, E, F). zse.pensoft.net 490 Pedro de Souza Castanheira et al..: Taxonomy of Neotropical Tetragnatha lich (1992: 365), but we disagree with this synonymy and consider T. nitens kullmanni as a synonym of T. bogoten- sis (see Synonymy and notes for that species). Mainly based on our observations of the morphology of the chelicerae, we propose three new synonymies for T. nitens: T. peruviana from Peru, T. decipiens from Par¬ aguay, and T. caporiaccoi from Guyana. The type mate¬ rials of all species were studied; for T. peruviana images from MIZ were examined, and for T. decipiens and T. caporiaccoi specimens were studied on visits to NHM and MZUF, respectively. Tetragnatha decipiens and T. caporiaccoi were de¬ scribed only from females, while descripitions of T. pe¬ ruviana were based on males and females, with males de¬ scribed first, even though only females were illustrated. Females of the three species and males of T. peruviana clearly show the diagnostic characters of T. nitens (see Diagnosis above). Variation. Males (n = 23): total length, 7.22-9.04; fe¬ males (n = 30): total length, 7.42-11.44. The internal gen¬ italia of females is also variable, with CS stalk shorter or longer than CS head, which places this structure at the same level or anterior to the spermathecae, which also vary in size (Fig. 15K, L; Zhu and Zhang 2011: fig. 125G) Distribution. This species was first described from Af¬ rica (Egypt), but it has a cosmopolitan distribution, with many new records from the Neotropics (Fig. 22E). Tetragnatha vermiformis Emerton, 1884 Figs 17-19, 20F, 22F Type material. UNITED STATES OF AMERICA: 6' lectotype, $ paralectotype (Eevi 1981). $ lectotype, 3$ paralectotypes in MCZ database, Beverly, Essex, Massa¬ chusetts (Coll. J. H. Emerton, 15.xiii.l8xx), not examined. Extended diagnosis. Males and females of T. ver¬ miformis are most similar to T. pallescens F. O. Pick- ard-Cambridge, 1903. Males have similar length and width of chelicerae (Figs 17D-G, 19A, Banks 1892: 51, pi. 5, fig. 88 as T. pallida., Okuma 1992: 236, fig. 16A-E); ‘T’ and ‘rsu’ very alike, sclerotized and point¬ ed; G1 very sclerotized, thick and pointed, with a very large base, remaining teeth set apart by similar gaps. They also share similar elongated paracymbia, with finger-like notches and straight lateral knobs (Figs 17J, 19E; Okuma 1992: 236, fig. 16D). The epiandrous field sets this species apart as it is flat and wide, with 20 fu- sules in two bands (Fig. 19F). Females of both species have similar small, rounded and laterally bulging chelicerae (Figs 18D-G, 19B; Oku¬ ma 1992: 236, fig. 16F, G); Gu isolated from U2; G1 from E2 by large gaps, with all teeth very pointed; and simi¬ lar short genital folds (Fig. 18H; Okuma 1992: 236, fig. 16J). Nonetheless, males and females of T. vermiformis differ from T. pallescens in having eyes much smaller and delicate and abdomen not as long and projecting (Figs 17A-C, 18A-C; Okuma 1992: 236, fig. 16H, I). Males differ by the following characters: absence of ‘sF (Figs 17D-F, 19A); ‘a’ bending downward and closer to fang base (Figs 17D-G, 19A); Gu not so close to fang base, larger and with thicker base (Figs 17D, E, 19A); Gu and ‘T’ placed apart from the row proper, towards lower side and following fang’s closing (Figs 17D, E, 19A); G1 with wider basis and close to AXl and fang ba¬ sis (Figs 17E, F, 19A); presence of an inner cusp on fang (Figs 17D, F, 19A) and more elongated conductors, with thicker projected tips completely enfolding the emboli, not ending in long tails (Figs 17H-J, 19C, D, 20F). Fe¬ males differ in lacking both AXu and AXl and having smaller and triangular Gu, longer and wider U2, and lack of a small denticle and groove near the base of E2 (Figs 18D-F, 19B; Okuma 1992: 236, fig. 16G). Females of T. vermiformis and T. pallescens have similar internal genitalia, with two curved kidney-shaped spermathecae on edge of plate, lacking central membranous sacs (Fig. 181; Eevi 1981: 311, fig. 131). However, T. vermiformis has longer spermathecae, without a median membranous area (Fig. 181). Variation. Males {n = 4): total length, 6.29-7.29; females {n = 7): total length, 6.99-10.98. The spermathecal lobes are variable in size and form. Both lobes may be more regularly cylindrical and the external lobe may be much smaller than the inner one (Eevi 1981: figs 178-180) or both lobes may be curved and about the same size (Fig. 181; Zhu and Zhang 2011: fig. 133H). Distribution. Temperate and tropical Asia, North and Central America, newly recorded from South America (Brazil) (Fig. 22F). Tetragnatha major Holmberg, 1876 and Tetragnatha riparia Holmberg, 1876 Remarks. Tetragnatha major and T riparia were de¬ scribed by Holmberg (1876), but the original descriptions are very short, without any suitable characters to correct¬ ly diagnose the species. There is a lack of illustrations and no type materials are specified because Holmberg did not collect the specimens he described. No specimens labeled as types of either species were found in MACN during our visit to that collection; Galiano and Maury (1979) had previously determined this. Despite the lack of in¬ formation on these two species, both were subsequently reported from many localities from Argentina (e.g. Mel- lo-Eeitao 1941, 1942). However, we re-examined many of these specimens at the MACN and MNRJ and they belong to either T argentinensis or T nitens. Therefore, we treat both species as nomina dubia. zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 465-505 491 Figure 17. Tetragnatha vermiformis Emerton, 1884, male (UFRJ 1556). A. Dorsal habitus. B. Lateral habitus. C. Ventral habitus; D-G. Left chelicera: D. Upper view; E. Inner view; F. Lower view; G. Outer view; H-J. Left male palp: H. Mesal view; 1. Dorsal view; J. Ventral view (paracymbium). Scale bars, 2 mm (A, B, C); 0.5 mm (D, E, F, G); 0.2 mm (H, I, J). zse.pensoft.net 492 Pedro de Souza Castanheira et al..: Taxonomy of Neotropical Tetragnatha Figure 18. Tetragnatha vermiformis Emerton, 1884, female. A. Dorsal habitus (UFRJ 1556); B. Lateral habitus (UFRJ 1556); C. Ventral habitus (UFRJ 1556); D-G. Left chelicera (UFRJ 1556); D. Upper view; E. Inner view; F. Lower view; G. Outer view; H, 1. Genital area: H. Genital fold, ventral view (UFRJ 1556); 1. Internal genitalia, cleared, ventral view (MCTP 43339 ex 11333). Scale bars: 2 mm (A, B, C); 0.2 mm (D, E, F, G, H, I). zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 465-505 493 Figure 19. Tetragnatha vermiformis Emerton, 1884, SEM photos (MCTP 43339 ex 11333). A. Eeft male chelicera, upper and lower views; B. Eeft female chelicera, upper and lower views; C. Eeft male palp, mesal view; D. Eeft palp conductor detail, mesal view; E. Male palp paracymbium, ventral view; F. Epiandrous field, ventral view Scale bars: 0.5 mm (A, B, C); 0.05 mm (D); 0.1 mm (E); 0.02 mm (F). Discussion In the literature the importance of the chelicerae for cor¬ rectly diagnosing species has often been ignored. Old publications (e.g. Audouin 1826; Walckenaer 1841; Emerton 1884) usually had short descriptions, basical¬ ly describing the body size, shape of abdomen, and eye length. On the other hand, some later authors like F. O. Pickard-Cambridge (1903), Chickering (1957c), Levi (1981), Okuma (1983, 1987, 1988a, 1988b, 1992) and Gillespie (1992a, 1992b), paid attention to chelicerae morphology and also genitalic features. Levi (1981), for example, based most of his determinations only on genital morphology, because he believed intraspecific variation on chelicerae was too high to allow clearcut species separation. In contrast, Okuma (1983, 1987, 1988a, 1988b, 1992) based her determinations mostly on chelicerae morphology, with additional drawings of male palps and genital folds. Gillespie (1992a, 1992b) followed Okuma, but also paid attention to leg spines on some Tetragnatha. In the same way, we also heavily rely on cheliceral features for species diagnoses, but add genitalic and other somatic characters whenever pos¬ sible. In our opinion, taxonomy in Tetragnatha cannot zse.pensoft.net 494 Pedro de Souza Castanheira et al..: Taxonomy of Neotropical Tetragnatha Figure 20. Embolous and conductor tip detail, dorsal view, SEM photos. A. T. bogotensis (MCTP 4299); B. T. elongata (MCTP 43306 ex 0370); C. T. keyserlingi (MCTP 43319 ex 6944); D. T. mandibulata (UFRJ 1356); E. T. nitens (MACN 2252); F. T ver- miformis (MCTP 43339 ex 1133). Scale bars; 0.05 mm (A, B, C, E, F); 0.02 mm (D). zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 465-505 495 P yEnsPFr. BERLIN A new species of Aphyocharax Gunther, 1868 (Characiformes, Characidae) from the Maraca 9 ume river basin, eastern Amazon Ptoella Silva de Erick Cristofore Guimaraes^’^’^, Luis Fernando Carvalho-Costa^, Felipe Polivanov Ottoni^’^’^’^ 1 Universidade Federal do Maranhao, Programa de Pos-Graduagao em Biodiversidade e Biotecnologia da Amazonia Legal. Av. dos Portugueses 1966, Cidade Universitdria do Bacanga, CEP 65080-805, Sdo Luis, MA, Brasil 2 Universidade Federal do Maranhao, Departamento de Biologia, Laboratorio de Genetica e Biologia Molecular, Av. dos Portugueses 1966, Cidade Universitdria do Bacanga, CEP 65080-805, Sdo Luis, MA, Brasil 3 Universidade Federal do Maranhao, Programa de Pos-Graduagao em Biodiversidade e Conservagdo. Av. dos Portugueses 1966, Cidade Universitdria do Bacanga, CEP 65080-805, Sdo Lius, MA, Brasil 4 Universidade Federal do Maranhdo, Laboratorio de Sistemdtica e Ecologia de Organismos Aqudticos, Centro de Ciencias Agrdrias e Ambientais, Campus Universitdrio, CCAA, BR-222, KM 04, S/N, Boa Vista, CEP 65500-000, Chapadinha, MA, Brasil 5 Universidade Federal do Maranhdo, Programa de Pds-graduagdo em Ciencias Ambientais, Centro de Ciencias Agrdrias e Ambientais, Campus Universitdrio, CCAA, BR-222, KM 04, S/N, Boa Vista, CEP 65500-000, Chapadinha, MA, Brasil http://zoobank.org/CFA54088-AFEE-4CFC-9187-AC9AE2E21A21 Corresponding author: Erick C. Guimardes (erick.ictio@yahoo.com.br) Academic Qditor: Nicolas Hubert ♦ Received 18 June 2019 ♦ Accepted 3 September 2019 ♦ Published 23 October 2019 Abstract A new species of Aphyocharax is described from the Maraca^ume river basin, eastern Amazon, based on morphological and mo¬ lecular data. The new species differs from all its congeners, mainly by possessing the upper caudal-fin lobe longer than the lower one in mature males, and other characters related to teeth counts, colour pattern, and body depth at dorsal-fin origin. In addition, the new species is corroborated by a haplotype phylogenetic analyses based on the Cytochrome B (Cytb) mitochondrial gene, where its haplotypes are grouped into an exclusive lineage, supported by maximum posterior probability value, a species delimitation method termed the Wiens and Penkrot analysis (WP). Key Words Freshwater, integrative taxonomy. Neotropical ichthyology, sexual dimorphism Introduction The Neotropical fish genus Aphyocharax Gunther, 1868 is distributed along the river basins of the Orinoco, Am¬ azon, and La Plata systems, as well as in the river sys¬ tems drainaing the Guiana Shield (Gery 1977; Taphorn and Thomerson 1991; Tagliacollo et al. 2012; Brito et al. 2018; Fricke et al. 2019), with highest diversity in the Amazon basin (Fricke et al. 2019). According to Brito et al. (2018), the genus comprises 11 valid species: Aphy¬ ocharax agassizii (Steindachner, 1882), A. anisitsi Eigen- mann & Kennedy, 1903, A. ovary Fowler, 1913, A. coli- fax Taphorn & Thomerson, 1991, A. dentatus Eigenmann & Kennedy, 1903, A. erythrurus Eigenmann, 1912, A. gracilis Fowler, 1940, A. nattereri (Steindachner, 1882), A. pusillus GfmfhQX, 1868, A. rathbuni Eigenmann, 1907, Copyright Pamella Silva de Brito etal. This is an open access articie distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the originai author and source are credited. 508 Pamella Silva de Brito et al.: A new species of Aphyocharax Gunther, 1868 and A. yekwanae Willink, Chernoff & Machado-Allison, 2003. However, there are at least four undeseribed spe¬ cies (Souza-Lima 2007). Tagliacollo et al. (2012) included seven valid species of Aphyocharax in their phylogenetics analysis, and pro¬ vided a hypothesis of interspecific relationships based on both molecular and morphological datasets. Their parsimony-based total evidence analysis (TE) indicates that Aphyocharax and Prionobrama Fowler, 1913 form a clade supported by three morphological synapomorphies: (1) interrupted lateral line with a single perforated scale on the posterior region of caudal peduncle; (2) absence or reduction of the fourth infraorbital bone canal; and (3) presence of a single large cusp on anterior maxillary teeth. In addition, three morphological synapomorphies have been proposed fox Aphyocharax: (1) narrow trigem- inofacialis foramen like a cleft with sphenotic almost ex¬ cluded from its margin; (2) dorsal projection of maxilla overlaping the second infraorbital; and (3) dorsal margin of third postcleithrum not projecting dorsally to poste¬ rior region of scapula (Mirande 2010; Tagliacollo et al. 2012). However, several other morphological features have been commonly used to characterize Aphyocharax species, such as the red caudal-fin colouration, moderate¬ ly elongated body, single series of tricuspid teeth on the premaxilla and mandible, and maxilla with teeth on up to two-thirds of its ventral margin (Taphom and Thomerson 1991; Willink et al. 2003; Tagliacollo et al. 2012; Brito et al. 2018). During recent fieldwork at the Maraca^ume river ba¬ sin, eastern Amazon, specimens of an additional unde¬ scribed species of Aphyocharax were collected and is herein described, based on both morphological and mo¬ lecular evidence, in accordance to an integrative taxono¬ my perspective. Methods Taxa sampling, specimens collection, and preservation Individuals collected for this study were euthanized with a buffered solution of Tricaine methanesulfonate MS-222 at a concentration of 250 mg/L for a period of 10 min or more until opercular movements completely ceased. Specimens selected for morphological analysis were fixed in 10% formalin and left for 10 days, after which they were preserved in 70% ethanol and specimens se¬ lected for molecular analysis were fixed, and preserved in absolute ethanol. Specimens for morphological analysis are listed in type and comparative material lists. Specimens for mo¬ lecular analysis are listed in Table 1. We also retrieved sequences from other species of Aphyocharax and al¬ lied genera for a comparative analysis from the National Center for Biotechnology Information (NCBI) databases (Table 1). Morphological analysis Measurements and counts were made according to Fink and Weitzman (1974) and Brito et al. (2018), except for the count of scale rows below lateral line, which were counted to the insertion of pelvic-fin. Vertical scale rows between the dorsal-fin origin and lateral line do not include the scale of the median pre¬ dorsal series situated just anterior to the first dorsal-fin ray. Counts of supraneurals, vertebrae, procurrent caudal-fin rays, unbranched dorsal and anal-fin rays, branchiostegal rays, gill-rakers, and teeth were taken only from cleared and stained paratypes (C&S), pre¬ pared according to Taylor and Van Dyke (1985). The four modified vertebrae that constitute the Weberian apparatus were not included in the vertebrae counts and the fused PUl + U1 was considered as a single element. Osteological nomenclature follows Weitzman (1962). Institutional abbreviations are: ANSP Acade¬ my of Natural Sciences, Philadelphia, Pennsylvania, USA; BMNH Natural History Museum, London, UK; CAS California Academy of Sciences, San Francisco, California, USA; CICCAA Cole^ao Ictiologica do Centro de Ciencias Agrarias Ambientais, Universidade Federal do Maranhao, Chapadinha, Brazil; FMNH Division of Fishes, Department of Zoology, Field Museum of Natural History, Chicago, Illinois, USA; LBP Laboratorio de Biologia e Genetica de Peixes, Departamento de Morfologia, Institute de Biociencias, Universidade Estadual Paulista “Julio de Mesquita Fil- ho”. Campus de Botucatu, Sao Paulo, Brazil; MNRJ Museu Nacional, Departamento de Vertebrados, Setor de Ictiologia, Universidade Federal do Rio de Janeiro, Rio de Janeiro, Brazil; UFRJ Cole 9 ao Ictiologica do Institute de Biologia, Universidade Federal do Rio de Janeiro, Rio de Janeiro, Brazil; UFRO Universidade Federal de Rondonia, Porto Velho, Brazil. DNA extraction, amplification, and sequencing DNA extraction was carried out with the Wizard Genom¬ ic DNA Purification kit (Promega) following manufactur¬ er’s protocol. DNA quality was evaluated by 0.8% aga¬ rose gel electrophoresis stained with GelRed (Biotium). DNA was stored in -20 °C until further procedures. Sam¬ ples (Table 1) were amplified using standard PCR (Poly¬ merase Chain Reaction) for partial Cytochrome B gene (CytB), using primers developed by Ward et al. (2005) (CytB2F 5' - GTG ACT TGA AAA ACC ACC GTT G-3' and CytB2R 5' - AAT AGG AAG TAT CAT TCG GGT TTGATG-3'). Amplification reactions were performed in a total volume of 15 pi comprising lx buffer, 1.5 mM MgCl 2 , 400 pM dNTP, 0.2 uM of each primer, 1 U of Taq Pol¬ ymerase (Invitrogen), 100 qg of DNA template, and ul- trapure water. The amplification program consisted of a denaturation of 94 °C for 3 min, followed by 35 cycles zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 507-516 509 Table 1. List of species, specimens and their respective GenBank sequence accession numbers, study in bold. Sequences made available by this Species Catalog number Genbank accession Aphyocharacidium bolivianum LBP9055-42219 HQ289710 Aphyocharax anisitsi LBP 25524 JQ820081 Aphyocharax anisitsi LBP3764-22190 HQ289581 Aphyocharax avary CICCAA2344-1 MK409660 Aphyocharax avary CICCAA2344-3 MK409661 Aphyocharax brevicaudatus sp. nov. (female) CICCAA02306 MK409668 Aphyocharax brevicaudatus sp. nov. (male) CICCAA02308 MK409669 Aphyocharax brevicaudatus sp. nov. (male) CICCAA02310 MK409670 Aphyocharax dentatus LBP 26163 JQ820082 Aphyocharax dentatus LBP 3604 JQ820083 Aphyocharax cf. erythrurus LBP 15819 JQ820076 Aphyocharax cf. erythrurus LBP 15820 JQ820077 Aphyocharax nattereri LBP 22345 JQ820070 Aphyocharax nattereri LBP 22132 JQ820071 Aphyocharax pusillus LBP 23546 JQ820078 Aphyocharax pusillus LBP4046-22920 HQ289590 Aphyocharax rathbuni LBP 36496 JQ820079 Aphyocharax rathbuni LBP 40434 JQ820080 Aphyocharax sp. LBP1587-11774 HQ289533 Aphyocharax sp. LBP 16349 JQ820084 Prionobrama paraguayensis LBP 19465 JQ820073 Prionobrama paraguayensis LBP 19468 JQ820072 Prionobrama filigera LBP 23664 JQ820075 Prionobrama filigera LBP 23663 JQ820074 Leptagoniates steindachneri LBP 4137-23661 HQ289600 Paragoniates alburnus LBP9208-43156 HQ289712 Phenagoniates macrolepis LBP6105-35623 HQ289678 Xenagoniates bondi LBP3074-19694 HQ289563 of 94 °C for 30 s, 46-48 °C for 45 s, and 72 °C for 80 s, and an extension phase of 5 min at 72 °C. Ampli- cons were visualised in 1% agarose gel electrophoresis stained with GelRed (Biotium) and purified with Illustra GFX PCR DNA and Gel Purification Kit (GE Health¬ care). Samples were sequenced using both forward and reverse primers and BigDye Terminator 3.1 Cycle Se¬ quencing kit in ABI 3730 DNA Analyser (Thermo Fish¬ er Scientific). Data partition, evolution models, and alignment The dataset included the partial Cytochrome B (CytB) mitochondrial gene (754bp). Sequences were aligned us¬ ing ClustalW (Chenna et al. 2003), and were translated into amino acids residues to test for the absence of pre¬ mature stop codons or indels using the program MEGA 7 (Kumar et al. 2016). Substitution Saturation tests were performed in DAMBE5 (Xia 2013) according to the al¬ gorithm proposed by Xia et al. (2003). The best-fit evo¬ lutionary model (GTR+G) was selected using Akaike In¬ formation Criterion (AIC) by iModelTest 2.1.7 (Darriba etal. 2012). Phylogenetic analysis A Bayesian inference-based phylogenetic (BI) tree was estimated in MrBayes (Huelsenbeck and Ronquist 2001) plugin in Geneious 9.0.5 to reconstruct the evolutionary relationships among terminals using General Time Revers¬ ible (GTR+G) as evolutionary model; and following pa¬ rameters: two Markov chain Monte Carlo (MCMC) runs of four chains each for 3 million generations and sampling frequency of 1,000. We used sequences of Aphyocharacid- ium bolivianum Gery, 1973, Leptagoniates steindachneri Boulenger, 1887, Paragoniates alburnus Steindachner, 1876, Phenagoniates macrolepis (Meek & Hildebrand, 1913), Prionobrama filigera (Cope, 1870), Prionobrama paraguayensis (Eigenmann, 1914), dindXenagoniates bon- di Myers, 1942 as outgroups. Species concept, species delimitation, and diagnoses The unified species concept is herein adopted by express¬ ing the conceptual definition shared by all traditional species concepts, “species are (segments of) separately evolving metapopulation lineages”, disentangling opera- zse.pensoft.net 510 Pamella Silva de Brito et al.: A new species of Aphyocharax Gunther, 1868 tional criterion elements to delimit taxa from species con¬ cepts (de Queiroz 2005,2007). According to this concept, species are treated as hypothetical units and could be test¬ ed by the application of distinct criteria (species delim¬ itation methods) (de Queiroz 2005, 2007). It allows for any criterion to separately provide evidence about species limits and identities, independently from other criteria (de Queiroz 2005, 2007). However, evidence corrobo¬ rated from multiple operational criteria is considered to produce stronger support for hypotheses of lineage sep¬ aration (de Queiroz 2007; Goldstein and Desalle 2010), a practice called “integrative taxonomy” (Dayrat 2005; Goldstein and Desalle 2010; Padial et al. 2010). Two distinct and independent operational criteria for species delimitation, based on morphological and molec¬ ular data, were implemented here: the population aggrega¬ tion analysis (Davis and Nixon 1992) (hereafter PAA); and a tree-based method as proposed by Wiens and Penkrot (2002) (hereafter WP, following Sites and Marshall 2003). Population aggregation analysis (PAA) The PAA (Davis and Nixon 1992) is a character-based method, in which species are delimited by unique com¬ bination of morphological character states occurring in one or more populations (Costa et al. 2014). The mor¬ phological data was based on both examined material and literature (e.g. Gunther 1869; Cope 1870; Eigenmann and Kennedy 1903; Eigenmann and Ogle 1907; Fowler 1913; Eigenmann 1915; Fowler 1940; Gery 1977; Taphorn and Thomerson 1991; Britski et al. 1999; Souza-Eima 2003a, 2003b; Willink et al. 2003; Gongalves et al. 2005; Tagli- acollo et al. 2012; Brito et al. 2018). Wiens and Penkrot analysis (WP) The WP analysis was based on CytB haplotypes, sup¬ ported on the direct inspection of the haplotype tree gen¬ erated by the phylogenetic analysis having as terminals at least two individuals (haplotypes) of each focal spe¬ cies. In this method, the term ‘exclusive’ is used instead of monophyletic, as the term monophyly is considered inapplicable below the species level (Wiens and Penkrot 2002). Clustered haplotypes with concordant geographic distribution forming mutual and well supported clades (exclusive lineages) are considered strong evidence for species discrimination (absence of gene flow with other lineages). When haplotypes from the same locality fail to cluster together, there is potential evidence of gene flow with other populations (Wiens and Penkrot 2002). Statistical support for clades is assessed by the poste¬ rior probability, considered as significant values about 0.95 or higher (Alfaro and Holder 2006). When only one haplotype (specimen) from one putative population was available, the species delimitation was based on the exclusivity of the sister clade of this single haplotype. supported by significant values, allowing us to perform the test in populations with only one haplotype (Wiens and Penkrot 2002). In addition, the method allows recog¬ nition of non-exclusive lineages as species if their sister clade is exclusive and supported by significant values (Wiens and Penkrot 2002). Results Aphyocharax brevicaudatus sp. nov. http://zoobank.org/C5D86CB2-B51B-4B45-AFF7-6E483533B680 Figs 1, 2 Holotype. CICCAA 02293, (male) 35.9 mm SE, Brazil, Maranhao state, Maraca^ume municipality, Maraca 9 ume River, 2°3T4"S, 45°57T6"W; 29 Jun 2018, E.C. Guim- araes and PS. Brito. Paratypes. All from Brazil, Maranhao state: CICCAA 02294, 1 (female), 32.4 mm SE, CICCAA 02295, 35 (males), 20.9-31.7 mm SE,CICCAA 02296,94 (females), 21-32.1 mm SE, CICCAA 02297, 30 (females) C&S, 22.2- 30.8 mm SE, CICCAA 02312, 2 (males) C&S, 28.3- 32.1 mm SE, UFRJ 11746, 10 (female), 24.2-30.2 mm SE; all collected with holotype. Diagnosis (PAA). Aphyocharax brevicaudatus sp. nov. differs from all its congeners by possessing the upper lobe of the caudal fin longer than the lower lobe in ma¬ ture males (vs upper and lower lobes similar in length, see Figs 1, 2; Tagliacollo et al. 2012: flg.4). Additionally, the new species is distinguished from Aphyocharax avary and A. pusillus by having hyaline middle caudal-fin rays (vs black or dark brown middle caudal-fin rays, Brito et al. 2018: fig. 3); from Aphyocharax colifax,A. yekwanae, and A. rathbuni by having caudal-fin light red colouration never surpassing the vertical line of the adipose-fin (vs red colouration extending to the lateral midline of body, Willink et al. 2003: fig. 1); from A. gracilis by having a larger body depth at dorsal-fin origin (body depth), 24.5-29.2% SE (vs 20.1-20.6% SE); and from A. pusillus by having teeth along 2/3 of the maxillary extension (vs along proximal half of the bone, Brito et al. 2018: fig. 4). Description. Morphometric data is presented in Table 2. Body shape is generally fusiform, slightly elongate, greatest body depth slightly anterior to dorsal-fin base; dorsal body profile straight or slightly convex from snout to vertical through anterior nostrils; straight or slight¬ ly convex from posterior nostrils to tip of supraoccip- ital bone; straight or slightly convex from this point to dorsal-fin origin; slightly convex along dorsal-fin base; postdorsal profile straight from base of last dorsal-fin ray to adipose-fin origin; slightly concave from adipose-fin to end of caudal peduncle; ventral profile convex from snout to pelvic-fin insertion; straight or slightly convex from this point to anal-fin origin; straight along anal-fin zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 507-516 511 Table 2. Morphometric data (A^= 141) of the holotype and paratypes of Aphyocharax brevicaudatus sp. nov. from the Maracagume river basin. SD; Standard deviation. Holotype (Male) Paratypes (Male) N = 35 Mean SD Paratypes (Female) N = 105 Mean SD Standard length (mm) 35.9 20.9-35.9 26.6 - 21.0-32.4 28.0 - Percentages of standard length Depth at dorsal-fin origin (body depth) 25.4 24.5-28.7 25.9 1.0 25.6-29.1 26.3 0.8 Snout to dorsal-fin origin 53.1 51.9-55.6 52.6 1.1 51.8-54.5 52.1 0.7 Snout to pectoral-fin origin 23.2 23.0-27.7 23.9 0.9 22.6-25.2 23.5 0.6 Snout to pelvic-fin origin 46.3 45.1-49.4 45.2 0.6 44.2-47.1 44.6 0.9 Snout to anal-fin origin 67.4 63.9-68.6 64.4 0.9 64.0-68.5 64.2 0.7 Caudal peduncle depth 10.8 10.1-12.5 11.3 0.5 10.9-12.2 11.3 0.3 Caudal peduncle length 13.2 12.2-17.2 14.0 1.2 12.2-14.9 13.1 0.7 Pectoral-fin length 20.4 17.9-22.5 19.7 0.3 18.6-21.1 19.3 0.6 Pelvic-fin length 15.9 14.6-20.6 15.6 0.5 14.0-17.1 15.3 0.7 Dorsal-fin base length 11.6 9.5-13.4 11.3 0.5 10.8-13.0 11.8 0.5 Dorsal-fin height 23.1 21.2-24.8 22.4 0.5 20.8-24.0 22.3 0.7 Anal-fin base length 18.9 16.7-21.1 18.1 0.4 16.8-20.7 18.3 1.0 Eye to dorsal-fin origin 42.6 40.6-54.6 42.1 0.6 41.4-52.4 41.8 1.9 Dorsal-fin origin to caudal-fin base 47.6 46.5-49.5 46.5 0.7 46.4-49.4 46.5 0.7 Head length 24.0 22.3-26.6 24.0 1.7 22.3-24.9 23.1 0.6 Percentages of head length Horizontal eye diameter 30.2 28.7-36.0 31.4 1.5 29.5-34.8 31.6 1.4 Snout length 24.2 19.7-28.8 23.5 0.6 22.8-29.3 25.4 1.2 Least interorbital v/idth 36.8 32.7-38.9 34.1 0.1 32.9-37.0 11.1 1.1 Upper jaw length 34.2 31.9-37.3 33.4 0.2 32.7-39.9 33.9 1.4 Figure 1. Aphyocharax brevicaudatus sp. nov. a. CICCAA 02293, holotype (male), 35.9 mm SL; b. CICCAA 02294, paratype (female), 32.4 mm SL, Brazil; Maranhao state: Maracafume river basin. (Photographed by Erick Guimaraes). zse.pensoft.net 512 Pamella Silva de Brito et al.: A new species of Aphyocharax Gunther, 1868 Figure 2. Caudal-fin oiAphyocharax brevicaudatus, holotype, CICCAA 02293, (male). base; long snout, with its length larger than orbital di¬ ameter; five infraorbital bones; fourth infraorbital absent and sixth infraorbital reduced; posterior border of maxilla rounded, extending vertically through anterior margin of orbit, not reaching third infraorbital. All teeth unicuspid or tricuspid and lateral cusps, when present, much smaller; premaxillary teeth in one rows with 6(9), 7(23) tricuspid teeth; maxilla with 11(3), 12(12), 13(14), or 14(3) unicuspid teeth; dentary with 6 (2) or 7 (30) larger tricuspid teeth followed by 6(26) or 7(6) smaller tricuspid teeth. Scales cycloid and same size over entire body generally. Predorsal scales mostly regular, but sometimes irregular just posterior to supraoccipital and/or slightly anterior to dorsal-fin. Scales covering anterior third of caudal-fin, with up to two, three, or four scales beyond posterior margin of hypural plate. Lateral line interrupted; last scale on cau¬ dal-fin base, with 9+1 (12), 10+1 (74), 11+1(50), or 12+1(5). Longitudinal scales series including lateral-line scales 35(3), 36(3), 37(56), 38(49), or 39(30). Longitudinal scales rows between dorsal-fin origin and lateral line 5(1), 6(93) or 7(47). Horizontal scale rows between lateral line and pelvic-fin origin 4 (141), Axillary scale present. Scales in median series between tip of supraoccipital spine and dor¬ sal-fin origin 13+1(24), 14+1(65), 15+1(26), or 16+1(26). Circumpeduncular scales 13(18), 14(115), or 15(8). Dorsal-fin rays i+10(99) or ii+10(42). Dorsal-fin origin situated posterior to vertical through pelvic-fin insertion, near middle of body. First dorsal-fin pterygiophore main body located of 8* and 9* vertebrae. Adipose-fin pres¬ ent. Anal-fin i+14(20), iii+15(18), ii+16(61), iii+16(24), ii+17(10), iii+17(5), ii+18 (3). Anteriormost anal-fin pterygiophore inserting at 14* and 15* vertebrae. Anteri¬ or anal-fin margin slightly convex, with anteriormost rays more elongate and slightly more thickened than remaining rays, forming a distinct lobe. Remaining rays smaller with straight distal margin. Pectoral-fin rays i+9(8), i+10(113), or i+11(20). Tip of pectoral-fin not reaching pelvic-fin ori¬ gin, when adpressed. Pelvic-fin rays i+7(120) or ii+7(21). Tip of pelvic-fin not reaching anal-fin origin, when zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 507-516 513 -50 -48 -46 -44 -42 -40 -6 Figure 3. Type locality of Aphyocharax brevicaudatus sp. nov. Atlantic Ocean 500km Wlaraca^ume River basin adpressed. Caudal-fin with a sexually dimorphic pat¬ tern, described below (Fig. 1). Principal caudal-fin rays 10+9(130) or 10+10(11); dorsal procurrent rays 8(2), 9(3) or 10(27) and ventral procurrent rays 7(2), 8(3) or 9(27). Branchiostegal rays 4(32). Supraneurals 6(4) 7(27) or 8(1). Total vertebrae 31 (1), 32(30) or 33(1). Colour in alcohol. Ground colouration light brown to yellowish brown. Inconspicuous light brown to light gray stripe from humeral spot to caudal-fin base, more con¬ spicuous on posterior half. Humeral region with one con¬ spicuous dark brown to black humeral spot. Smaller dark brown or black chromatophores homogeneously scat¬ tered. Smaller dark brown or black chromatophores ho¬ mogeneously scattered along body, except on chest. Head ground colouration similar to trunk, with dark brown chromatophores present on jaws, tip of snout, opercle, and dorsal portion of head. Dorsal, adipose, anal, caudal, pectoral, and pelvic fins hyaline to light brown. Sexual dimorphism. Caudal-fin of mature males with upper lobe longer (about 2/3 longer) than lower one, while both cauldal-fin lobes have similar leght in females (Fig. 1). Gill glands were found in all analyzed mature males of Aphyocharax brevicaudatus sp. nov. and were always absent in females. They were always located on anteriormost portion of lower branch of first gill arch, extending posteriorly through variable number of gill fil¬ aments. Etymology. The name brevicaudatus is a contraction of the Latin words brevis meaning “short” and cauda mean¬ ing “tail”, an allusion to the shorter caudal-fin lower lobe in the mature males of the new species. Geographic distribution. Aphyocharax brevicaudatus sp. nov. is currently known only from a single locality, the Maracagume river basin, a small and isolated coastal river basin of the eastern Amazon region (Fig. 3). zse.pensoft.net 514 Pamella Silva de Brito et al.: A new species of Aphyocharax Gunther, 1868 Discussion Several authors supported Aphyocharax as a monophyletic genus within Aphyocharacinae (Mirande 2010; Oliveira et al. 2011, Tagliacollo et al. 2012, Betancur-R. et al. 2018, Mirande 2018) and also the sister-group relationship be¬ tween Aphyocharax and Prinobrama (e.g. Oliveira et al. 2011; Tagliacollo et al. 2012; Betancur-R et al. 2018). On the other hand, few studies focused on the intrage¬ neric phylogenetic relationships vy'idcuu Aphyocharax (e.g Tagliacollo et al. 2012), and its diversity is probably un¬ derestimated, with at least four undescribed species (Sou- za-Lima 2007) and several populations or species waiting for a taxonomic revision (Lima et al. 2013; Ohara et al. 2017; Brito et al. 2018). Aphyocharax brevicaudatus sp. nov. is described here based on two distinct criteria and assumptions (PAA and WP). As mentioned in the Diagnosis (PAA), Aphyocharax brevicaudatus sp. nov. is unique among its valid conge¬ ners possessing the upper lobe of the caudal fin longer than the lower lobe in mature males (Souza-Lima 2003b; this study). This feature is generally rare among species of Characidae (Mirande 2010). In our Bayesian inference phylogenetic analysis (Fig. 4), haplotypes of A. brevicaudatus sp. nov. clustered as an exclusive lineage with high node support (maximum posterior probability value) (WP). The hypothesis of this new species is strengthened from an integrative taxono¬ my perspective, since it was based on evidence obtained from two independent criteria of species delimitation (see Dayrat 2005; de Queiroz 2007; Goldstein and Desalle 2010; Padial et al. 2010). The closer relationship between A. brevicaudatus sp. nov. and A. ovary is recovered with maximum posterior probability value. However the relationship between this clade {A. brevicaudatus sp. nov. and A. ovary) and other congeners have low phylogentic resolution, and discus¬ sions related to the phylogenetic positioning of this clade would be speculative with the data at hand. Comparative material Aphyocharax avaty: ANSP 39217,1 (Holotype), Madeira River, about 200 miles east, Brazil. UFRO 018489,3, Gua- jara municipality, Rondonia state, Brazil. LfFRO016159, 62, Porto Velho municipality, Rondonia state, Brazil. UFRO 014317,7, Novo Aripuana municipality, Amazonas state, Brazil. MNRJ 10968, 11, Borba municipality, lago de Borba (Madeira River Basin), Amazonas state, Brazil. CICCAA 02394, 38, Sororo River, Maraba municipality, Para state, Brazil. Aphyocharax anisitsi. CICCAA 00867, 14, Pontes Lacerda municipality, Mato Grosso state, Bra¬ zil. CICCAA 01267, 6 C&S, Pontes Lacerda municipality, Mato Grosso state, Brazil. CAS 59697, 1, Asuncion mu¬ nicipality (radiograph and photograph of holotype), Par- HQ289710.1 Aphyocharacidlum bolManum 0.59 0.92 0.99 0.74 0.7 1 r MK409660 Aphyocharax avary \ MK409661 Aphyocharax avary ^ I MK409668 Aphyocharax brevicaudatus sp. n. MK409669 Aphyocharax brevicaudatus sp. n. MK409670 Aphyocharax brevicaudatus sp. n . 1 ^ JQB2008^Aphyocharax anisitsi ^ HQ289581.1 Aphyocharax anisitsi ^ JQ820082.1 Aphyocharax dentatus 1 0.8 1 t JQ820083.1 Aphyocharax dentatus JQ820084.1 Aphyocharax sp. HQ289533.1 Aphyocharax sp. 1 _r JQ820QB0A Aphyocharax rathbuni ^ JQ820Q79.^ Aphyocharax rathbuni JQ820077.1 Aphyocharax ct. erythrurus JQ820076.1 Aphyocharax ct. erythrurus HQ289590.1 A phyocharax pusillus 1 JQ820078.1 Aphyocharax pusillus JQ820070.1 Aphyocharax nattereh 0.84 JQ820071.1 Aphyocharax nattereh JQ820072.1 Phonobrama paraguayensis JQ820073.1 Phonobrama paraguayensis 1 _r JQ820075.1 Phonobrama filigera JQ820074.1 Phonobrama filigera HQ289678.1 Phenagoniates macrolepis 0.83 HQ289712.1 Paragoniates alburnus 0.99 " HQ289600.1 Leptagoniates dteindachneri HQ289563.1 Xenagoniates bondi 006 Figure 4. Bayesian inference tree including Aphyocharax brevicaudatus sp. nov. (red bar) and other congeners. Red arrow indicates the posterior probability of A. brevicaudatus node. Number above branches are posterior probability values. zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 507-516 515 aguay. Aphyocharax dentatus. ANSP 128718, 21, Lake Mozambique, Colombia. UFRJ 5571, 2, Rio Verde mu¬ nicipality, Mato Grosso do Sul state, Brazil. CAS 59722, 1, Laguna del Rio Paraguay (radiograph and photograph of holotype), Asuncion municipality, Paraguay. Aphyo¬ charax erythrurus: FMNH 53406,1, Rockstone sandbank (photograph of paratype), Guyana. Aphyocharax nattere- ri. UFRJ 5783,2, Pocone municipality, Mato Grosso state, Brazil. Aphyocharax pusillus: ANSP 178013, 4 (photo¬ graphs of recently preserved specimens), Rio Napo (Am¬ azon river basin), right bank just upstream from mouth of Mazan River, near town of Mazan, Loreto, Peru. BMNH 1867.6.13.46, 1 (syntype), Amazon river basin, Huallaga and Xeberos, Peru. BMNH 1867.6.13.58-59,2 (syntypes), Amazon river basin, Huallaga and Xeberos, Peru. BMNH 1869.5.21.10,1 (lectotype of Chirodon alburnus), Amazon River, Peru. BMNH 1869.5.21.11-13, 3 (paralectotypes of Chirodon alburnus), Amazon River, Peru. Aphyocharax rathbuni. CAS 76467,1 (Radiograph and photograph of a Holotype), Paraguay basin, Arroyo Chagalalina, Paraguay. Aphyocharax yekwanae: FMNH 109278, 1 (radiograph of paratype), Bolivarian Republic, Venezuela. Aphyo¬ charax sp.: CICC7C4 00865,11, Pontes e Lacerda munici¬ pality, Mato Grosso state, Brazil. CICC7C4 00865,4 C&S, Pontes e Lacerda municipality, Mato Grosso state, Brazil. Acknowledgements The authors thank James Maclaine for providing photo¬ graphs, x-ray images, and information on the type material of Chirodon alburnus and A. pusillus', Harry Taylor, the photographer of C. alburnus specimens, and Kevin Webb, the photographer of A. pusillus specimens; Mark Sabaj Perez for providing photographs of the A. pusillus', Rosana Souza-Lima for providing photographs and x-ray images of A. avary', Paulo Buckup, Cristiano Moreira, James Ma¬ claine, Carolina Doria, Wilson Costa, and Mark Sabaj Pe¬ rez for allowing us to examine material in their care; Paulo Petry, Francisco Provenzano, Oscar Miguel Lasso-Alcala, and Elias Costa Araujo Junior for providing useful litera¬ ture. CAPES and FAPEMA for providing the scholarship to PSB under the process 88887.159561/2017-00. This paper benefited from suggestions provided by P. Bragan^a and F. Roxo. 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BERLIN Anatomical redescription of Cyrenoidafloridana (Bivalvia, Cyrenoididae) from the Western Atlantic and its position in the Cyrenoidea Barbara L. Valentas-Romera\ Luiz R. L. Simone\ Paula M. Mikkelsen^, Rudiger Bieler^ 1 Museu de Zoologia da Universidade de Sdo Paulo. Laboratorio de Malacologia. Avenida Nazare, 481. CEP: 04263-000, Brazil 2 Integrative Research Center, Field Museum of Natural History, 1400 South Lake Shore Drive, Chicago, IL 60605, USA http://zoobank.org/A9E2CElF-6293-425F-AE4A-56468E7E02F9 Corresponding author; BarbaraL. fh/entos-i?o/wera (barbarella.lou@gmail.com) Academic editor; Z vow ♦ Received 1 August 2019 ♦ Accepted 17 September 2019 ♦ Published 24 October 2019 Abstract Members of the small bivalve family Cyrenoididae inhabit brackish waters of the eastern and western Atlantic Ocean. Cyrenoida flor¬ idana (Dali, 1896) from the western Atlantic is poorly known aside from shell descriptions. A detailed shell and anatomical study of C. floridana is here presented and compared with available data for Cyrenidae and Glauconomidae, two families of closest relation¬ ship according to recent phylogenetic studies. The species is characterized by valves externally covered by thin light brown periostra- cum; muscle scars and pallial line (without sinus) weakly impressed on the internal shell surface; a unique hinge pattern composed of cardinal and lateral teeth joining each other, right hinge with two laterals and two cardinals forming two inverted-V-shaped teeth and left hinge with two cardinals and one lateral forming a horizontal reversed F-shaped tooth; and microtubules inside the shell walls. Anatomically, the species presents unequal adductor muscles; demibranchs fused to each other along their posterior ends; a pair of totally fused, pigmented siphons; two pairs of siphonal retractor muscles; and a stomach with conjoined style sac and intestine, a single typhlosole, and three sorting areas. Evidence of shell parasitism is described. Key Words Anatomy, biodiversity, brackish water, Cyrenidae, freshwater, Glauconomidae, Heterodonta, Mollusca, taxonomy Introduction Cyrenoida Joannis, 1835, is the single genus of the small bivalve family Cyrenoididae and currently includes only six recognized extant species. Although previously as¬ sumed to extend into the Austral-Asian region (Pilsb- ry and Bequaert 1927), its known distribution includes temperate to tropical estuaries, marshes, and mangrove swamps of western Africa, both sides of the North Amer¬ ican continent, and the western coast of Panama and ad¬ jacent islands (Joannis 1835; Morelet 1851; Dali 1896; Pilsbry and Zetek 1931; Morrison 1947; Huber 2010; MolluscaBase 2019). Because of its ecological position in the fringe area between freshwater and saltwater en¬ vironments, the group has been left out from larger-scale treatments of marine (e.g. Mikkelsen and Bieler 2007) and freshwater mollusks (e.g. Lydeard and Cummings 2019). Cyrenoida is poorly represented in museum col¬ lections, which, together with the fact that many of the original localities are difficult to access, has hindered de¬ tailed study of this taxon. Several nominal species were introduced for West African members of this genus. These have been interpreted, based on shell morphology, as belonging to two fairly wide- ranging species (Huber 2010), the type species Cyrenoida dupontia Joannis, 1835, described from Senegal and extending to the Congo River (with synonyms Cyrenella senegalensis Deshayes, 1855, and Cyrenoida rhodopyga Copyright Barbara L. Valentas-Romera etal. This is an open access articie distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the originai author and source are credited. 518 Valentas-Romera, B.L. et al..: Anatomical redescription of Cyrenoida floridana Martens, 1891), and C. rosea (d’Ailly, 1896), described from Cameroon (including nominal subspecies C. rosea brevidentata Pilsbry & Bequaert, 1927 from Senegal). The fossil record of the group is poorly known, with a Pliocene species from southern Florida, Cyrenoida caloosaensis (Dali, 1896), recognized by Campbell (1993), and the modem species Cyrenoida floridana (Dali, 1896) interpreted as extending to the middle Pleistocene of Florida (Portell and Kittle 2010) and the Holocene of southern Texas (Neck and Herber 1981). For the American Pacific coast (Coan and Valentich- Scott 2012), Cyrenoida panamensis Pilsbry & Zetek, 1931 was described from the western coast of mainland Panama (and is known from Costa Rica; Vargas-Zamora and Sibaja-Cordero 2011), and C. insula Morrison, 1946 from the Pearl Islands in the Gulf of Panama. Another two species occur in the western Atlantic region, Cyrenoida americana Morelet, 1851, described from Cuba (and with published records from Puerto Rico and the Bahamas; Dali 1905), and C. floridana (Dali, 1896), the focus of the current treatment. Although rare¬ ly studied, the latter is a wide-ranging species along the western Atlantic and Gulf of Mexico coasts and has been cited as a prey item for both fish and bird species (Heard 1975, 1982; Kat 1978). It can be found living infaunally in muddy and sandy sediment colonized by halophytic plants, in estuaries and waters surrounding river mouths (Dali 1896,1901; Kat 1982), the outer fringes of very low saline to freshwater marshes (Tunnell et al. 2010), as well as around freshwater ponds (this study). The position of the family Cyrenoididae within the Heterodonta remained unresolved for a long time, with most authors including it in a broad concept of Luci- noidea (e.g. Prime 1860; Dali 1901; Lamy 1920; Cha- van 1969; Yokes 1980; Boss 1982; Vaught 1989). Others placed it tentatively near the Corbiculidae (= Cyrenidae) (Thiele 1934) or as its own superfamily Cyrenoidoidea near groups such as Chamoidea and Galeommatoidea (Olsson 1961). A close relationship of Cyrenoididae with Lucinidae was questioned by Taylor and Glover (2006) and Williams et al. (2004) on anatomical grounds. The latter opinion was confirmed by Taylor et al. (2009) in the first molecular study that included a member of the fam¬ ily (C. floridana), which indicated a close relationship of Cyrenoididae with Cyrenidae (the latter as Corbiculidae) and Glauconomidae. They again elevated the rank to su¬ perfamily Cyrenoidoidea, which was adopted in some subsequent classifications (e.g. Bieler et al. 2010). Subse¬ quent studies with additional molecular markers (Sharma et al. 2012; Combosch et al. 2017, Lemer et al. 2019) and combined morphological and molecular datasets (Bieler et al. 2014) confirmed the close relationship of Cyrenoi¬ didae, Cyrenidae, and Glauconomidae. The latter work combined them in the superfamily Cyrenoidea within the Neoheterodontei (the crown group of Imparidentia), its current position. The recent transcriptomic study by Le¬ mer et al. (2019) included members of Cyrenidae \Cor- bicula fluminea (O. F. Muller, 1774) and Polymesoda caroliniana (Bose, 1801)], Glauconomidae {Glauconome rugosa Hanley, 1843), and Cyrenoididae {C. floridana). Interestingly, Polymesoda grouped with Glauconome, not Corbicula (which appeared as the basal taxon in this clade), indicating that Cyrenidae as currently understood (Bieler and Mikkelsen 2019) might not be monophyletic. Whereas morpho-anatomical studies on this group have been limited, molecular data of C. floridana have been in¬ volved in several analyses, including recent transcriptom¬ ic studies exploring questions ranging from synonymous codon usage bias (Gerdol et al. 2015) to Imparidentia phytogeny (Lemer et al. 2019). To improve the morpho¬ logical knowledge of Cyrenoididae and to contribute to the resolution in the Cyrenoidea clade, a detailed anatom¬ ical study of C. floridana is here presented. Its features are then compared to available data for members of Glau¬ conomidae and several genera of Cyrenidae sensu lato. Material and methods A detailed list of examined material is presented follow¬ ing the anatomical description. The anatomical study is divided in two parts: shell analysis and soft part analy¬ sis. The shell analysis consisted of measurements and scanning electronic microscopy of the shell. The shell measurements were taken using a caliper or, in case of photos, using Image! software. The measurements used were shell length, height, and width; umbo length and height; dental shelf length and height; hinge teeth length and height; dorsal shell margin length; adductor muscle length, height, and area; and pallial line spacing from the ventral shell margin. The soft part analysis collected de¬ tails of topology and morphology of systems and organs using photography and drawings under camera lucida. The soft part data were obtained from specimens pre¬ served in ethanol. Dissection occurred with the specimens immersed in 70% ethanol. Final drawings were initially made in graphite and later remade on translucent paper with China ink, scanned, and edited using Photoshop CS3 software. The final drawings are average anatomi¬ cal schemes based on information collected from several specimens. The number of specimens dissected varied ac¬ cording to the availability in collections and was expected to be sufficient to cover any feature affected by preser¬ vation methods, such muscular contractions, distensions limits, and presence or absence of delicate structures, or sexual stages of the specimens, and to detect maturation stages and gonadal filling. All soft parts here drawn are based on specimens in lot FMNH 328260. The type materials of C. floridana and C. guatemalen- sis were examined from photographs, whereas addition¬ al samples were physically studied. Scanning electron microscopy (SEM) was provided by the Laboratorio de Microscopia Eletronica do Instituto de Biociencias of the Universidade de Sao Paulo and by the Eaboratorio de Microscopia Eletronica from Museu de Zoologia of the Universidade de Sao Paulo. zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 517-534 519 The following abbreviations are used in the ana¬ tomical descriptions and figures: aa: anterior adductor muscle; an: anus; ar: anterior pedal retractor muscle; an: auricle; cc: cerebral connective; eg: cerebral gan¬ glia; cn: ctenidial nerve; cp: cerebropedal connective; cv: cerebrovisceral connective; dd: digestive divertic¬ ula; dg: digestive gland; dh: dorsal hood; dm: dorsal siphonal retractor muscles; eo: excurrent opening; er: esophageal rim; es: esophagus; ex: excurrent siphon; fg: food groove; ft: foot; gf: gill fusion; gi: gill; go: go¬ nad; gp: genital pore; gs: gastric shield; he: heart; id: inner demibranch; if: mantle border inner fold; in: intes¬ tine; io: incurrent opening; ip: inner palp; is: incurrent siphon; ki: kidney; Ic: left caecum; Ip: left pouch; Iv: large inverted-V-shaped tooth; mf: mantle border mid¬ dle fold; mo: mouth; mt: major typhlosole; na: anterior adductor muscle nerve; np: nephropore; nt: minor ty¬ phlosole; od: outer demibranch; of: mantle border out¬ er fold; op: outer palp; pa: posterior adductor muscle; pg: pedal ganglia; pi: pallial line; pm: pallial muscle; pn: pallial nerve; pp: papillae; pr: posterior pedal re¬ tractor muscle; rc: right caecum; rn: renal nerve; sal: sorting area 1; sa2: sorting area 2; sa3: sorting area 3; sm: siphonal membrane; sn: dorsal siphonal muscle nerve; sp: siphons; ss: style sac; st: stomach; sv: small inverted-V-shaped tooth; tl: large lateral tooth of right valve; 12: large cardinal tooth of right valve; 13: small lateral tooth of right valve; 14: small cardinal tooth of right valve; 15: lateral tooth of right valve; 16: posterior cardinal tooth of left valve; 17: anterior cardinal tooth of left valve; ve: ventricle; vg: visceral ganglia; vm: ven¬ tral siphonal retractor muscles. Institutional abbreviations: ANSP, Academy of Nat¬ ural Sciences of Drexel University, Philadelphia, Penn¬ sylvania, USA; UF, Florida Museum of Natural History, Gainesville, Florida, USA; FMNH, Field Museum of Natural History; USNM, National Museum of Natural History [United States National Museum], Smithsonian Institution, Washington, DC, USA. Systematics Family Cyrenoididae H. Adams & A. Adams, 1857 (1853). Synonym: Cyrenellidae J.E. Gray, 1853. Genus Cyrenoida Joannis, 1835 (type species by monotypy: Cyrenoida dupontia Joannis, 1835). Syno¬ nyms include: Deshayes, 1836 (established in synonymy of Cyrenoida, available because it was used as valid before 1960, e.g. by Gray 1853); Cyrenoidea Dali, 1896 (unjustified emendation; the earlier use by Hanley (1846) is considered an incorrect subsequent spelling); Cyrenodonta has been credited by some au¬ thors to H. Adams & A. Adams, 1857, but it was in¬ troduced in synonymy of Cyrenoida and no pre-1960 use as a valid name has been located); Cyrenoides auct. is an incorrect subsequent spelling for Cyrenoida (G.B. Sowerby II, 1839). Cyrenoida floridana (Dali, 1896) Figs 1-41 Cyrenoidea floridana Dali 1896; 52; Simpson 1887-1889; 66 [nomen nudum]; Dali 1889; 50, 208 [nomen nudum]; Rhoads 1899; 48; Heard 1975; 22; 1982a; 23, fig. 24; 1982b; 131. Cyrenoida floridana —Dali 1901; 817, pi. 42, fig. 7; Lamy 1920; 388; Pilsbry andZetek 1931; 69; Smith 1951; 45 (pi. 16, fig. 11, pi. 18, fig. 8); Pulley 1952; 114-115, pi. 9, fig. 15; Morrison 1954; 9-10; Van RegterenAltena 1968; 157,176; 1971; 5,41, fig. 14; Waas 1972; 123; Abbott 1974; 466 (fig. 5383); Leathern et al. 1976; 93, figs 1-3; Kat 1978; 1-168, figs 1-91, tables 1-7; A1-A6; Neck and Herber 1981; 35-39; Kat 1982; 47, figs 1-3 (oocytes); Heard 1982a; 25, fig. 28j; Yokes and Yokes 1983; 39, 62, pi. 39, fig. 7; Neck 1985; 5; Bishop and Hackney 1987; 141, fig. 6; Turgeon et al. 1988; 36; 1998; 39; Camp et al. 1998; 11; Abbott and Morris 1995; 53, pi. 24, fig. 12; Red- fern 2001; 219, pi. 92, fig. 898; Reece et al. 2004; 1116; Mikkelsen and Bieler 2000; 373; 2004a; 513; 2004b; 596; Lee 2009; 28, fig.; Turgeon et al. 2009; 728; Taylor et al. 2009; 10 (figs 4—8); Tunnell et al.2010; 345;Redfem 2013; 400, fig. 1067; Bieler etal. 2014; 45 (fig. 3N); Arzul and Carnegie 2015; 33; Gonzalez et al. 2015; 4, figs 1, 2; Combosch et al. 2017; table 1, figs 1, 2; Lemer et al. 2019; figs 1, 2. Cyrenellafloridana —Walker 1918; 88, fig. 232. Cyrenoidea guatemalensis Pilsbry 1920; 221 (pi. 11, fig. 9); Clench and Turner 1962; 60. Cyrenoida guatemalensis —Pilsbry and Zetek 1931; 69; Morrison 1946; 45. Description. Shell (Figs 11-23): External features: Out¬ line rounded, subcircular with ventral margin slightly pos¬ teriorly carinated (Figs 11, 12); equivalve, equilateral, ~6% longer than high, reaching maximum length of ~15 mm. Laterally inflated, width -59% of total shell length (Figs 13-15). Externally white, adorned only with growth lines, showing -3 thicker commarginal growth increments. Peri- ostracum thin, slightly wrinkled, light brown. Walls thin, fragile. Umbones prosogyrous, low, -5% of total shell height, large, length -25% of total shell length, located at midpoint of shell length. Ligament parivincular, opisthodet- ic, long, -39% of total shell length (Figs 15-17). Nymph long, -20 times longer than wide, rectangular. Lunule and escutcheon absent. Internal features: Internal surface opaque white (Figs 16,17). Adductor muscle scars and pal- lial line weak, very faintly impressed (outlined in Fig. 16). Anterior adductor muscle scar reniform, occupying -1.5% of total internal surface; ventral portion -2 times wider than dorsal portion; positioned at median third of valve height. Posterior adductor muscle scar oval, slightly pointed dorsal- ly, occupying -1.6% of total internal surface; located slight¬ ly more ventral than anterior muscle scar. Pallial line weak, formed by row of small pallial muscle scars, connected to middle portion of ventral surface of anterior adductor mus¬ cle to middle portion of ventral surface of posterior adductor muscle, inset from ventral shell margin by -19.5% of total shell height, without pallial sinus. Microtubules of elongat¬ ed conical shape, beginning with circular opening in interior shell wall, tapering toward but not reaching external surface (Figs 20-22). Internal surface of shell usually with aragonit- ic nodules of various sizes and quantities (Figs 16,17, 23). zse.pensoft.net 520 Valentas-Romera, B.L. et al..: Anatomical redescription of Cyrenolda floridana Figures 1-10. Syntypes of Cyrenoida floridana and lectotype of C. guatemalensis. 1-4. Cyrenoida floridana (USNM 46846, length 12 mm, height 13 mm). 1. Left valve, external view; 2. Right valve, external view; 3. Left valve, internal view; 4. Right valve, internal view; 5-10. Cyrenoida guatemalensis (ANSP 107532; length 8.6 mm, height 8.9 mm). 5. Left valve, external view; 6. Right valve, ex¬ ternal view; 7. Left valve, internal view; 8. Right valve, internal view; 9. Detail of left hinge; 10. Detail of right hinge. Scale bar: 2 mm. Hinge (Figs 16-19): Hinge restricted to central and anterior portion of dorsal margin, composed of lateral and cardinal teeth. Right hinge (Figs 17,19): Dental shelf short and wide, triangular, running along entire length of anterior portion of dorsal margin, height equivalent to ~10 times dorsal margin width; composed of two laterals and two cardinals. Each cardinal tooth joining posterior¬ ly with a lateral tooth, forming two inverted-V-shaped teeth, one large (Fig. 19: Iv), one small (Fig. 19: sv). Large V-shaped tooth located near dorsal shell margin, formed by long and laminar lateral tooth (Fig. 19: tl) and short cardinal one (Fig. 19: t2). Lateral tooth length equivalent to -56% of total dental shelf length, cardinal tooth length equivalent to -22% of lateral tooth length; small V-shaped tooth located ventral to large V-shaped tooth (Iv). Small lateral tooth (Fig. 19: t3) length -30% shorter than large lateral tooth, whereas small cardinal tooth (Fig. 19: t4) -40% shorter than dorsally located cardinal tooth (Fig. 19: t2). Left hinge (Figs 16, 18): Dental shelf narrow, fusiform, running along 30% of an¬ terior portion of dorsal margin, height equivalent to -3 times dorsal margin width; composed of three cardinal teeth, two cardinal and one lateral, forming horizontal reversed F-shaped tooth (Fig. 18: fs). Both cardinal teeth originating parallel and close to each other, (Fig. 18: t6, t7). Lateral tooth laminar (Fig. 18: t5), joining anterior cardinal tooth (Fig. 18: t7), length equivalent to 50% of total length of dorsal shelf. Both cardinal teeth equiva¬ lent to 20% of lateral tooth length. When articulated, left valve tooth complex (t5-t7) fits within groove between right valve t3-t4 and tl-t2. Muscular system (Figs 25-27, 31, 34, 33): Anterior adductor muscle (aa) reniform in cross section, -3 times taller than wide; ventral portion -2 times wider than dor¬ sal portion; occupying -3% of total shell internal volume; located at middle third of shell height; clearly divided into quick and slow components (Figs 25, 26, 35), quick com¬ ponent occupying -39% of anterior portion of muscle, dark grey in color, slow component occupying -61% of posterior portion of muscle, light cream in color. Posterior adductor muscle (pa) elliptical in cross section, -1.5 times wider than tall, -20% shorter and -2 times wider than an¬ terior adductor muscle, occupying -3% of total shell in¬ ternal volume; located slightly ventral to anterior adductor muscle; clearly divided into quick and slow components (Figs 25-27, 34, 35), quick component occupying -52% of posterior portion of muscle, dark grey in color, slow component occupying -48% of anterior portion of muscle, light cream in color. Paired anterior pedal retractor muscles (ar) oval in section, thin, attached on shell at posterodor- sal side of anterior adductor muscle insertion, area -3% of that of adductor, length -20% of shell length, left and right branches fused at mid-length. Paired posterior pedal re¬ tractor muscles (pr) oval in cross section, slightly laterally compressed, thin, -40% longer than anterior pedal retrac¬ tors; inserting on shell dorsally posterior adductor muscle, in area -3% of that adductor, left and right branches fus¬ ing at dorsal -20% of total muscle length. Pedal protractor zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 517-534 521 Figures 11-24. Cyrenoida floridana, shell and gills details. 11-22, 23. UF 246126; 24. FMNH 328260. 11. Left valve, external view; 12. Right valve, external view; 13. Dorsal view of shell; 14. Anterior view of shell; 15. Posterior view of shell; 16. Left valve, internal view, muscle scars and pallial line outlined; 17. Right valve, internal view; 18. Left hinge, SEM; 19. Right hinge, SEM; 20. Internal surface of shell, SEM, showing microtubule orifices; 21. Detail of microtubule patch; 22. Fractured shell showing microtubules partially through shell thickness; 23. Detail of nodules at shell internal surface; 24. Gill fragment, transverse section, SEM. Scale bars: 1 mm (11-17, 23); 200 pm (18,19); 20 pm (20-22), 0.5 mm (23); 10 pm (24). zse.pensoft.net 522 Valentas-Romera, B.L. et al..: Anatomical redescription of Cyrenoida floridana muscles absent. Two pairs of siphonal retractor muscles (Figs 27, 31, 33); dorsal siphonal retractors (dm) ~3 times longer than wide; insertion at mantle bifid for half of total muscle length, 2 times as long as excurrent opening, origi¬ nating laterally at half of siphonal base height; ventral siph¬ onal retractors (vm) thin and translucent, ~3 times longer than wide, length -50% of total length of dorsal siphonal muscle, originating at ventral end of incurrent siphon base. Foot (Figs 25, 26, 33): Foot short, wedged-shaped, length equivalent to -35% of total shell length, eontract- ed height equivalent to -27% of total shell height, later¬ ally compressed, with small heel of length equivalent to -23% of total foot length. Distal end aeuminate. Byssal groove and byssus absent in adults. Mantle (Figs 25-29): Mantle lobes symmetrical, thin, translucent white. Pallial muscles long, triangular, insert¬ ing from inner mantle fold region to -16% of total mantle lobe height, arranged sparsely at ventral margin of mantle lobe; separated from each other by -4 times pallial mus¬ cle basal width (Fig. 25: pm). Mantle border with three folds (Fig. 29); outer fold (of) thin, width -5% of shell thickness, 5 times higher than wide; middle fold (mf) similar to outer fold, -30% shorter; inner fold (if) short, -3 times taller than wide. Middle fold with 30 small and short papillae, bordering entire pedal gape portion (Fig. 26: pp); each papilla taller than wide, with rounded tip, separated from adjacent papillae by width equivalent to 4 times papillar width. Periostracum between outer and middle folds. Mantle lobes totally free exeept for siph¬ onal area. Anterior mantle fusion occurring at -42% of anterior adductor muscle height; posterior mantle fusion occurring at -70% of posterior adductor muscle height (Fig. 26). Siphonal area eorresponding to -30% of total mantle lobe length (Fig. 26). Incurrent and excurrent si¬ phons originating from inner mantle fold; siphonal area equivalent to -35% of total animal height and -7% of length (Figs 26, 27); siphons externally fused, eovered by small brown spots, internally separated by thick, smooth muscular wall (Figs 27, 28, 31); siphonal internal open¬ ings free, opening directly into pallial cavity; incurrent and excurrent siphons similar in size; -5 times longer than wide; incurrent siphonal external tip bordered by one row of short papillae, papillae length equivalent to -10% of total siphon length (Fig. 28: pp); excurrent siphonal tip with siphonal membrane (Fig. 28: sm). Pallial cavity (Figs 25, 26, 30, 32, 33): Occupying -50% of total internal shell volume (Fig. 25). Labial palps small, -2% of total internal shell volume, trian¬ gular (Figs 26, 32), external surfaee smooth; outer (op) and inner hemipalps (ip) of similar size, -60% narrow¬ er and 55% shorter than anterior adductor muscle inser¬ tion; outer hemipalp connected to mantle lobe by dorsal edge, at -30% of palp length; inner hemipalp conneeted to visceral mass by dorsal edge, at -20% of palp length; internal surface of each palp with -10 tall, rounded trans¬ verse folds covering -90% of inner palp surface, leaving thin smooth area at palp edges, eorresponding to -10% of total inner palp area. Folds decreasing in length toward mouth, forming shallow channels direeted to anterior and posterior portions of mouth (Fig. 32). Gill wide, -60% times wider than outer hemipalp, equivalent to -30% of total valve area (Fig. 25). Ctenidia eulamellibranch with two demibranchs (Fig. 30). Outer demibranch, fusiform, twice as long as wide; folded upon -30% of its own area; covering perieardial and renal areas; connected to mantle lobe by tissue for -15% of posterodorsal border length (Fig. 25); inner demibraneh triangular, -1.5 times longer than wide; folded upon 50% of its own area; covered by outer demibranch in area equivalent to -20% of its own area; food groove along ventral surface of inner demi¬ branch (Figs 30, 33: fg); demibranchs connected to eaeh other at posterior end by tissue (gf), fusion length equiv¬ alent to 25% of total gill length (Figs 25, 33); each dem¬ ibranch thin, fragile, without signs of chemosymbiotic bacteria (Figs 24, 30). Suprabranehial chamber -1/3 of infrabranchial ehamber volume (Fig. 26). Visceral mass (Fig. 26): Triangular, occupying half of total internal shell volume, laterally flattened, 2 times as wide as museular base; -40% of anterodorsal portion filled by brown digestive gland (dg); remaining area filled by cream-colored gonad (go). Stomach and style sac lo¬ cated vertically in central portion of visceral sac. Circulatory and excretory systems (Figs 26, 34): Pericardium located in posterodorsal region of visceral sac, between posterior region of umbonal cavity and dor¬ sal surface of kidney (Fig. 34), -2 times as long as wide; oecupying -25% of total viseeral mass volume. Paired auricles (au) anteroposteriorly elongated, connecting to main axis of gills along -1/3 of gill length; walls thin, translucent. Ventricle (ve) elongated, thick, located at eentral perieardial region, surrounding -45% of intestine crossing pericardial area, connected to auricles at median portion of lateral walls. Kidney light brown, triangular, located posteroventral to visceral mass, between ventral wall of perieardium and dorsal surfaee of posterior pedal retractor muscles, occupying -25% of total visceral mass volume. Nephropores (Fig.26: up) small, located at ante¬ rior third of kidney length, near genital pore (Fig. 26: gp). Digestive system (Figs 35-37): Palps and digestive gland as described above. Mouth small, located centrally between pairs of inner and outer labial palps. Esophagus (es) long, narrow, length -30% and height -10% of vis- eeral sae length and height (Fig. 35), eylindrieal, running separate from anterior adductor muscle between and par¬ allel to anterior portion of paired anterior pedal retractor muscles; internal surface covered by longitudinal folds, forming esophageal rim at stomaeh entrance (Fig. 37, er) in anteroventral region. Stomach (st) wide, occupy¬ ing -30% of visceral sac volume, conical, funnel-like, located anterior to umbo (Fig. 35); length -60% of total visceral sac length, -30% of its height; posterior portion -60% taller than anterior portion. Paired apertures to di¬ gestive caeca located ventrolaterally, turned toward ven¬ tral portion of visceral sac, located side by side at anterior portion of stomach. Dorsal hood (dh) long, thin, length -40% of total stomach length, anteriorly bluntly pointed. zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 517-534 523 Left pouch (Ip) located below anterior portion of dorsal hood, shallow, wide, occupying -20% of external area of left stomach wall, with ducts to digestive gland connect¬ ing at its central region (Fig. 36: dd). Stomach internal surface mostly smooth, with three well-developed sorting areas (Fig. 37); first sorting area starting at left side of es¬ ophageal rim, running along dorsal wall of anterior stom¬ ach chamber, penetrating dorsal hood, narrow, comprised of small transverse folds (sa 1). Second sorting area origi¬ nating ventral to first sorting area, at left side of esophage¬ al rim, running along left wall of anterior stomach cham¬ ber, entering left pouch and dorsal hood, both on their ventral surfaces, broad, formed by thickening of stomach wall (sa 2).Third sorting area starting inside dorsal wall of dorsal hood, running along dorsal and right walls of posterior stomach chamber, until diffusing on ventral portion of right wall (sa 3). Gastric shield (gs) located at central dorsal wall, occupying -40% of total gastric area, with two anterior projections, one dorsal at left border, penetrating dorsal hood, and one left ventral, penetrating left pouch. Two narrow, tall gastric ridges running along ventral stomach chamber, forming major and minor ty- phlosoles at style sac entrance (Fig. 37). Longer ridge originating posterior to left caecum, penetrating caecum and exiting its anterior end, running toward anterior por¬ tion of stomach, performing curve, penetrating anterior end of right caecum, exiting that caecum at its posterior end, penetrating style sac at its right side, forming ma¬ jor typhlosole (mt). Shorter fold originating at style sac entrance, at region of major typhlosole penetration into style sac, forming rim bordering style sac entrance and ul¬ timately minor typhlosole (nt). Style sac (ss) connecting stomach ventrally (Fig. 35), tapering ventrally, -3.3 times longer than wide, occupying -12% of visceral sac total volume; style sac height equivalent to 50% of visceral sac length, and -10% of its width. Intestine (in) thin, long, originating between typhlosoles, merging with style sac initially, narrowing after ventral end of style sac, passing ventrally below central stomach, penetrating pedal mus¬ culature at -5% of foot height, contacting dorsal surface of posterior pedal retractor muscles, curving toward right, following posterodorsal portion of visceral sac, parallel to style sac; intestine total length -7 times longer than style sac. Anus simple, sessile, on ventral surface of posterior adductor muscle (Fig. 31, 35: an). Reproductive system (Fig. 26): Gonads with branched aspect, opaque, cream-colored. Paired gonoducts con¬ nected to gonadal acini branches along posterodorsal por¬ tion of visceral sac. Genital pores simple (gp), located at posterior portion of visceral mass, at -20% of visceral mass height, near nephropore (up). Central nervous system (Figs 38-41): Paired cere¬ bral ganglia (Figs 38, 41: eg) surrounding dorsal surface of anterior esophagus, dorsal to external surface of outer labial palp, triangular, longer than wide (Fig. 38), length 50% of esophageal length. Each cerebral ganglion -50% width of transverse section of esophagus. Cerebral com¬ missure -50% longer than ganglia length; anterior ad¬ ductor muscle nerve (na) originating at anterior end of cerebral ganglion, contacting posterior surface of anterior adductor muscle, bifurcating into two main branches; in¬ ternal branch penetrating posterodorsal third of muscle, diffusing into muscle; outer branch bordering posterior surface of anterior muscle until contacting pallial re¬ gion and diffusing into muscle. Two additional pairs of nerves originating dorsally on cerebral ganglia, anterior to cerebrovisceral connective (cv) crossing visceral mass, contacting gonopore dorsally, bordering anterior portion of kidney and connecting dorsally with visceral ganglia, connecting cerebropedal connective (cp) running im¬ mersed in pedal muscles, connecting to anterior region of paired pedal ganglia (Figs 40, 41: pg). Paired visceral ganglia (Figs 39, 41: vg) fusiform, of similar length and height, length -60% of cerebral ganglia length, partially fused medially, with subcentral groove; located ventral to paired posterior pedal retractor muscle, parallel with posterior adductor muscle, at dorsal tip connecting to cerebrovisceral connective (cv, as described above) and renal nerve (rn), penetrating kidney area; laterally origi¬ nating ctenidial nerves (cn) running through central axis of posterior portion of gills; dorsally originating posterior adductor muscle nerve, penetrating mid-region of ante¬ rior surface of posterior adductor muscle; at ventral tip originating pallial nerve (pn), contacting anterior surface of ventral portion of posterior adductor muscle, running toward incurrent and excurrent siphonal muscles, reach¬ ing excurrent opening, originating single, short nerve (sn) that runs parallel to -25% of dorsal siphonal mus¬ cle length, continuing parallel to mantle border, diffusing into mantle lobe edge. Paired pedal ganglia totally fused (Figs 40, 41: pg), oval, longer than wide, -20% wider than visceral ganglia; located internal to posterior pedal retractor muscles, dorsal to foot insertion, at anterior tip connecting with cerebropedal connectives from cerebral ganglia; at posterior tip connecting two pairs of nerves, with dorsal pair running toward posterior region, inside posterior pedal retractor muscles; posteroventral pair curving ventrally, running into foot. Habitat. Infaunal, in muddy sand; usually positioned vertically in about 2 cm depth (Kat 1978), in mangrove areas and brackish waters. Measurements, (length by height by width, in mm): FMNH 328260 (specimen #1 of 4): 14.2 x 13.4 by 8.5; UF 122840 #1: 12.5 x 11.4 x 8; UF 264025 #1: 10.8 x 10.3 X 5.6; #2: 14.18 x 13.55 x 8.55. Distribution. USA: eastern coast from Delaware to the Florida Keys, and Gulf of Mexico coast from western Florida to Texas; Bahamas; Mexico: Yucatan, Quintana Roo; Guatemala; Bonaire (Lee 2009); Suriname. Type material. Syntypes: Cyrenoideafloridana'. United Stated Oe America • Florida, Fort Myers, Everglades; 2 specimens; USNM 87735. Marco Island; 3 specimens; zse.pensoft.net 524 Valentas-Romera, B.L. et al..: Anatomical redescription of Cyrenoida floridana Figures 25-34. Anatomy of Cyrenoida floridana (FMNH 328260). 25. Right view valve removed, some structures seen by trans¬ parency of mantle lobe; 26. Same, with mantle and gill removed; 27. Incurrent and excurrent siphons, posterolateral view, right mantle lobe partially removed, some adjacent structures shown; 28. Siphonal tips; right view, both partially sectioned longitudinally; 29. Mantle border, section in its ventromedial portion; 30. Gill, transverse section in its central portion; 31. Incurrent and excurrent siphons, interior view, with details of their base and siphonal muscles; 32. Labial palps, ventral view, outer hemipalps deflected dor- sally; 33. Posteroventral visceral region, ventral view, showing fusion of inner demibranchs in siphonal base; 34. Pericardial region, posterodorsal view, dorsal mantle wall partially removed. Scale bars; 2 mm (25-28, 31, 33-34); 1 mm (29, 30, 32). zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 517-534 525 Figures 35-41. Anatomy of Cyrenoida floridana (FMNH 328260). 35. Digestive tubes as in situ; right lateral view; 36. Stomach, left lateral view; 37. Stomach, right lateral view, right wall opened and deflected to show inner gastric surface; 38. Cerebral gan¬ glia, ventral view; 39. Visceral ganglia, ventral view; 40. Pedal ganglia; right flgure in lateral right view, left figure in dorsal view; 41. Nervous system topology, right lateral view. Scale bars; 2 mm (35-41, 40); 0.5 mm (38-40). zse.pensoft.net 526 Valentas-Romera, B.L. et al..: Anatomical redescription of Cyrenoida floridana USNM 60974. 3 specimens, USNM 60975; Boca Ciega Bay; 3 specimens; USNM 60973. St Johns River mouth; 10 specimens; USNM 46846 (Figs 1^). Georgia, Bruns¬ wick Island; 30 specimens; USNM 129197. Cyrenoidea guatemalensis'. Lectotype: Guatemala • Livingstone; ANSP107532 (Figs 5-10). Note: Pilsbry’s (1920) descrip¬ tion of C. guatemalensis can be read as having been based on a single specimen and Van Regteren Altena (1971: 41) interpreted the specimen of ANSP 107532 as a holotype. However, Pilsbry is known for imprecisely indicating the type material at hand (P. Callomon, G. Rosenberg pers. comm.) and the existence of more than one original type specimen cannot be excluded. We accept Van Regteren Altena’s (1971) action as a fixation of lectotype by infer¬ ence of holotype under ICZN (1999) Article 74.6. Examined material. United Stated Oe America • 10 valves; Delaware, Kent County, Bombay Hook; 01 Aug. 1954; Morrison and Rosso leg.; USNM 777892. • 10 valves; New Jersey, Delaware Bay, Cumberland County, Fortescue; 15 Jul. 1957; J.P.E. Morrison leg.; USNM 777894. • 6 valves; Delaware Bay, Cumberland County, Dividing Creek; J.P.E. Morrison leg.; 15 Jul. 1957; USNM 777895. • 20 valves; Maryland, Dorches¬ ter County; J.P.E. Morrison leg.; 11 Jul. 1954; USNM 777893. • 10 valves; Dorchester County, near Elliot, gul¬ let of black duck; F.M. Uhler leg.; USNM 592260. • 8 valves; Queen Anne’s County; 11 Jul. 1954; J.P.E. Mor¬ rison leg.; USNM 777890. • 12 valves; Arundel County, Deale, marshy, head of small inlet; 11 Jul. 1953; J.P.E. Morrison leg.; USNM 777887. • 6 valves; Arundel Coun¬ ty; 15 Jul. 1953; J.P.E. Morrison leg.; USNM 777889. • 4 valves; Arundel County, Deale; 13 Jun. 1954; J.P.E. Morrison leg.; USNM 777888. • 6 valves; North Caroli¬ na, Beaufort, under algal mats; R.W. Heard leg.; USNM 678947. • 1 specimen; South Carolina, Horry/George¬ town counties, Murrell’s Inlet, in black muddy sand un¬ der log near high tide line, S. edge of inlet along road; 03 Dec. 1955; J.P.E. Morrison leg.; USNM 1437782. • 6 valves; Georgia, McIntosh County, Fort King George Historic Site, Darien, exposed under drift logs and boards; 15 Dec. 1954; Cmdr. Miller leg.; USNM 707264. • 4 valves; Glynn County, Saint Simons Island; Oct. 1938; H.A. Rehder leg.; USNM 535386. • 1 valve; Mississip¬ pi, Jackson County, Halstead Bayou; Gulf Coastal Ma¬ rine Eaboratory leg.; UF 246126. • 15 valves; Florida, Wakulla County, St. Marks; 17 Jun. 1958; United States Fish and Wildlife Service leg.; USNM 612256. • 5 valves; Saint Johns County, Saint Augustine; F.E. Spinner leg.; ANSP 54330. • 6 valves; Saint Johns County, Halifax Riv¬ er; USNM 253659. • 12 valves; Volusia County, Daytona [Beach]; C.W Johnson leg.; USNM 336943. • 2 valves; Marion County, creek SE of Ocala; 15 May. 1928; T. Van Hyning leg.; ANSP 152656. • 8 valves; Citrus County, Homosassa; E. Roper leg.; USNM 131462. • 4 speci¬ mens; Hernando County; G. Prime leg.; ANSP 68457. • 25 specimens; Hernando County, Aripeka; G. Prime leg.; ANSP 73905. • 20 valves; Pasco/Hemando counties. Aripeka; E. Pine leg.; USNM 149932. • 10 valves; Her¬ nando County, Eittle Blind Creek; 04 Dec. 1927; T. Van Hyning leg.; ANSP 149568. • 1 valve; Pasco/Hillsbor¬ ough counties, Hillsborough River; E.J. Post leg.; USNM 591792. • 6 valves; Charlotte County, Punta Gorda; 1928; J.E. Madden leg.; USNM 592290. • 1 specimen; Glades County, Caloosahatchee River; C.W. Johnson leg.; ANSP 62888. • 5 valves; W Florida; C.W. Johnson leg.; ANSP 59610. • 30 valves; Collier County, Carnestown; 12 Apr. 1928; T. Van Hyning leg.; ANSP 152655. • 4 valves; Dade County, Miami; Olsen leg.; USNM 153404. • 10 valves; Dade County, Miami; 07 Apr. 01; Benedict leg.; USNM 330959. • 10 specimens; Dade County, Miami; S. N. Rhoads leg.; ANSP 77046. • 8 valves; Dade County, Miami; S. N. Rhoads leg.; ANSP 189416. • 16 valves; Eee County, Fort Myers; Hend. leg.; USNM 455820. • 16 valves; Eee County, Fort Myers; Henderson leg.; UNSM 425820. • 2 valves; Eee County, Fort Myers, Everglades; 1896; C.W. Johnson leg.; USNM 87735; syntype. • 20 valves; Monroe County, Big Pine Key; C. Margaret leg.; UF 122840. • 14 valves; Florida Keys, Monroe County, Big Pine Key; 27-28 Dec. 1956; C. Phillips, F. Philips leg.; FMNH 63059. • 31 valves; Florida Keys, Monroe County, pond on Big Pine Key; 1968; M. Teskey leg.; FMNH 293174. • 4 specimens +15 valves; Florida Keys, Monroe County, Blue Hole quarry on Big Pine Key, Flor¬ ida Keys; 024°42'21"N, 081°22'49"W; shoreline sedi¬ ment at base of vegetation, salinity measured at 3 ppt; sta. FK-727; 03 May. 2004; R. Bieler, PM. Mikkelsen leg.; FMNH 328260. • 67 valves; Florida Keys, Monroe County, Blue Hole quarry on Big Pine Key; 024°42'24"N, 081°22'48"W; sta. FK-794; 18 Nov. 2007; R. Bieler, P. Sierwald, E.A. Glover, J.D. Taylor leg.; same locality as in study by Taylor et al., 2009; FMNH 333534. • 5 valves; Florida Keys, Monroe County, off mangrove is¬ land SE of Cudjoe Key; 024°38T2"N, 081°18T2"W; 1 m; sta. FK-745; 15 May. 2005; R. Bieler, P. Mikkelsen leg.; FMNH 333533. • 11 valves; Florida Keys, Monroe County, quarry on Big Pine Key, in sediment on shore¬ line rock; 024°4F56"N, 081°23'03"W; sta. FK-728; 03 May. 2005; R. Bieler, P. Mikkelsen leg.; FMNH 333535. • 3 valves; Florida Keys, Monroe County, mosquito ditch on Big Pine Key; 024°42'30"N, 081°23'02"W; sta. FK- 939; 25 Apr. 2010; R. Bieler, P. Mikkelsen leg.; FMNH 333532. • 3 valves; Florida Keys, Monroe County, Span¬ ish Harbor Key; sta. JG-708-0; 08 Jun.2000; J. Gerber leg.; FMNH 308431. • 1 valve; Florida Keys, Monroe County, Ohio Key, land-locked pond adjacent to Ohio- Missouri Key bridge; 024°40'20'’N, 081°14'36"W; sta. FK-723; 29 Apr. 2004; R. Bieler, P. Mikkelsen leg.; FMNH 314324. • 4 valves; Monroe County, Boca Chica Key; H. Hemphill leg.; ANSP 7983. Bahamas • 14 valves; Grand Bahama Island; 26°3F00'’N, 78°46'30"W; J.N. Worsfold leg.; ANSP 374956. • 1 valve; Dover Sound, 26°35'05"N, 78°13'20"W, May.1983; J. N. Worsfold leg.; ANSP 374350. • 2 valves; 26°31'00"N, 78°46'30"W; J.N. Worsfold leg.; ANSP 374957. • 6 valves; Abaco; C.W. Johnson leg.; USNM 425821. • 2 valves; S side of Abaco; zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 517-534 527 O. Bryant leg.; USNM 180503. Mexico • 10 valves (1 figured specimen); Quintana Roo, Boca de Paila; Tulane University leg.; UF 264025. • 20 valves; Tulane Univer¬ sity; Emily & Harold Yokes leg.; UF 264026. GUATE¬ MALA • 2 valves; Livingstone; 1913; A. A. Hinkley leg.; syntype; ANSP 107532. Discussion Cyrenoida floridana and other Cyrenoidea anatomy Recently, the family Cyrenoididae was classified together with Cyrenidae and Glauconomidae based on analyses of nuclear 18S and 28S rRNA genes (Taylor et al. 2009). This statement was corroborated using an integrative approach, with anatomical, molecular, and ontogenetic views (Bieler et al. 2014). In both cases, only one species each from Cyrenidae and Glauconomidae was used, and a robust discussion comparing morphological characters among the families was still lacking. Here a detailed dis¬ cussion is provided, including morphological traits of the three families using available data from general traits of the family to details of species anatomy. Very little information regarding Cyrenoida floridana has been available. What morphological data have been published are fragmented and lack details (e.g. Taylor et al. 2009; Bieler et al. 2014). Kat (1978) provided a histo¬ logical and ecological study on the species, but only part of those results were formally published (Kat 1982). Habitat: All three families that comprise Cyrenoidea are infaunal in soft sediments, inhabiting fresh to brackish waters worldwide. Most species present geographic dis¬ tributions in Asia (e.g., Glauconomidae), but Cyrenidae includes species naturally distributed worldwide, except for Arctic and Antarctic regions, as well as introduced in¬ vasive species (Bieler and Mikkelsen 2019). Cyrenoidi¬ dae is known only from African and American continents in brackish waters. Kat (1978) commented that C. flori¬ dana presents hermaphroditism, brooding behavior, juve¬ nile dispersion by buoyance, thermal resistance, and site selection as adaptations to survive in intertidal areas of severe conditions subject to rapid environmental change. Additional details about C. floridana habitat in Canary Creek, Delaware were provided by Kat (1978). There, the species lives buried horizontally in the first two centim¬ eters of sediment. High sulfide concentration in the sedi¬ ment appears to inhibit colonization, although the species can tolerate a wide range of salinity, suspended particles, temperature, pH, and moisture content. Highest popula¬ tion densities were found in moist sediment protected by a primary layer of halophytic vegetation and a secondary cover of filamentous algae. Shell: As is common in fresh and brackish water bi¬ valves (Cummings and Graf 2015), C. floridana presents a persistent periostracum covering the entire shell sur¬ face. In contrast to African Cyrenoida species, Cyrenidae, and Glauconomidae that are covered in thick, brown to greenish periostracum (Huber 2015), C. floridana bears a thin periostracum. Periostracum in fresh and brackish water bivalves is a common feature, because such envi¬ ronments present much more corrosive properties (Cum¬ mings and Graf 2015) and this organic layer protects against shell corrosion. Kat (1978) noted that C. floridana periostracum is thicker near the shell border and almost invisible and iridescent near the umbo. Kat (1978) also described parallel folds in the thicker portions of per¬ iostracum in adult specimens of C. floridana. In living specimens, these folds create ridges that act to channel water when the bivalve is partially exposed, helping the animal to stay hydrated and thermally stable. Shape among Cyrenoidea varies from rounded (Cyre¬ noididae), trigonal (Cyrenidae), to anteroposteriorly elon¬ gated with a straightening of the posterior shell margin (Glauconomidae) (Joannis 1835; Owen 1959; Boss 1982; Huber 2015). Some representatives present a gap between the valves at the posterior end of the shell, as in Polymes- oda (Morton 1976) and Glauconome (Owen 1959). Shell size among Cyrenoida species can be variable. The African Cyrenoida can reach 30 mm (e.g. C. dupon- tia) whereas the American C. floridana reaches only 14 mm. Despite the size difference, ecological and mor¬ phological characteristics allow African and American species assignment to the same genus, i.e., the habitat in brackish waters and the unique hinge pattern. Compared with other Cyrenoidea, C. floridana can be considered the smallest representative of the superfamily; Cyrenidae attain lengths of 39 to more than 150 mm and Glaucono¬ midae between 20 and 79.5 mm (Huber 2015). Cyre¬ nidae is the only family within this group that presents concentric ribs; the remaining species are adorned only with growth lines (Boss 1982; Huber 2015). Commargin- al growth increments are here described for C. floridana. These thickened lines could be associated with seasonal growth anomalies or seasonal metabolic changes (Lewis and Cerrato 1997). Hinge: Hinge pattern in Cyrenoidea is somewhat var¬ iable. Glauconomidae and Cyrenidae share the presence of at least three cardinal teeth positioned perpendicular to the shell umbo, and one of these teeth can be bifid (Boss 1982; Huber 2015). Members of Cyrenidae share with C. floridana the presence of lateral teeth, but those in Cyrenidae can be either smooth or serrate. Cyrenoidi¬ dae, including African Cyrenoida and C. floridana, pre¬ sents a combination of lateral and cardinal teeth forming a unique pattern. Microtubules: The C. floridana shell presents mi¬ crotubules that partially penetrate the shell. This feature is shared with some Cyrenidae, e.g., Corbicula species (Araujo et al. 1994), but is absent in others, e.g., Polymes- oda (see Tan Tiu and Prezant 1989). According to Waller (1980), microtubules are more common in epifaunal than in infaunal bivalves and, based on studies in Arci- dae, could be associated with photoreception, anchorage sites for the mantle, improvement of the surface for res- zse.pensoft.net 528 Valentas-Romera, B.L. et al..: Anatomical redescription of Cyrenoida floridana piratory change, protection against boring organisms, or sites of ionic regulation. Rosso (1954) commented that microtubules could be involved in embryonic nourish¬ ment. Robertson and Coney (1979) stated that they could be used for monitoring water conditions, although this is unlikely given that they do not fully penetrate the shell. Tiu Tan and Prezant (1989) hypothesized that microtu¬ bules could act to lighten juvenile shells, aiding in plank¬ tonic dispersal, and to assist in anchoring the mantle to shell during locomotion or biomineralization. At any rate, microtubules occur in several bivalve families as a post- larval feature (Malchus 2010) and resemble the aesthetes found in some gastropods and polyplacophorans (Simone 2011), which mostly have receptor functions. Kat (1978) observed that the vast majority of tubules in C. floridana do not fully penetrate the shell and are filled with non-se- cretory, finger-like projections from the external layer of the mantle. The mantle includes blood sinuses that, when filled, could extend the projections inside the tubules, improving tissue attachment to the shell. Therefore Kat (1978) discarded any notion of secretory or sensory func¬ tion and believed that the microtubules serve to anchor the mantle to the inner shell surface. Pallial line: Another feature variable among Cyre¬ noidea is the pallial line. In Cyrenoida floridana it is weak, discontinuous, and without a sinus, whereas Glau- conomidae presents a narrow, deep sinus, and in Cyre- nidae, it varies between entire to including a shallow or deep sinus (Huber 2015). The difference between the presence of a sinus and pallial line intensity is due to size and form of the siphonal muscles, and the quantity of pallial muscles, respectively. Siphonal muscles in C. floridana are paired, short, and thin and insert on the pal¬ lial muscle insertion line; Glauconomidae and Cyrenidae have strong, long muscular bands that dislocate the pallial muscle insertion toward the interior of the valve (Owen 1959; Kurniushin and Glaubrecht 2002). Pallial muscles in C. floridana are spaced along the mantle border, creat¬ ing a discontinuous pallial line, whereas Glauconomidae and Cyrenidae present powerful pallial muscles strongly marked on the internal surface of the shell (Owen 1959; Glaubrecht et al. 2003). Main muscular system: All known species of Cyre- noidea present anterior and posterior adductor muscles, a pair of anterior pedal retractor muscles, and a pair of posterior retractor muscles (Owen 1859; Huber 2015); a pair of pedal protractor muscles are present in some Cyre¬ nidae (Simone et al. 2015). The overall symmetry of the adductor muscles in Cyrenoidea and Glauconomidae is slightly anisomyarian, with the anterior adductor mus¬ cle reniform and the posterior adductor muscle oval, but some Cyrenidae are isomyarian (Morton 1976; Simone et al. 2015; Huber 2015). Foot: Foot shape among Cyrenoidea varies between Cyrenidae, which bear a well-developed, strong, wide, axe-shaped foot, whereas Cyrenoididae and Glaucono¬ midae have a wedged-shaped foot (Owen 1959; Mansur and Meier-Brook 2000; Huber 2015). Dali (1898) de¬ scribed the Cyrenoida floridana foot as filiform, and this condition was used to classify the species in Lucinidae. The image of a living specimen included by Bieler et al. (2014, fig. 3 N) can be referenced to verify shape and color of foot in living condition. Kat (1978) commented that the foot of C. floridana can be extended by more than one shell length in the sediment, lacks a byssal gland, and in large individuals required up to 10 minutes to complete a burrowing cycle. Mautle: The number of mantle folds and presence of papillae are very diverse among Cyrenoidea. In Cyreni¬ dae, mantle papillae are relatively common (Boss 1982; Huber 2015), although they are not shared among all genera and species (Simone et al. 2015). Although three mantle folds are found in Cyrenidae (Morton 1976), the genera Corbicula and Cyanocyclas present four mantle folds, with the middle one doubled and forming papillae at the anteroventral portion of mantle border (Mansur and Meier-Brook 2000). Cyrenoida floridana and Glaucono¬ midae present a three mantle folds arrangement, but in C. floridana the middle fold bears papillae, whereas this does not occur in Glauconomidae (Owen 1959). The de¬ gree of mantle fusion of both mantle lobes varies greatly. In Cyrenidae (Huber 2015; Simone et al. 2015) and C. floridana, fusion occurs only at the siphonal area, where¬ as Glauconomidae presents a wider fusion along the in¬ ner fold and internal surface of middle fold (Owen 1959), forming an anterior pedal gape. Siphon: In Cyrenidae, the siphons can originate from the inner fold, as in Polymesoda (see Morton 1976) and Cyanocyclas (see Mansur and Meier-Brook 2000; as Ne- ocorbicula) or from fusion of the inner fold and the internal surface of middle fold, as in Corbicula (see Mansur and Meier-Brook 2000). This second pattern is also found in Glauconomidae (Owen 1959), classified as type B by Yonge (1957). In C. floridana, the siphon originates as 'mPolymes- oda, classified as a type A (Yonge 1957), but it presents a row of papillae described as type B by Yonge (1982). Siphon length is variable in Cyrenoidea from long or short, fused or separated (Huber 2015). Glauconomidae presents long siphons totally fused, whereas Cyrenidae can show separated short to long siphons (Owen 1959; Morton 1976; Mansur and Meier-Brook 2000; Huber 2015). The siphons of C. floridana were shown in liv¬ ing condition by Bieler et al. (2014, fig. 3N); showing siphons with different lengths, with the excurrent siphon two times longer than incurrent, although the incurrent one has a wider opening than the excurrent; this can be explained by the different degree of contraction of the two siphons. Kat (1978) illustrated C. floridana siphons with the excurrent one being longer, but half of its length is due to the extroverted siphonal membrane. At any rate, the cyrenoidid siphons are not as long as those of the Glauconomidae, which reach -50% of the animal length, but they are not as short as those of the cyrenid Polymes¬ oda erosa (Solander, 1786) (Morton 1976). All three families share the pattern of the incurrent opening wider than the excurrent (Owen 1959; Morton 1976; Boss 1982; zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 517-534 529 Mansur and Meier-Brook 2000; Simone et al. 2015). The different sizes of incurrent and excurrent siphons in C. floridana is a reversed pattern from that of the Glaucono- midae, in which the excurrent siphon is slightly shorter than the incurrent (Owen 1959); Cyrenidae presents si¬ phons of similar lengths (Morton 1976; Mansur and Mei¬ er-Brook 2000; Simone et al. 2015). The presence of papillar rows at the siphonal apertures is a common feature in all three families. They have, at least, one row of papillae at the external rim of the incur¬ rent siphon (Morton 1956; Owen 1959; Glaubrecht et al. 2003). Glauconomidae, C. floridana, and some Cyreni¬ dae present a siphonal membrane in the external excur¬ rent siphonal opening, but some Cyrenidae genera, e.g. Corbicula, also can present papillar rows at this opening (Araujo et al. 1993; Glaubrecht et al. 2003). Glauconomi¬ dae and some Cyrenidae, such as Corbicula and Polymes- oda erosa present papillae beyond the siphonal tip. In Glauconomidae, the external surfaces of both siphons present small papillae, especially on the ventral and dor¬ sal surfaces (Owen 1959). In Corbicula and P. erosa, the papillae occur at the siphonal base in parallel rows, which can surround the siphonal base, as in P. erosa (see Mor¬ ton 1976), or can be distributed along the entire mantle border up to the siphonal tip, as in Corbicula (Glaubrecht et al. 2003). According to Kat (1978), the siphonal pa¬ pillae in C. floridana serve a regulatory function for the incurrent siphon. When incurrent water presents a high concentration of suspended material or the siphon tips are below the sediment surface, the papillae are positioned over the siphonal entrance, whereas in conditions of low suspended material, the siphons are held at maximum dil¬ atation with the papillae away from the entrance. The siphonal base or tips in all three families show pigment as rings or spots in pale to dark brown, orange, or black (Owen 1959; Morton 1976; Araujo et al. 1993; Glaubrecht et al. 2003). Siphonal musculature is composed of siphonal retrac¬ tor muscles that can be present as only one muscle band or divided into two bands, one dorsal and one ventral. In Glauconomidae and most Cyrenidae, the siphonal retractor muscle appears as one wide muscular band; it is fan-shaped in Glauconomidae (Owen 1959; Korniushin and Glaubre¬ cht 2002), but in C. floridana and Polymesoda floridana (Conrad, 1846) each present two muscular bands (Simone et al. 2015) with the dorsal band bifid in C. floridana. Gills: Gills in Cyrenoidea are eulamellibranch and both demibranchs are present (Huber 2015). Both demi- branchs are wide in C. floridana, and the anterior portion of the inner demibranch inserts between the labial palps. In Cyrenoidea, gill size is usually small, without insertion of the inner demibranch between the palps (Owen 1959; Morton 1976; Simone et al. 2015). Stomach: The stomach of C. floridana presents the style sac and the midgut conjoined, the major typhlo- soles penetrating both left and right caeca, presence of a normal gastric shield that penetrates at dorsal hood and left pouch, ducts from digestive diverticula opening into the stomach via left pouch and both caeca, presence of sorting area at roof of anterior side of the dorsal hood and sometimes extending over the right wall of stomach, sorting area in the left pouch, and a sorting area at an¬ terior roof of stomach from esophagus to dorsal hood. Gastric shield teeth and a cuticular lining of the stomach, coded in the morphological/molecular analysis by Biel- er et al. (2014) and reaffirmed as having been observed by I. Temkin who conducted that part of the study (2019 pers. comm.), were not detected in the histological study by Kat (1978) nor in the present study. Kat (1978) also described two short caeca, one adjacent to the gastric shield and another near the intestinal groove, but those structures were not observed during this study. The main differences between C. floridana and other Cyrenoidea is that in Corbiculidae and Glauconomidae, digestive ducts open independently on the lateral wall of the stomach and Cyrenidae presents a sorting area on the anterior roof and posterior wall of the stomach. Intestinal coiling among Cyrenoidea shows a simple, loose pattern, with few loops. Midgut course can be sum¬ marized as starting as style sac, running ventrally in the visceral sac, followed by a portion running anteriorly, forming a dorsal loop directing the intestine posteriorly, then following parallel to the style sac until leaving vis¬ ceral sac, passing through the pericardium, crossing the dorsal surface of the kidney, and ending on the surface of posterior adductor muscle (Owen 1959; Morton 1976; Simone et al. 2015). This pattern is a little more complex in Corbicula, which presents several spiral coils at the anterior portion of the midgut (Araujo et al. 1993). The anus of C. floridana is located on the ventral sur¬ face of the posterior adductor muscle (Fig. 35: an). This is a unique position. The anus in the Cyrenidae can be found at different points on the posterior surface of the posterior adductor muscle (Owen 1959; Morton 1976; Araujo et al. 1993; Simone et al. 2015). In living specimens of C. floridana, Kat (1978) ob¬ served that the portion of the intestine posterodorsal to the anterior adductor muscle is folded when the intestine was empty, but straightened when the intestine was filled. Excretory system: The kidney did not present any unusual gross features during this study. In histological sections, Kat (1978) described U-shaped tubules differ¬ entiated into anterior and posterior portions based on cell type. Also Kat (1978) described soft, rounded concretions of unknown composition in the kidney lumen and, be¬ cause some were too large to be expelled, hypothesized that the kidney acts as a storage area for such concretions as a strategy to survive in intertidal environments. Reproductive cycle: Little is known about the repro¬ ductive cycle of C. floridana. Kat (1978, 1982) described the reproductive physiology of C. floridana, identifying the species as a simultaneous hermaphrodite, describing four gonadal stages and observing developing juveniles in the demibranchs, indicative of brooding behavior, as in many Cyrenidae (Huber 2015). Kat (1978) also noticed that a few specimens in the studied population were purely males. zse.pensoft.net 530 Valentas-Romera, B.L. et al..: Anatomical redescription of Cyrenolda floridana Based on the reproductive cycle of C. floridana, Kat (1978) discussed differing fertilization strategies, based on the number of embryos developing inside the gonad. Kat (1978) noticed that southern populations of C. floridana present a characteristic gonadal development and spawn twice a year, whereas northern populations present dis¬ creet alterations on gonadal development and successfully spawn only once. Due these differences Kat hypothesized that both populations are in process of differentiation. Nervous system: Kat (1978) performed a histological study on C. floridana specimens and provided addition¬ al details about the nervous system, especially regarding microscopic nerve branching and the presence of stato- cysts on the pedal ganglia. Parasitism: Calcareous nodules on the inside of C. floridana shells are sometimes visible in published pho¬ tographs of this species, e.g. those by Abbott (1974) and Abbott and Morris (1995). They have been variously re¬ ported, e.g. by Van Regteren Altena (1971: 41) who re¬ ferred to their presence on the type material of Cyrenoida guatemalensis (see Figs 7, 8) and stated that “the present Suriname specimens also possess blisters interiorly and I think that their presence is caused by some outward influ¬ ence in all.” Nodules were noted frequently during this study, although the small numbers of specimens examined do not present a reliable percent occurrence in the species or any living population. They were neither mentioned nor figured in the morphological study by Kat (1978), which involved an unquantifled “large number” of shells collected over nine months from the coast of Delaware. Each nodule seen during the present study presents as a small orifice on an igloo-shaped structure that could in¬ dicate parasitism by Trematoda (Huntley and De Baets 2015). Kat (1978) detected trematodes in sporocyst stages throughout the bivalve’s soft tissues in a small percentage of histological sections and notice that the infection nega¬ tively affected gonadal development and excretion. Also, histozoic and coelozoic parasitism by the haplosporidian protist Minchinia sp. has been reported in samples of C. floridana (Reece et al. 2004; Arzul and Carnegie 2015). Conclusions 1. Cyrenoida floridana is morphologically charac¬ terized by valves externally covered by thin light brown periostracum; muscle scars and pallial line only faintly visible on the internal shell surface, and a unique hinge pattern. 2. The species presents microtubules on the interior shell wall. 3. Anatomically the species presents slightly unequal adductor muscles; few pallial muscles that are well separated from each other; an inner demibranch in¬ serted between the labial palps; demibranchs fused to each other along their posterior ends; totally fused and pigmented siphons that originate from the inner mantle fold; two pairs of siphonal retractor muscles; loose intestinal coiling; and the anus located on the ventral surface of the posterior adductor muscle. 4. Cyrenoida floridana shares a similar habitat, its gill morphology, most of the stomach complexity, and si- phonal pigmentation with members of Cyrenidae and Glauconomidae. It differs from the latter two fami¬ lies in its hinge composition, small size, weak and discontinuous pallial line, few and separated pallial muscles at the mantle border, the presence of papillae along the entire ventral border of the mantle except the siphonal area, an excurrent siphon longer than the incurrent one, a bifld dorsal siphonal retractor muscle tip, demibranchs inserting between the labial palps, the absence of independently digestive ducts opening into the lateral side of stomach, and the anus located at the ventral surface of the posterior adductor muscle. 5. Calcareous nodules presenting single circular open¬ ings are common on the internal shell surface and could be associated with trematode parasitism. Acknowledgments We thank Dr Ellen Strong (USNM) for providing images of the syntypes of Cyrenoida floridana and providing ac¬ cess to the mollusk collection of USNM during the first author’s visit; her and Dr Ilya Temkin for information con¬ cerning the stomach structure of this species in their earli¬ er study; Dr Gary Rosenberg, Amanda Eawless, and Paul Callomon (ANSP) for providing images of C. guatemal¬ ensis and associated information concerning the material’s type status; Dr Gustav Paulay and Mandy Bemis (UF) for loaning samples of C. floridana, Eugene V. Coan for com- /V ments and literature suggestions; Enio Mattos and Phillip Eenktaitis (Eaboratorio de Microscopia Eletronica, Uni- versidade de Sao Paulo); and Eara Guimaraes (MZSP) for assistance with scanning electron microscopy, and Rob¬ ert Forsyth for providing comments and corrections. The specimens from the Blue Hole site on Big Pine Key, Flor¬ ida, were collected under Florida Keys National Wildlife Refuge Special Use Permit 41580 to RB. This project was partly suported by Fapesp (Funda^ao de Amparo a Pesqui- sa) proc. #2010/11401-8 and CNPq (Conselho Nacion- al de Desenvolvimento Cientiflco e Tecnologico) proc. #134425/2010-3, #159490/2012-0 and #203533/2014- 3. Support by the Bivalve Assembling the Tree of Fife project (http://www.bivatol.org) through the US National Science Foundation (NSF) Assembling the Tree of Fife (AToE) program (DEB-0732854) is also acknowledged. References Abbott RT (1974) American Seashells (2"‘‘ edn). Van Nostrand Reinhold Company, New York, 663 pp. [23 pis] Abbott RT, Morris PA (1995) A Field Guide to Shells of the Atlantic and Gulf Coasts and the West Indies. Houghton Mifflin Co., Boston, 350 pp. [74 pis] zse.pensoft.net Zoosyst. 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Z)e Grave ♦ Received 18 August 2019 ♦ Accepted 19 September 2019 ♦ Published 29 October 2019 Abstract A new mysid, Heteromysoides songkhlaensis, is reported from shallow water in the Songkhla Lagoon, southern Thailand. The new species is closely similar to H. nana in having a triangular rostrum and the eyestalk without a process at the anteromesial corner. However, the new species can be readily distinguished from H. nana by three setae on the inner margin and five setae on distal margin of the second segment of the mandibular palp; the carpopropodus of the fifth and sixth thoracic endopod of the new species is composed of four articles, and the telson of the new species is distally rounded, shorter than the sixth abdominal somite, and 1.3 times longer than its basal width; the spines on the telson form elongated spines from the distal to posterior margin. The new species also resembles H. dennisi, H simplex, and H. stenoura; however, it differs by (1) the presence of a rounded distal margin of telson, and (2) absence of a sharp process on the distolateral corner of the eyestalk. Key Words Brackish water, Heteromysoides songkhlaensis, Thale Sap, Thale Sap Songkhla Introduction In Southeast Asia, about 210 species in the order Mysida (Crustacea) have been reported (Sawamoto 2014), and 48 of these species have been documented in Thailand (Tatter- sail 1921; Murano 1988,1995; Fukuoka and Murano 2002; Hanamura et al. 2008, 2011; Moriya et al. 2015). Informa¬ tion about the mysid fauna in the Songkhla Lagoon system of southern Thailand has not been updated since Tattersall (1921), who reported Rhopalophthalmus egregius Hansen, 1910 and Nanomysis siamensis W. Tattersall, 1921. During a study of variability of the recruitment abun¬ dance of Metapenaeus spp. in the hyperbenthos of Thale Sap and Thale Sap Songkhla in the Songkhla Lagoon system in 2018, several mysid specimens were collect¬ ed. Among them, an undescribed species was discovered. The species showed morphological characteristics of the genus Heteromysoides Bacescu, 1968: (1) the cornea of the eye, which is restricted to the anterolateral part of the eyestalk, is reduced in size, and (2) the pleopods are not sexually dimorphic. The known species of the genus pri¬ marily occupy shallow marine waters, including marine caves, in the tropical and subtropical regions of the world. To date, 10 species have been reported in the genus Heteromysoides: H berberae Bacescu & Muller, 1985 from Somalia, east Africa; H cotti Caiman, 1932 from the Canary Islands; H. dennisi Bowman, 1985 from the Caribbean Sea; H. longiseta Bacescu, 1983 from Heron Island, eastern Australia; H. macrops Murano, 1988 from northern Australia; H. nana Murano, 1998 from Channel Island, Northern Territory, Australia; H. sahulensis Mura¬ no, 1998 from the Sahul Shelf, Australia; H. simplex Han- Copyright Rofiza Yolanda etal. This is an open access articie distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the originai author and source are credited. 536 Yolanda, R. et al.: A new species of Heteromysoides amura & Kase, 2001 from Okinawa, Japan; H. spongicola Bacescu, 1968 from Cuba and Grand Cayman; and H. ste- noura Hanamura & Kase, 2004 from the Caribbean Sea. In this paper, Heteromysoides songkhlaensis is newly described based on specimens collected in the Songkhla Lagoon system. This, the fourth mysid species known to occur in the Songkhla Lagoon and also the 49* in Thai¬ land, is newly added to the Southeast Asian mysid fauna. Material and methods The Songkhla Lagoon system, also known as Songkh¬ la Lake, is a large, shallow body of water located on the east coast of the Thai Peninsula in southern Thailand sit¬ uated between 7°08'N and 7°48'N and between 100°07'E and 100°35'E. The lagoon system covers approximately 1,082 km^ and comprises four distinct bodies of water: Thale Noi, Thale Euang, Thale Sap, and Thale Sap Songkh¬ la (Fig. 1). The lagoon exhibits three water regimes, fresh, brackish, and salt water, which are arranged from north to south, respectively. The salty, southern end of the lagoon is connected to the Gulf of Thailand. Within the lagoon sys¬ tem, salinity ranges from 0 to 34 psu in Thale Sap Songkh¬ la to almost zero in Thale Noi. In both Thale Sap and Thale Sap Songkhla, the substratum is primarily mud. Figure 1. The map of the Songkhla Lagoon system, southern Thai¬ land, shows seven sampling stations (black dot) where Heteromy¬ soides songkhlaensis sp. nov. was collected. St. 1. Ban Bang Kh- iat; St. 2. Ban Koh Nang Kum; St. 3. Ban Laem Chak; St. 4. Ban Tai; St. 5. Ban Pa Khad; St. 6. Ban Bo Pab; St. 7. Ban Hua Khao. Specimens were collected by using a modified Riley’s hand-pushed net. Its mouth frame was 30 x 50 cm (height X width), the mesh sizes were 2 mm and 0.5 mm, and its side length 2.5 m. The net was pushed forward for 30 m along the shallow zone of Songkhla Lagoon at seven sta¬ tions (Fig. 1). The specimens collected were fixed in 4% formalin in the field and brought back to the laboratory. In the laboratory, mysid specimens were sorted and then transferred to 70% ethanol for further study. Using a micrometer installed in the eyepiece of the mi¬ croscope, body length (BE) was measured from the tip of the rostmm to the distal end of the telson, excluding apical denticles. Illustrations were made with the aid of a camera lucida. The marginal setae of some appendages, especial¬ ly the antennal scale, thoracopodal exopods, and uropod, were omitted from the illustrations. Terminology was based mainly on Tattersall and Tattersall (1951) and Wittmann et al. (2014). Specimens examined in this study are deposited in the collection of Prince of Songkla University Zoological Collection (PSUZC), at the Princess Maha Chakri Sirind- hom Natural History Museum, Prince of Songkla Universi¬ ty in Hat Yai, Songkhla, Thailand; Phuket Marine Biological Centre (PMBC) in Phuket, Thailand; Zoological Reference Collection (ZRC) of the Fee Kong Chian Natural History Museum, National University of Singapore and National Museum of Nature and Science, Tokyo (NSMT), Japan. Systematic account Heteromysoides songkhlaensis sp. nov. http://zoobank.org/DE6E674F-CEB3-4CAD-829C-8221F21ElF77 Figs 2-4 Type material. Holotype. Adult male (BE 3.2 mm) (NMST-Cr 26744), Thale Sap, 7°20'58.68"N, 100°25'3L56"E, Ban Bang Khiat, Tambon Bang Khiat, Singha-Nakhon District, Songkhla Province, Thailand, 19 January 2019, at 1.3 m of depth with salinity of 0.39 psu, over a muddy substrate, coll. V. Eheknim, N. Tubtim- tong and R. Yolanda. Allotype. Adult female with empty marsupium (BE 3.7 mm) (PSUZC 20190119-02.01), Thale Sap Songkhla, 7°15T8.77"N, 100°28TL86"E, Ban Pa Khad, Singha-Nak¬ hon District, Songkhla Province, Thailand, 19 January 2019, at 1.3 m of depth with salinity of 0.47 psu, over a muddy substrate, coll. Y Eheknim, N. Tubtimtong and R. Yolanda. Paratypes. 1 adult male (BE 3.6 mm, dissected) (NS- MT-Cr 26745), Thale Sap, 7°19'34.50"N, 100°24'3L45''E, Ban Koh Nang Kum, Koh Nang Kum, Pak Payoon Dis¬ trict, Phattalung Province, Thailand, 18 May 2018, at 1.3 m of depth with salinity of 3.68 psu, over muddy substrate, coll. V. Eheknim, N. Tubtimtong and R. Yolanda; 1 adult male (BE 3.4 mm, dissected) (PSUZC 20180618-01.01) Thale Sap Songkhla, 7°13T4.67''N, 100°31'24.12'’E, Ban Bo Pab, Sathing Mor, Singha-Nakhon District, Songkhla Province, Thailand, 18 June 2018, at 1 m of depth with salinity of 1.67 psu, over muddy substrate, coll. V. Ehek- zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 535-542 537 Figure 2. Heteromysoides songkhlaemis sp. nov. Holotype, male (BL 3.2 mm, A, C, E, F, G, I) (NSMT-Cr 26744), allotype, fe¬ male (BL 3.7 mm, B, D, H, J) (PSUZC 20190119-02.01), paratype, male (BL 3.6 mm, 1^0) and female (BL 3.6 mm, K) (ZRC 2019.1095). A, B. Lateral view of whole body; C, D. Dorsal view of anterior body; E. Dorsal view of right eye; F, H. Dorsal view of right antennule; G. Ventral view of right antennule; I, J. Ventral view of antenna; K. Ventral view of labrum; L. Ventral view of mandibles with palps; M. External view of mandibles enlarged; N. Right maxillule; O. Right maxilla. Scale bars: 0.5 mm (A, B); 0.3 mm (C, D, F-J); 0.2 mm (L); 0.1 mm (E, K, M-O). zse.pensoft.net 538 Yolanda, R. et al.: A new species of Heteromysoides nim, N. Tubtimtong and R. Yolanda; 1 adult male (BL 3.5 mm, dissected) (PMBC 11806), Thale Sap, 7°20'58.68"N, 100°25'31.56"E, Ban Bang Khiat, Tambon Bang Khiat, Singha-Nakhon District, Songkhla Province, Thailand, 18 July 2018, at 0.6 m of depth with salinity of 2.72 psu, over a muddy substrate, coll. V. Lheknim, N. Tubtimtong and R. Yolanda; 1 adult female (BL 3.7 mm, dissected) (PMBC 11807), Thale Sap, 7°16'30.89"N, 100°25T7.21"E, Ban Laem Chak, Pak Ror, Singha-Nakhon District, Songkh¬ la Province, Thailand, 18 April 2018, at 1.5 m of depth with salinity of 15.1 psu, over muddy substrate, coll. V. Lheknim, N. Tubtimtong and R. Yolanda; 1 adult female (BL 3.1 mm, dissected) (NSMT-Cr 26746), Thale Sap Songkhla, 7°ir45.16"N, 100°33'33.76"E, BanHuaKhao, Hua Khao, Singha-Nakhon District, Songkhla Province, Thailand, 18 April 2018, at 1.4 m of depth with salinity of 24.8 psu, over a muddy substrate, coll. V. Lheknim, N. Tubtimtong and R. Yolanda; 2 adult females (BL 3.2 mm [dissected], 3.6 mm [not dissected]) (NSMT-Cr 26747), Thale Sap, 7°20'58.68"N, 100°25'31.56"E, Ban Bang Kh¬ iat, Tambon Bang Khiat, Singha-Nakhon District, Song¬ khla Province, Thailand, 18 May 2018, at 1.1 m of depth with salinity of 7.43 psu, over muddy substrate, coll. V. Lheknim, N. Tubtimtong and R. Yolanda; 2 adult males (BL 3.6 mm, [dissected], 4.0 mm [not dissected]); 1 adult female (BL 3.6 mm, dissected) (ZRC 2019.1095), Thale Sap Songkhla, 7°14'32.41'’N, 100°25'50.57"E, Ban Tai, Pak Ror, Singha-Nakhon District, Songkhla Province, Thailand, 18 November 2018, at 1.2 m of depth with salin¬ ity of 5.25 psu, over muddy substrate, coll. V. Lheknim, N. Tubtimtong and R. Yolanda. Description. Head and cephalic appendages. Carapace with anterior margin obtusely produced into wide, triangu¬ lar rostrum (Fig. 2C, D); cervical groove distinct at anterior two-fifths, posterior margin excavated, leaving last thoracic somite uncovered in dorsal view, but sufficiently covered laterally (Fig. 2A, B): antero-ventral comer rounded (Fig. 2A, B). Eye slightly depressed, subglobular in dorsal as¬ pect; cornea comprise of small tube-like ommatidia situat¬ ed in antero-lateral part without ocular process (Fig. 2C-E). Antennule with first segment of peduncle longer than wide, with dorsal projection bearing 3 setae (Fig. 2F, H); distolat- eral comer of first segment greatly produced anteriorly, dis¬ tal part with 5 setae; second segment shortest, with dorsal projection bearing 2 setae and distal part with 2 setae; third segment subequal to first segment, with dorsal projection bearing 4 setae dorsally, middle part with 2 setae at the me¬ sial margin for female while no seta on male, respectively and 2 setae at distomesial comer for male while 3 setae for female; the lobe completed with several long setae on male (Fig. 2G) while no long setae on female. Antennal peduncle more robust in male than female and reaching 0.8 length of antennular peduncle and the sympod rounded (Fig. 21, J); antennal scale elongated, elliptical with apical suture, setose all round, reaching middle part of third segment of antennular peduncle, nearly 3 times as long as wide, not reaching distal end of antennal peduncle (Fig. 21, J). La- bmm triangular, without process or spine in anterior part but with expanded disto-lateral parts (Fig. 2K). Mandibular palp 3-segmented, second segment longest and widened at mid-length, with barbed setae on both margins, 3 on middle part of inner margin, 1 seta on proximal part of outer mar¬ gin and 5 setae on the distal margin of the second segment (Fig. 2L); incisor process well developed and comprised of a series of teeth forming serrated sharp ridge; lacinia mo- bilis showing different shape in right and left mandibles, and spine row and molar process clearly visible (Fig. 2M). Maxillule well developed, basal lobe with spines densely, wider than precoxal lobe (Fig. 2N). Maxilla with 4 distal setae and smooth setae on exopod (Fig. 20); distal segment of endopod longer than proximal one; basal and coxal en- dites well developed, with dense setae. Thoracopods. Flagelliform part of first and eighth thoracopodal exopods composed of 8 segments (Figs 3A, 4C), while second to seventh thoracopodal exopods with 9 segments (Figs 3C, E, G, I, 4A, B). First thoracopo¬ dal endopod short and basis well developed, larger than endite; medial margins of carpus, propodus and dactylus heavily setose (Fig. 3A, B). Second thoracopodal endo¬ pod stout (Fig. 3C); basis with 2 setae; preischium with 1 seta, shorter than basis; ischium longer than preischium with 6 setae; merus longest, with 2 setae; carpopropodus 0.75 times as long as merus, with several barbed setae (Fig. 3D); dactylus 0.5 times as long as carpopropodus, 1.6 times as long as width, with several barbed setae (Fig. 3D). Third and fourth thoracopodal endopods sim¬ ilar in form (Fig. 3E, G) and more slender than second (Fig. 3C); basis with 2 setae; ischium slightly longer than merus, with 4-6 setae on inner margin; merus slightly shorter than carpopropodus; carpopropodus constituting 3 sub-segments with barbed setae, basal segment long¬ est and second shortest; dactylus short, apex with long barbed seta and several setae (Fig. 3E-H). Fifth thoracop¬ odal endopod longest (Fig. 31), with similar morpholog¬ ical characters to sixth, seventh, and eighth (Fig. 4A-C). Sixth thoracopodal endopod (Fig. 4A) subequal in length to seventh and eighth (Fig. 4B, C); basis with 1 seta, is¬ chium longer than merus, with several setae; merus with 6 setae; carpopropodus constituting 4 sub-segments, with several setae, basal segment longest, third segment short¬ est; dactylus short, with several setae. Penis (Fig. 4C) long, 0.8 times as long as ischium of eighth thocacopodal endopod, apex rounded, with several smooth setae. Pleon andpleopods. Abdominal somites smooth, with¬ out hairs, spines or folds, ventral sternites without process, anterior 5 somites subequal in length, sixth somite 1.3 times as long as preceding somite (Fig. 2A, B). Five pleopods reduced to unsegmented lobes, not modified; first pleopod shortest, second to fourth ones subequal in length and fifth pleopod longest, 1.6 times as long as fourth (Fig. 4D-H). Uropod and telson. Uropodal endopod slightly shorter than exopod, without spine on ventral side of statocyst region (Fig. 41). Telson (Fig. 4K-N) subtriangular, 0.8 times as long as sixth abdominal somite, 1.3 times as long as basal width, excluding apical denticles, with 11-18 ar- zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 535-542 539 Figure 3. Hetewmysoides songkhlaensis sp. nov. Paratype, male (BL 3.6 mm, A-I) (ZRC 2019.1095). A. Right first thoracopod; B. Right first thoracopodal endopod enlarged; C. Right second thoracopod; D. Carpopropodus, dactylus and setae enlarged; E. Right third thoracopod; F. Distal part of carpopropodus, dactylus and setae enlarged; G. Right fourth thoracopod; H. Distal part of carpo¬ propodus, dactylus and setae enlarged; 1. Left fifth thoracopod. Scale bar; 0.2 mm (A, C, E, G, I); 0.1 mm (B, D, F, H). zse.pensoft.net 540 Yolanda, R. et al.: A new species of Heteromysoides Figure 4. Heteromysoides songkhlaensis sp. nov. Holotype, male (BL 3.2 mm, D-H, K), (NSMT-Cr 26744), allotype, female (BL 3.7 mm, L) (PSUZC 20190119-02.01), paratype, male (BL 3.6 mm, A-C, I, J, M) and female (BL 3.6 mm, N) (ZRC 2019.1095). A. Right sixth thoracopod; B. Right seventh thoracopod; C. Right eighth thoracopod; D-H. Right first to fifth pleiopods; I. Dorsal view of telson and uropods; J. Ventral view of right uropod; K-N. Dorsal view of telson. Scale bars; 0.2 (A-C, I-N); 0.1 mm (D-H). zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 535-542 541 Table 1. Body length and the number of spines on the telson of Heteromysoides songkhlaensis sp. nov. southern Thailand. from Songkhla lagoon, No. Type of specimen Sex Body length (mm) Number of spines 1 Holotype Male 3.2 15 2 Paratype Male 3.4 12 3 Paratype Male 3.5 13 4 Paratype Male 3.6 14 5 Paratype Male 3.6 11 6 Paratype Male 4.0 18 Range 3.2-4.0 11-18 7 Allotype Female 3.7 15 8 Paratype Female 3.1 12 9 Paratype Female 3.2 17 10 Paratype Female 3.6 19 11 Paratype Female 3.6 17 12 Paratype Female 3.7 16 Range 3.1-3.7 12-19 ticulated denticles in males and 12-19 in females on dis¬ tal quarter margin (Table 1), increasing in length distally. Etymology. This species is named after the locality, Songkhla Lagoon, where of the specimens were found. Distribution. This species was captured in brackish wa¬ ters above a muddy substratum at Thale Sap and Thale Sap Songkhla, Songkhla Lagoon, southern Thailand. Discussion Heteromysoides songkhlaensis is closely similar to H. nana in (1) having a triangular rostrum of the carapace, and (2) lacking a process at the anteromesial of the eye- stalk. However, the new species can be distinguished from H. nana by several features: (1) the mandibular palp displays three setae on the inner margin and five setae at the distal margin of the second segment as op¬ posed to two setae on the inner margin and three setae at the distal margin of the second segment in H. nana, (2) the carpopropodi of the fifth and sixth thoracic en- dopods of the new species is composed of four articles compared to five articles in H. nana, (3) the telson of the new species is distally rounded, shorter than the sixth abdominal somite, and 1.3 times longer than its basal width, while in H. nana the telson is slightly concave, longer than the sixth abdominal somite, and 1.6 times longer than its basal width; and (4) the spines on the telson are noticeably different in their arrangement from those of H. nana. The new species also exhibits similarities to several other Heteromysoides species, e.g. H. dennisi, H. sim¬ plex, and H. stenoura, in having (1) the telson without a cleft or sinus at its distal end, and (2) the uropodal endopod without a spine on the inner mesial margin. However, the new species differs from them in having (1) the eyestalk without a sharp process at the disto-lat- eral part, and (2) the telson rounded, not truncated, at the distal margin. Several species from the family Mysidae in South¬ east Asia have been found from brackish to full-strength seawater, e.g. Mesopodopsis orientalis (e.g., Hanamura et al. 2008), Acanthomysis thailandica (e.g, Ramarn et al. 2012), and Rhopalophthalmus spp. (e.g., Hanamura et al. 2011). So far, all described species of Heteromy¬ soides have been found in seawater, whereas the new species was found in brackish waters. Further study would be needed to elucidate the exact relationship of the current geographical pattern and the origin of Heter¬ omysoides species. Acknowledgements The present study is part of a research project on “Distri¬ bution patterns and variability in abundance of post larvae and juvenile of Metapenaeus spp. for fishery status and management guidelines in Thale Sap Songkhla, southern Thailand” which is supported by a grant (SCI 6003643) from Prince of Songkhla University. The first author ex¬ presses his deep gratitude to the Graduate School, Prince of Songkla University for the scholarship award on Thai¬ land’s Education Hub for ASEAN Countries (TEH-AC) (contract no. TEH-AC 042/2017) and for financial sup¬ port of this research. We also thank Mr Naratip Tubtim- tong for the fieldwork and Mr Sompong Pachonchit for driving us during this study. Finally, we thank an anon¬ ymous reviewer and Dr Yukio Hanamura for their useful comments which improved the manuscript. 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The Ray Society, Eondon, 460 pp. Wittmann KJ, Ariani AP, Eagardere JP (2014) Orders Eophogastrida Boas, 1883, Stygiomysida Tchindonova, 1981, and Mysida Boas, 1883 (also known collectively as Mysidacea). In; Klein JCVV, Char- mantier-Daures M, Schram FR (Eds) Treatise on zoology - anatomy, taxonomy, biology. The Crustacea. Revised and updated, as well as extended from the Traite de Zoologie, 4B. Brill Publisher, Eeiden, 189-396. https;//doi.org/10.1163/9789004264939_006 zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 543-556 | DOI 10.3897/zse.95.38727 4>yEnsPFr. BERLIN The evolutionary terrestrialization of planarian flatworms (Platyhelminthes, Trieladida, Geoplanidae): a review and research programme Ronald Sluys^ 1 Naturalis Biodiversity Center, P.O. Box 9517, 2300 RA Leiden, The Netherlands http://zoobank. org/49967409-C3F7-4005-9F7B- 7EAECBB93AA8 Corresponding author: RonaldSluys (Ronald.Sluys@naturalis.nl) Academic editor. A. Schmidt-Rhaesa ♦ Received 1 August 2019 ♦ Accepted 20 September 2019 ♦ Published 29 October 2019 Abstract The terrestrialization of animal life from aquatic ancestors is a key transition during the history of life. Planarian flatworms form an ideal group of model organisms to study this colonization of the land because they have freshwater, marine, and terrestrial represen¬ tatives. The widespread occurrence of terrestrial flatworms is a testament to their remarkable success occupying a new niche on land. This lineage of terrestrial worms provides a unique glimpse of an evolutionary pathway by which a group of early divergent aquatic, invertebrate metazoans has moved onto land. Land flatworms are among the first groups of animals to have evolved terrestrial adap¬ tations and to have extensively radiated. Study of this terrestrialization process and the anatomical key innovations facilitating their colonization of the land will contribute greatly to our understanding of this important step in Metazoan history. The context and scien¬ tific background are reviewed regarding the evolutionary terrestrialization of land flatworms. Furthermore, a framework of a research programme is sketched, which has as its main objective to test hypotheses on the evolution of land planarians, specifically whether particular anatomical and physiological key innovations have contributed to their evolutionary successful terrestrial colonization and radiation. In this context special attention is paid to the respiration in aquatic and terrestrial planarians. The research programme depends on a comprehensive phylogenetic analysis of all major taxa of the land flatworms on the basis of both molecular and anatom¬ ical data. The data sets should be analyzed phylogenetically with a suite of phylogenetic inference methods. Building on such robust reconstructions, it will be possible to study associations between key innovations and the evolutionary terrestrialization process. Key Words adaptations, evolution, key innovations, land flatworms, model organisms, respiration, terrestrialization Introduction Charles Darwin was fascinated by planarian flatworms, and he was particularly struck by the fact that there is a group of planarians that actually live on land. As he wrote in a letter from 23 July 1832 to his mentor Henslow: “Amongst the lower animals, nothing has so much interested me as finding 2 species of elegantly coloured true Planariae inhabiting the dry forest.” And in a letter from 15 August 1832: “I have today to my astonishment found 2 Planariae living under dry stones....”. Darwin thought for a long time that he was the first person to have discovered terrestrial flatworms. It was only in 1846 that it came to his attention that already in 1774 the Danish naturalist O. F. Muller had described the land flatworm Microplana terrestris (Muller, 1774) (Porter and Graham 2016). Darwin (1983: 25) appropriately and succinctly described land flatworms as follows: “In general form they resemble little slugs, but are very much narrower in proportion, and several of the species are beautifully coloured with longitudinal stripes.” He took some of Copyright Ronald Sluys. This is an open access articie distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the originai author and source are credited. 544 Sluys, R.: The evolutionary terrestrialization of planarian flatworms the land flatworms that he had collected in Tasmania on board of the Beagle and managed to keep them alive for two months. He experimented on the animals by cutting some animals into half, and he discovered that after 25 days each piece had regenerated to an almost complete animal (Darwin 1983; Sluys 2016). Darwin was so much intrigued by his flatworm findings that in a letter of 22 May 1833, written when he was in Tierra del Fuego, he asked his sister Catherine to send him DalyelTs (1814) paper on several species of British freshwater planarians (Thomson 2009). In this paper Daly ell (1814) reported many observations on the biology of these worms, including a detailed study of their fission and regeneration (Ball and Reyoldson 1981). Since the days of Darwin and Dalyell our knowledge on the regenerative capacity of land planarians, and that of triclad flatworms in general, has greatly increased (cf Reddien 2018; Rink 2018), as well as our knowledge on the morphology, taxonomy, phylogeny, and distribution of land flatworms (cf Sluys and Riutort 2018 and references therein). The fact that delicate organisms such as free-living flatworms, most of which occur in freshwater or marine environments, are able to survive on land has received little attention. Thus far, no detailed studies have been undertaken that attempted to analyze how and when during their evolutionary history planarians conquered the terrestrial environment, which of their anatomical and physiological features enabled their colonization of the land, and which adaptations currently facilitate their occupation of the terrestrial niche. Therefore, in the following I do not so much present the results of such studies, but provide a review of this subject and sketch the context, scientific background, and framework of a research programme in which land flat- worms form the model group through which we may not only learn about their own terrestrialization but may be enlightened also on the early evolutionary terrestrializa¬ tion of animal life in general. In this context, special at¬ tention is paid to the respiration in aquatic and terrestrial planarians. In addition, the results obtained during this putative research programme will also provide data for some collateral topics, such as biodiversity assessment and historical biogeography. Early evolutionary terrestrialization of animal life, as exemplified by land flatworms The terrestrialization of animal life from marine or freshwater ancestors is a key event in the history of life on earth, particularly because in the course of evolution “.. .transitions among physically different habitats... are rare” (Vermeij and Dudley 2000: 546; see also Vermeij 2010). Moving from an aquatic to a terrestrial niche is challenging due to the dramatically different demands each environment places on the physiology and structure of an organism, thus requiring numerous innovations. Key innovations afford enhanced performance, promot¬ ing ecological opportunity. These innovations enable subsequent species diversification and radiation. For ex¬ ample, marine gastropod snails in several cases evolved terrestriality independently from amphibious ancestors by side-stepping a major constraint on land snail evo¬ lution, viz., the need to produce mucus for locomotion (Rosenberg 1996). Interestingly, mucus plays an impor¬ tant role in land flatworms: forming a slime trail for lo¬ comotion, slime threads by which they can cross spaces or lower themselves from heights, and a protective coat against drying. Apparently, the worms have followed a different adaptive pathway than the molluscs and were able to colonize the land without conserving mucus. The land flatworms or planarians (Fig. 1) likely represent one of the first groups of animals that during evolution have colonized the land and have extensively radiated (see below: Impact and innovative aspects). Study of the evolutionary terrestrialization of the land flatworms and the key anatomical innovations facilitating this process will contribute greatly to our understanding of the early steps onto land of the Metazoa. Planarian flatworms (Platyhelminthes Claus, 1887, Tricladida Lang, 1884) form an ideal group of model organisms to study this process because they have freshwater, marine, and terrestrial representatives. Preadaptations for life on land are the conditions that (1) flatworms have internal fertilization, (2) the cocoons enveloping the eggs have a relatively hard shell, (3) the young hatch directly as young worms and not as larvae (Little 1983), and that (4) the worms produce mucus. Traditionally, three major groups of triclads were recognized: Paludicola Hallez, 1892 (freshwater planarians), Maricola Hallez, 1892 (marine triclads), Terricola Hallez, 1892 (land planarians). A fourth clade, the Cavemicola Sluys, 1990, was proposed by Sluys (1990). More recently, molecular phylogenetic studies have shown that one of the three paludicolan families is more closely related to the land planarians than to the other freshwater planarians (Baguna and Riutort 2004; Alvarez- Presas et al. 2008; see also Sluys and Riutort 2018). Figure 1. Photograph of the South American land planarian Polycladus gayi (from Grau and Carbayo 2010). zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 543-556 545 Model organism: land planarians Terrestrial planarians (Platyhelminthes, Tricladida, Geo- planidae Stimpson, 1857) are a relatively species-rich group (approx. 910 nominal species) with a worldwide, mainly pan-tropical, distribution (Fig. 2). The animals live typically in tropical jungles and wooded areas. Terrestri¬ al planarians colonized the land hundreds of millions of years ago, but they still lack any special mechanism for water conservation (Kawaguti 1932). Although they re¬ quire a humid environment, they cannot endure long sub¬ mersion in water, in contrast to their marine and freshwa¬ ter relatives (Froehlich 1955). The animals cannot endure heat and direct sunlight, otherwise they desiccate quickly (Kawaguti 1932). Because of these limitations they tend to remain hidden during the day in humid, but not wet, refuges, only emerging at night when the relative humid¬ ity of the air is high. Land planarians are fully terrestrial cryptozoic organisms because they are not tied to aquatic systems for reproduction. The worms are part of the soil ecosystem, living within the habitat of their prey (Ogren 1955; Ball and Sluys 1990; Ogren and Sheldon 1991). Terrestrial planarians are successful top-predators of other invertebrates such as snails, slugs, earthworms, isopods, insect larvae, and springtails; they themselves are rarely predated upon by other organisms, although some of their predators may be beetles, snails, and other land planarians (Boll and Leal-Zanchet 2018 and references therein). Land planarians search for, attack, and capture prey much larg¬ er than themselves, employing various techniques such as physical force, adhesive mucus, pharyngeal action, and pouring very effective digestive secretion over the surface of the live prey, or into it, by the protrusible pharynx. The widespread occurrence of these terrestrial flat- worms (Fig. 2) is a testament to their remarkable success occupying a new niche on land for hundreds of millions of years. This lineage of flatworms thus provides a unique glimpse of an evolutionary pathway by which a group of early divergent aquatic, invertebrate metazoans has moved onto land. Objectives The main objective of the research programme described here is to test hypotheses on the evolution of land flat- worms, speciflcally whether particular anatomical and physiological key innovations have contributed to their successful terrestrial colonization and subsequent radia¬ tion (see below: Hypotheses testing). This first requires a comprehensive phylogenetic analysis of all major taxa of the land flatworms (e.g., for the current 55 genera; cf Sluys et al. 2009). The project needs to construct this phytogeny on the basis of both molecular and anatomical data. This reconstruction forms the necessary basis and framework for subsequent comparative studies of key innovations, adaptive radiation, and historical biogeography of land flatworms, and on the early terrestrialization of animal life. Key innovations, adaptive radiation, and terrestrialization Understanding ofthe evolutionary dynamics of the following presumed key innovations, for example, may contribute to our insight in the evolution of land flatworms: (1) colonization of the land (on which occasions did the transition from water to land occur and were there reversals?); (2) contribution of the various kinds of anatomically complex creeping soles and (3) of mesenchymal body musculature (absent in freshwater and marine forms) to the effective terrestrialization; (4) contribution of cephalic specializations for the capture of prey to the adaptive radiation process; (5) the relation between the ecology and anatomy of the various taxa and their various types of pharynges (frequently totally different from freshwater and marine forms) for capturing and digesting prey; (6) the extent to which the various kinds of multi-cellular eyes of land flatworms (completely different from marine and freshwater forms) facilitated terrestrialization and adaptive radiation; (7) adaptation of particular sense organs, such as olfactory chemoreceptors, to the humid air of the terrestrial environment, in contrast to taste chemoreceptors that evolved in aquatic habitats; (8) the way in which the worms are able to cope with a major evolutionary constraint: the need to produce mucus for their locomotion, mucus for the most part being water; (9) the correlation between various body shapes (cylindrical, flat, etc.) on the one hand and water conservation and various terrestrial habitats (ranging from humid to rather dry) on the other hand, a cylindrical body considered to be more economical in terms of water conservation (Clark and Cowey 1958); (10) the manner in which the protonephridia adapted from an osmoregulatory system in aquatic ancestors to a resorptive system (saving water and/or eliminating metabolic wastes) in land flatworms; (11) the change from aquatic to terrestrial respiration. Impact and innovative aspects Phylogenetic studies, including those on flatworms in general and land flatworms in particular, are generally based on one-sided approaches, incorporating either morphological/anatomical data, or molecular data. Molecular studies might plot some morphological data on the resulting phytogenies to legitimize the molecular trees (‘pseudo-morphology’; cf Mooi and Gill 2010; Assis and Rieppel 2011; see also Williams and Ebach 2010). But for the land flatworms there are no studies that take an integrative approach with both kinds of data. This contrasts with the fact that many interesting scientific questions in this group of animals concern the evolution of their structures, key innovations, and adaptations (see above: Key innovations, adaptive radiation, and terrestrialization). Land flatworms likely are among the first groups of animals that during evolution have colonized the land zse.pensoft.net 546 Sluys, R.: The evolutionary terrestrialization of planarian flatworms Figure 2. Map of species richness in land planarians on an equal area grid map; maximum in red, minimum in dark blue (from Sluys 1998). Over the past 20 years new discoveries have added to the number of native and introduced species. Nevertheless, the current pattern of biodiversity remains very similar to the one shown in this map, although now, for example, about 20 new species are recognized in Europe, 40 in southeastern and southern Brazil and northeastern Argentina, nine in the Australian territories, and six in New Zealand (Sluys 2016). and have subsequently radiated extensively. According to fossil information arthropods would be the first ani¬ mals to have colonized the land, with atmospheric oxy¬ gen levels as the major driver of successful colonization (Ward et al. 2006). However, phylogenetic trees suggest flatworms as one of the early colonizers (cf. Hedges and Kumar 2009). This is generally neglected and, therefore, the study proposed here will contribute to a more bal¬ anced representation of the evolutionary history of ter¬ restrial animal life. The fossil record of flatworms is sparse and hard¬ ly provides adequate calibration points for a molecular clock (cf. Pierce 1960; Alessandrello et al. 1988; Ruiz and Lindberg 1989; Poinar 2003); calibration points may have to be based on paleogeographical information, am¬ ber fossils, coprolites, or estimates of mutation rates (cf. Blaxter 2009 and references therein). Thus, the proposed project will also form a first step towards our understand¬ ing of the absolute timing of the early terrestrialization of animal life. It will facilitate the test of the hypothesis that terrestrialization is linked to atmospheric oxygen levels, as was proposed for arthropods (Ward et al. 2006). The project may thus contribute to the first few insights into the terrestrialization of the Lophotrochozoa Halanych et al., 1995 because our current understanding is mainly re¬ stricted to the Ecdysozoa Aguinaldo et al, 1997 and the Deuterostomia Grobben, 1908 (cf. Labandeira 2005). Such issues may only be adequately addressed with the help of the phylogenetic trees generated during the re¬ search programme outlined here. These phylogenetic reconstructions will also form the long-awaited robust backbone for conservation biology studies in which land planarians function as indicator taxa (see Sluys 1999). Furthermore, triclad flatworms are a key group in his¬ torical biogeography because they do not possess larval dispersive stages. Therefore, they are excellent models for vicariance scenarios as explanations for current bio¬ geographic patterns. Thus, the research programme will facilitate tests of the hypothesis that plate tectonics has been a major factor in their historical biogeography (cf. Sluys 1994, 1995). For the morphological data I envision this programme to develop a formal knowledge representation (ontology) of planarian phenotypes and character states that leverages progress in this field (for a review, see Mabee et al. 2007) such that one of the outcomes will be a ‘Rosetta stone’ of concepts in planarian morphology that can be used not just to disambiguate characters and their states in the putative study but also to inform subsequent research questions in connecting genomic data and planarian phenotypes. zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 543-556 547 Phylogeny Knowledge on the phylogenetic relationships within the Tricladida is based on phylogenetic analyses of molecular and morphological datasets (for a review, see Sluys and Riutort 2018). Attention has been paid mostly to: (1) the relationships between the higher taxa within the Tricladida (cf Sluys 1989a; Carranza etal. 1998; Alvarez- Presas et al. 2008); (2) the affinities within the suborder of the Maricola (Sluys 1989b); (3) the higher taxon relationships within the freshwater planarians in general and within the family Dugesiidae Ball, 1974 in particular (Ball 1974; De Vries and Sluys 1991; Sluys 2001; Lazaro et al. 2009). Sluys and Kawakatsu (2006) explored the phylogenetic relationships within the freshwater families Dendrocoelidae Hallez, 1892 and Kenkiidae Hyman, 1937. More recent studies have resulted in an expanded character state matrix, including several new species and gene sequences, and in a robust phylogenetic tree for the higher taxa (Alvarez-Presas et al. 2008). For decades the evolutionary relationships within the group of land flatworms have been neglected. Partial taxonomic revisions of the group have been published but these were rarely based on a phylogenetic analysis (cf Ogren and Sluys 1998; Kawakatsu et al. 2005). It was only with the advent of molecular techniques that studies started to address the phylogenetic structure within the group (cf Alvarez-Presas etal. 2008; Carbayo etal. 2013). On the basis of the results of these molecular studies, and in combination with anatomical data, a revised higher classification of planarian flatworms was proposed (Sluys et al. 2009), in which the subordinal taxonomic rank of the terrestrial planarians was downscaled to the level of family, viz., the Geoplanidae. However, these molecular studies suffer from the fact that they only incorporate relatively few exemplar species and then only for a small selection of land planarian taxa. Methodology Separate and eombined analyses It is proposed here to use both molecular and morpho¬ logical data sets. To characterize their phylogenetic signal the character state matrices should be analyzed separately, using parsimony (morphology) and Bayesian methods (morphological as well as molecular). In addi¬ tion, molecular and morphological data sets should be combined into one joint analysis. With respect to mor¬ phological and molecular characters two approaches may be followed: (1) morphology is merely optimized in post-tree analysis of the molecular results, or (2) mor¬ phological and molecular characters are combined into one data matrix. The first approach is favoured in many recent studies and considered to be the only contribution of morphology to phylogenetic analysis by Scotland et al. (2003); but see Jenner (2004) for a rebuttal. It is here suggested that the second approach, i.e., combined anal¬ ysis, is applied. From an empirical perspective, it has been shown that morphology can have a profound effect on the com¬ bined analysis, irrespective of the fact that the number of molecular characters generally exceeds the number of morphological features (Jenner 2004; Assis 2009). Com¬ bined analyses may be different from the analyses of the separate data matrices and consensus trees may hide a phylogenetic signal that is generated by a total evidence analysis. Positive contributions of morphology to quanti¬ tative clade support measures in combined analyses have been observed for a large number of taxa (Jenner 2004). Timetree calibration As the fossil record of flatworms is sparse and does not provide adequate calibration points for a molecular clock, calibration of the phylogenetic timetrees has to be based on other kinds of data, such as, for example, paleogeo- graphical information (see above: Impact and innovative aspects). This means, for example, that we will be look¬ ing for closely related taxa a and b that are endemic to the areas A and B, respectively. In addition, we will be looking for those areas A and B inhabited by endemic taxa for which paleogeographic data indicate the time since the two areas have fragmented from a single ances¬ tral area. These two pieces of information, together with the molecular clock hypothesis, will enable one to date all cladogenetic and biogeographic events in the entire lineage of which a and b only form a part. Within the triclad flatworms there is a good number of such disjunctions, due to vicariance events, within species or between closely related species, that may be tested as possible calibration points. For example, land planarians of the genus Othelosoma Gray, 1869 are restricted to Africa and India and have attained their current distribution when India and Africa started to separate at about 150 Mya. Table 1 specifies the taxa, and their presumed vicariance events and divergence times that may profitably be used for calibration. One may perhaps be inclined to consider paleogeo- graphical calibrations to be less ideal than fossil calibra¬ tions. In point of fact, the opposite may be the case. Fos¬ sil-calibrated molecular clocks at best provide minimum dates (Wilke et al. 2009). Therefore, it has been argued that clocks are best calibrated with reference to the distri¬ bution of molecular clades and associated tectonics (e.g., Azuma et al. 2008; Heads 2014). Molecular dating is a rapidly developing field and there¬ fore there is currently no single best method; each approach has its advantages and disadvantages. It is also important to note that in the phylogenetic tree of the Platyhelminthes, the triclads constitute one of the crown groups (cf Baguna and Riutort 2004). This implies that the nodes below the Tricladida represent older taxa that therefore might be used as potential outgroups in the phylogenetic analysis. zse.pensoft.net 548 Sluys, R.: The evolutionary terrestrialization of planarian flatworms Datasets and phylogenetic analysis The molecular data matrix for this project may be derived from published data from GenBank, and new molecular sequences generated from fresh material of new taxa examined during the project. Previous studies have shown that the following genes provide the best resolution at the hierarchical levels of the phylogenetic tree of the triclads that form the focus of the putative research programme: nuclear 18S rDNA and 28S rDNA for resolving the deeper, more ancient branches in the tree, and mitochondrial COI to contribute signal on more recent splits (e.g. between and within genera) (Alvarez- Presas et al. 2008). Subsequently, another nuclear marker, the elongation factor 1 -alpha (EF), was added to the list of informative markers for an intermediate level between family and species, contributing to resolution of the relationships between members of the Geoplaninae subfamily (Carbayo et al. 2013). It has been suggested that many of the problems associated with the amplification and sequencing of planarian molecular markers may be solved by applying next generation sequencing (NGS) methodologies (Sluys and Riutort 2018). This new era has started already, and in recent years genomes and transcriptomes of freshwater planarians have already been sequenced (Egger et al. 2015; An et al. 2018; Grohme et al. 2018; Rozanski et al. 2018), contributing new anonymous nuclear markers that can be used as phylogenetic tools. There are even bioinformatic pipelines that can help in the discovery of these markers using NGS data as input (Frias-Eopez et al. 2016). This has already been applied in freshwater planarians, and will soon be used also for terrestrial fiatworms. Moreover, there is now available also information on other mitochondrial markers, in addition to COI, as whole mitogenomes were sequenced and annotated (Sakai and Sakaizumi 2012; Sola et al. 2015; Gastineau et al. 2019; Yang et al. 2019), contributing to the combination of nuclear and mitochondrial markers, thus obtaining better resolved phylogenies. The project requires a large morphological data matrix of all major taxa of land planarians (e.g., for the current 55 genera; cf Sluys et al. 2009). For this, the programme may be taking as leads the doctoral dissertations of both Dr E. Winsor (2003, James Cook University, Townsville) and Dr F. Carbayo (2003, University of Sao Paulo). Both researchers made detailed character codings and scorings for taxa of land planarians. Dr Winsor mostly focussed on Australian taxa, while Dr Carbayo concentrated on South American species, although neither used these scorings for a subsequent phylogenetic analysis. During the project character codings and scorings may be extracted from these two doctoral theses and, subsequently, be combined, refined and also supplemented by scorings for taxa not examined by Winsor or Carbayo. Definition and scoring of morphological character states should comply with the most recent insights (cf Sereno 2007). The morphological, molecular, and combined data matrix may be analyzed using maximum parsimony (e.g., TNT; Goloboflf et al. 2003), maximum likelihood (e.g., RaxME; Stamatakis 2014; IQtree; Nguyen et al. 2015), and Bayesian Inference (MrBAYES, Ronquist et al. 2012, and BEAST, Drummond et al. 2012). Inform¬ ative genes should first be analyzed separately, but also combined in the form of concatenated datasets. Substi¬ tution model parameters may either be selected using nested likelihood ratio tests or explored using reversible jump Markov chain Monte Carlo methods (jModeltest, Darriba et al. 2012; Partitionfinder, Eanfear et al. 2017). Saturation tests should be performed using DAMBE (Xia 2017). Molecular data may be analyzed also under a dynamic approach to homology. In the dynamic approach deline¬ ations are dependent upon the topology of the phyloge¬ netic trees on which they are optimized. In this context hypotheses of homology are part of phylogenetic hypoth¬ eses and are subject to the same optimality criteria as the trees, viz., minimisation of evolutionary transformation events. The computer program POY (ver. 4.0 beta 2635; Wheeler et al. 2006, Varon et al. 2010) allows for phy¬ logenetic tree searches under this dynamic homology or direct optimization approach. Aquatic planarians should form one of the outgroups for the phylogenetic analysis. Particularly the freshwater family Dugesiidae has been shown to share a close rela¬ tionship with the land fiatworms (see Sluys et al. 2009). But other freshwater groups, as well as marine triclads, should be included also as outgroups. Comparison with these freshwater and marine forms will reveal the adapta¬ tions that made possible the transition to land. Table 1. Taxa, and their presumed vicariance events and divergence times that may be used for calibration of the timetrees. Taxon Vicariant distribution Divergence time (Mya) Othelosoma species Africa (31 sp.)/lnclia (7 sp.) <150 Bipalium species Madagascar (23 sp.)/lndia & SE Asia (160 sp.) 90-50 Girardia species N. America (4 sp.)/ S. America (39 sp.) 3,5 genus G/rard/a/genus Dugesia N. & S. America (42 sp.)/Africa (21 sp.) 130-100 Dugesia species E. Med. (12 sp.)/W. Mediterranean (10 sp.) 38-3 Procerodes littoralis E. Atlantic/W. Atlantic 150 Foviella affinis E. Atlantic/W. Atlantic 150 Uteriporus vulgaris E. Atlantic/W. Atlantic 150 genus Amblyplana/genus Geoplana Africa (9 sp.)/S. America (64 sp.) 100 Romankenkius species S. South America (1 sp.)/Australia (12 sp.) 120 zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 543-556 549 Hypotheses testing Creeping sole The research proposed here seeks to explore the evolu¬ tion of terrestrialization of land flatworms in time and in correlation with presumed key morphological adap¬ tations. As regards the time axis, this will be important both for direct reconstructions of when terrestrializa¬ tion happened, as well as in the subsequent analysis of whether hypothesized key innovations co-vary in their location on the phylogeny with elevated diversification rates under a model of adaptive radiation (e.g., using the method of Ree 2005). It is here proposed that divergence dates estimates may be reconstructed from molecular data calibrated using previously published node ages (Hedges and Kumar 2009) in a relaxed molecular clock approach as implemented in BEAST (Drummond et al. 2012). As for covariance between traits (see above: Key innovations, adaptive radiation and terrestrialization), the general model will be that the radiation of fiatworms, with presumably repeated terrestrializations, constitutes a natural experiment such that morphological changes hy¬ pothesized to be associated with terrestrialization co-vary with and co-occur in ancestral state reconstructions on a phylogenetic tree. Sophisticated comparative methods to correct for autocorrelation (because the different lineages are related to each other) have been developed, including Bayesian statistical methods that allow for phylogenetic uncertainty (e.g., Pagel and Meade 2006). Understanding of the evolutionary dynamics and consequences of such innovations will contribute to the unraveling of their ge¬ netic basis and their role in speciation events and, conse¬ quently, the adaptive radiation of this group of animals. To give an example, the study will provide insight into the evolution of creeping soles, the latter defined as: “A fiat or ridged modified strip of epithelium on the ventral surface of geoplanid triclad fiatworms characterized by the presence of cilia .... which provides propulsive forces by ciliary or muscular action, or by a combination of both” (Sluys et al. 2009: 1773). Traditionally, a narrow creeping sole is considered as the primitive character state, while medium to broad soles and loss of a creeping sole are seen as derived states. However, narrow soles may also secondarily result from partial reduction, while absence of a creeping sole altogether may be the ancestral state, as aquatic forms lack one. However, the evolution of creeping soles has never been analyzed within a phylogenetic context; the research programme developed here will be the first to do so. If the traditional view holds true, one would expect to infer the narrow creeping sole as the character state at the root of the tree, with transitions to medium or broad soles, or loss of a creeping sole, nearer the tips. Under the opposite scenario, the absence of a creeping sole being the ancestral state, one would consequently expect this state to be reconstructed for the root of the phylogeny with alternately polarized state changes reconstructed nearer the tips. Likewise, expectations can also be formulated for body shape: if cylindrical body shapes indeed conserve water one would expect transitions to this morphology to coincide with, or follow terrestrialization when reconstructed on the phylogeny. In a similar way, all of the hypotheses mentioned above (see above: Key innovations, adaptive radiation and terrestrialization), as well as others that may arise during the study, can be tested. Respiration in planarians Introduction A particular crucial feature that may be among the most difficult to examine and to plot its character states on the phylogenetic trees concerns the adaptation to terrestrial respiration. Planarian flatworms possess neither circula¬ tory nor respiratory systems for transporting oxygen or digested food substances to the internal tissues. In these animals, oxygen is absorbed across the entire body wall and for this diffusion process water is required to dissolve oxygen and carbon dioxide in order to cross cell mem¬ branes. Clearly, this poses no problem for freshwater and marine planarians as they live in an aquatic habitat, but when ancestral planarians colonized the land, leaving this aquatic milieu must have formed a major hurdle, as the physical properties of water and air are so different. It is true that many land planarians live in habitats with a high humidity, but still these conditions greatly differ from a fully aquatic environment, while there are also terrestrial planarians that occur in mesophile and xerophile habitats (Froehlich 1955; Winsor et al. 1998). Thickness of flatworms The analysis of this subject is complicated by the fact that not much is known about the respiratory physiology of free-living flatworms in general and planarians in par¬ ticular. Furthermore, most of these studies concern aquat¬ ic species. A striking example is McNeill Alexander’s (1979) conclusion that the maximum possible thickness of a free-living flatworm is 1.0 mm or at most 1.5 mm. His conclusion is based on an argument that involves the calculation of the diffusion rate of oxygen. In philosoph¬ ical context McNeill Alexander’s type of functional ex¬ planation has been presented as an example of viability explanations, which are distinct from causal or historical explanations (Wouters 1995). McNeill Alexander’s (1979) calculation is based on a number of factors, viz., (a) partial pressure of oxygen in water, (b) rate of gas diffusion, (c) the fact that oxygen mostly will diffuse through the dorsal body surface, as the ventral surface is in contact with the substrate, (d) the diffusion constant for oxygen diffusing through con¬ nective tissue, (e) density of flatworm tissue. When the known or estimated values for these variables are used zse.pensoft.net 550 Sluys, R.: The evolutionary terrestrialization of planarian flatworms in a derivation using Pick’s law of diffusion, “This cal¬ culation indicates that the maximum possible thickness for a flatworm ... is about 0.5 mm if oxygen diffuses in only from the dorsal surface, or 1.0 mm if it diffuses equally from the ventral surface.... A similar calculation for a cylindrical turbellarian indicates that the maximum possible diameter would be 1.5 mm .... This is proba¬ bly why large flatworms are flat.” (McNeill Alexander 1979: 185). As flatworms rely entirely on oxygen diffusion through the surface, it is indeed advantageous to have a large sur¬ face area:volume ratio, i.e., to be flat and not cylindrical. In general, this holds true for aquatic species, while with¬ in species their thickness hardly or not at all increases with an increase in plan area of the body (Calow 1987). Another interpretation of the broad bodies of large land flatworms is that this facilitates capture and subdual of prey (cf Cseh et al. 2017), i.e., forms an evolutionary ad¬ aptation to new kinds of terrestrial prey not encountered by their aquatic ancestors. For an organism that relies on direct diffusion of oxygen Prosser (1973) provided a formula for the calculation of the thickness of the animal, based on the oxygen concentration of the medium, a diffusion coeflicient K, and the rate of oxygen consumption. When the calculation is done for both water and air (Table 2) it reveals that air-breathers can be six times thicker because there is more oxygen in air than in water (it should be noted that there is uncertainty about the units in which thickness is expressed). Nevertheless, the conclusion of McNeill Alexander (1979), as well as the calculation on which it is based, is open to a number of criticisms. First of all, it is certainly not the case that large flatworms are always very flat or thin. For example, “giant” freshwater species from Lake Baikal, such as Bdellocephala bathyalis Timoshkin & Porflrjeva, 1989, are certainly thicker than 0.5-1.0 mm {B. bathyalis measuring 3.3 mm in thickness in pre¬ served condition; Sluys et al. 1998). And also the large land planarians are usually thicker, such as Polycladus gayi Blanchard, 1845 (4 mm thick), Pseudogeoplana lumbricoides (Schirch, 1929) (3.5 mm), P. nigrofusca (Darwin, 1844) (3 mm), Geoplana rufiventris Schultze & Miiller, 1857 (2.3 mm) (Von Graff 1899). Although such measurements on the thickness are usually made on preserved specimens, and that thus fully stretched live specimens will be thinner, this does not necessarily com¬ promise the picture. It should be realized that most of the time the flatworms hide under stones or fallen logs, etc. and that during this resting period they are highly contracted, thus presumably approaching the condition of preserved specimens. Another example of a thick flatworm-like animal is the basal hiXdXQx'idin Xenoturbella bocki Westblad, 1950, which may reach a length of 2-3 cm and a thickness of 5 mm (Franzen and Afzelius 1987). Another objection that may be raised against McNeill Alexander’s (1979) calculation concerns his assumption that diffusion in flatworms is more or less equal to that in frog muscle and connective tissue, i.e., 2 X 10“^mm^atm“'s“'. However, it is doubtful that the mesenchyme and gut tissue—forming the major component of the planarian body—have the same diffusion constant as frog muscles and connective tissue. Evidently, the diffusion constant of planarian tissue will not be equal to that in water (6 x 10“^mm^atm“'s“') but perhaps 4 x 10“^mm^atm“*s“’ would be a value that is more realistic for flatworms. This would then imply a higher rate of diffusion and, consequently, would allow the worms to be thicker. For the rate (m) of oxygen consumption per unit vol¬ ume of tissue, McNeill Alexander (1979: 185) used 0.1 cm^ oxygen g“'h“’, “or a little more”. However, accord¬ ing to Hyman (1951: 207 and references therein) this may be higher, viz., 0.2-0.31 cm^ oxygen g“*h“*, that is “0.2-0.3 cc per gram per hour in adult worms.” Ac¬ cording to Moore (2006) it would be about 0.1-0.2 ml oxygen g“'h“' at 15 °C. A variable that is not taken into account by McNeill Alexander (1979) is the relationship between size and metabolic rate in flatworms. Although data are scant, generally respiratory rate decreases as size increases (Vernberg 1968). Although McNeill Alexander’s view on the maxi¬ mum thickness of flatworm-like animals has been ex¬ plicitly or implicitly endorsed (e.g., Ruppert et al. 2004; Moore 2006), he considered it merely “a very rough cal¬ culation in a textbook” and he conceded that “There are many doubtful assumptions in the calculation” (McNeill Alexander in litt., 22 August 1996). Respiratory pigments Apart from the doubtful assumptions in McNeill Alexander’s calculation, another explanation for the empirical fact that flatworms frequently are thicker than 0.5-1.0 mm may lie in the presence of Table 2. Data and results of thickness calculation according to Prosser’s (1973) formula: thicknesss = V(8 Cq(K7Vq 2)); oxygen con¬ centration in medium (C^) and K value (for muscle) from Prosser (1973); oxygen consumption (VO 2 ) converted from value of 0.2 ml/g/hr (= 0.0033 ml/g/min; see this paper). Medium Oxygen medium (ml/ml) K (muscle; cm2/min/atm) Oxygen consumption (ml/g/min) Thickness (units ?) Water 0.00651 0.000014 0.0033 0.014864234 Air 0.2095 0.000014 0.0033 0.084322613 zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 543-556 551 respiratory pigments, transporting oxygen across tissues. Hemoglobin is the most common respiratory pigment among invertebrates in general (Brusca and Brusca 2003). With respect to flatworms it has been reported mostly from parasitic helminths (Weber and Vinogradov 2001) as well as for a number of ecto- and endosymbiotic turbellarians, such as Paravortex Wahl, 1906, Triloborhynchus Bashiruddin & Karling, 1970, Cleistogamia Faust, 1924, Serita Cannon, 1982, Paranotothrix Cannon, 1982 (Jennings and Cannon 1987; Jennings 1988, 1997). Hemoglobin is known also from the free-living flatworms, viz., a species of Phaenocora Ehrenberg, 1835 (Vernberg 1968; Weber and Vinogradov 2001). It has been hypothesized that presence of hemoglobin in the entosymbiotic graflilid rhabdocoel Paravortex scrobiculariae (Graff, 1882) represents an adaptation to the oxygen-poor conditions to which this species is subjected, in contrast to the well-aerated habitats of other species, such as P. cardii (Hallez, 1908) andGraffillabuccinicolaiamQSon, 1897, which lack hemoglobin (Jennings 1981). Unfortunately, oxygen carrying components such as hemoglobin have not been studied in planarians. Winsor (1998) mentioned the possible respiratory role of uroporphyrins (other than hemoglobin) present in the rhabdoids of Platydemus manokwari de Beauchamp, 1962. Potential presence of hemoglobin may be deduced from the presence of non- epidermal pigments, often situated in or around specific organs, such as the brain and the pharynx (Jennings 1981, 1988; Jennings and Cannon 1987). In planarians, body colouration generally is due to the presence of granular pigments located in the mesenchyme directly below the dorsal epidermis (Sluys and Riutort 2018), while in most species of the freshwater genus Girardia Ball, 1974 the pharynx is also pigmented. Therefore, red pigment at other locations in the planarian body may point to the presence of hemoglobin. However, 1 have rarely observed pigment to be present at such other locations, at least in histological preparations, suggesting that hemoglobin is absent in triclad flatworms. Respiration in triclads and other free-living tnrbel- larians Oxygen consumption in planarians and other free-living turbellarians has been studied chiefly in freshwater spe¬ cies (cf Vernberg 1968; Heitkamp 1979) and only once in a terrestrial species (Daly and Matthews 1982). An impor¬ tant result that emerged from these studies is the relation¬ ship between body size and respiration, in that generally smaller animals have a higher rate of oxygen consumption than larger specimens, when determined on a weight-spe¬ cific basis; however, some species showed the reverse correlation, while for others no relationship could be es¬ tablished between body size and oxygen uptake (Vernberg 1968 and references therein; Heitkamp 1979). Rates of oxygen consumption may be determined and expressed in different ways and are generally influenced by the temperature of the habitat. The type of response to changes in temperature varies per species, as some spe¬ cies are eurythermal and others much more stenothermal in their ecological requirements (Vernberg 1968 and ref¬ erences therein). Therefore, respiration rate frequently is expressed as microliters- 02 -per-(milli-)gram-wet weight- per hour (|il 02 /(m)gAVW/h) at a particular temperature. Another way to express oxygen consumption is by calcu¬ lating the coeflicient b, or the regression slope of a particu¬ lar power function for the relation between body size and respiration. Unfortunately, the value of b varies much as it greatly depends on the size of the animals, their physiolog¬ ical condition, and many external factors (Heitkamp 1979). For five species of freshwater planarians (Dugesia gonocephala(DugQS, 1830), Crenobiaalpma(D3m, 1766), Polycelis nigra (Muller, 111 A), P felina (Dalyell, 1814), Schmidteapolychroa (Schmidt, 1861)) the following values were found for oxygen consumption (plO^/gAVW/h), measured at a temperature of about 15 °C: 170, 240, 135, 199,116, respectively (Vernberg 1968). Unfortunately, these values for triclad flatworms cannot be compared directly with those determined for various microturbellarians as these were calculated per milligram wet weight ( 11 IO 2 / mgAVW/h), resulting in the following values, measured at a temperature of 15 °C: 1.242 (Dalyellia viridis (Shaw, 1791)), 0.688 {Opistomum pallidum Schmidt, 1848), 0168 (Mesostoma ehrenbergi (Focke, 1836)), ranging between 0.298 and 0.700 (eight different populations of Mesostoma lingua (Abildgaard, 1789)) (Heitkamp 1979). The habitat temperature of the tropical terrestrial land planarian Bipalium kewense Moseley, 1878 is usually much higher than that of the aquatic triclads mentioned above, albeit that this invasive species has established itself outdoors in, for example, several North American states, the West Indies, Portugal, French Guiana, and France (Sluys 2016; Justine et al. 2018). The oxygen consumption of B. kewense specimens from outdoor localities in Arkansas, USA was determined at temperatures varying between 27-33 °C, which yielded respiration rates (plO^/gAVW/h) ranging between 113- 290 (Daly and Matthews 1982). It is noteworthy that these values are in the same order of magnitude as those determined for freshwater planarians. The coeflicient b based on oxygen consumption of entire specimens of B. kewense ranged between 0.686- 0.753, as measured at temperatures ranging between 27-33 °C (Daly and Matthews 1982). These values are in the same order of magnitude as those determined for the freshwater triclads Crenobia alpina (0.66), Dugesia gonocephala (0.82), and Polycelis felina (0.82) and the microturbellarians Mesostoma ehrenbergi (0.625), M. lingua (0.850), and Opistomum pallidum (0.880) (Heit¬ kamp 1979 and references therein). zse.pensoft.net 552 Sluys, R.: The evolutionary terrestrialization of planarian flatworms Terrestrial respiration Terrestrial flatworms face two problems that involve mutually conflicting adaptations, viz., desiccation and respiration. A cylindrical body, with less surface areawolume ratio, will minimize water loss but restricts diffusion of oxygen to the internal tissues. Probably this is the reason why smaller terrestrial species tend to be round or oval in cross-section, e.g., species of the land planarian genus Microplana Vejdovsky, 1890. Therefore, large species tend to be flattened to create a large surface areawolume ratio in order to facilitate diffusion of oxygen to the deep tissues. Nevertheless, the generally large and particularly long species of the land planarian subfamily Bipaliinae Von Graff, 1896 also have a more or less cylindroid body in cross-section. The partial pressure of oxygen in well-aerated water in equilibrium with air is 0.21 atm (McNeill Alexander 1979) and results in about 9 mgO^/l at 20 °C. Evidently, the actual amount of dissolved oxygen available at a particular aquatic habitat depends on the temperature, depth, altitude, and the mixing properties of the water (e.g., running water in shallow streams mixing better with air). The amount of oxygen in air is about 30 times that of water (Little 1990; Moore 2006), amounting to about 210 ml O 2 , weighing 280 mg, in 1 L of air (Schmidt- Nielsen 1975), while the diffusion rate of oxygen through air is much faster than through water (3 million times faster through air; Prosser 1973). In the present context it suflices to realize that, thus, availability of suflicient oxygen would not have formed a stumbling-block during an evolutionary transition from water to a terrestrial environment. But, clearly, the organisms needed to evolve respiratory adaptations enabling them to extract oxygen from the air, as opposed to their ancestors, which had evolved in an aquatic habitat. One such adaptation may be the production of mucus, which is secreted by both aquatic and terrestrial planarians. Mucus plays several important roles in the life of a planarian flatworm and is produced by various kinds of gland. Secretions from glands at the body margin produce a slime trail that facilitates the gliding movement of both aquatic and terrestrial triclads, effectuated by the propulsive force of cilia on the ventral body surface (Jones 1978). Many species of land planarians do not have such dedicated marginal glands, and various types of mucus are secreted from their creeping sole and/or ventral surface. Sticky mucus discharged by cephalic glands partakes in the capture of prey and has neurotoxic properties (cf Thielicke and Sluys 2019 and references therein). The mucus of triclad flatworms also has repellent properties as the worms are only rarely eaten by other animals since their surface secretions appear to have repugnatorial function (Hyman 1951; Winsor 1998). As mucus is produced also in aquatic triclads it could well be that these various functions already formed evolutionary preadaptations of similar functions in land planarians. Perhaps the secretion produced by the marginal adhesive zone is an exception as it has been suggested that it may not provide a lubricant for locomotion in the land planarians but form a moisture-retaining sealant in a resting animal (Winsor 1998). However, one function does not exclude the other. Production of surface secretions may also have formed a preadaptation for respiration in the terrestrial environment as it covers the body with a “watery” layer that presumably improves the uptake of oxygen. This may be related to the possible role, including respiration, of substances (porphyrins) in rhabdoids that are conspicuous in the dorsal/dorso-lateral epithelium (and microrhabdites over the ventral surface) of land planarians (see above). To the best of my knowledge, this aspect of mucus secretion and respiration in land flatworms has never received any attention. Evidently, uptake of oxygen through the body wall is only the first step in the respiration process. Hereafter, the oxygen needs to be transported to tissues deeper inside the planarian body. In the absence of respiratory pigments (see above: Respiratory pigments) this can be achieved only by means of diffusion. This implies that also the internal tissues must have a suflicient amount of water in order to be able to dissolve oxygen and carbon dioxide. Maintaining a suflicient level of hydration may be unproblematic for freshwater planarians but may require certain adaptations in marine and terrestrial forms. In particular, land planarians have no physiological or anatomical adaptations for water retention (Kawaguti 1932). On the one hand land planarians have practically no water-saving adaptations, while on the other hand they avoid wet environments and thus are considered to be stenohygric hygrocoeles (Froehlich 1955). It should here be noted that the situation may not be as absolute as suggested by Kawaguti’s (1932) findings, in that land flatworms may prevent desiccation by encasing themselves in mucus. Further, there may be also biochemical adaptations (secondary metabolism; see Campbell 1965) in land planarians for the elimination of carbon dioxide and nitrogenous waste that appear to facilitate water retention, such as the secretion of calcium salts (Percival 1925), which is consistent with being a stenohygric hygrocoele. In contrast to land planarians, freshwater species are subjected to osmotic influx of water and therefore must continuously regulate its volume with the help of their protonephridial system. One way to assess the hydration of the planarian body is to determine the osmolarity of the tissues, which is expected to vary inversely with the degree of hydration (Jones et al. 2004). The osmolarity of the freshwater planarians Schmidtea polychroa and Girardia dorotocephala (Woodworth, 1897) was 125-128 mOsm kg“' and 126 mOsm kg“\ respectively, while that of the marine triclad Procerodes littoralis (Strom, 1768) ranged between 217-272 mOsm kg“' (Jones et al. 2004 and references therein). Tissue osmolarity for the terrestrial planarian Arthurdendyus triangulatus (Dendy, 1896) ranged between 187.8-257.5 mOsm kg“‘, depending upon the laboratory conditions (relatively dry or fully hydrated) zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 543-556 553 under which the specimens were kept (Jones et al. 2004). These data show that tissue osmolarity in land flatworms is higher than that of freshwater planarians. This means that terrestrial flatworms are less hydrated than freshwater species and that the former are able to maintain a higher osmolarity because they are less prone to influx of water. 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Albano\ Sara-Maria SchnedP, Ronald Janssen^, Anita Eschner^ 1 Department of Palaeontology, University of Vienna, Althanstrasse 14, 1090 Vienna, Austria 2 Natural History Museum Vienna, Third Zoological Department, Burgring 7, 1010 Vienna, Austria 3 Malacology Section, Senckenberg Research Institute and Natural History Museum, Senckenberganlage 25, 60325 Franlfurt am Main, Germany http://zoobank.org/4F7BA0CB-813C-467B-B4FB-B6023834AE82 Corresponding author; Paolo G. Albano (pgalbano@gmail.com) Academic editor: M G/awZjrecM ♦ Received 13 July 2019 ♦ Accepted 24 September 2019 ♦ Published 3 December 2019 Abstract Rudolf Sturany, the curator of molluscs of the Natural History Museum of Vienna between the late 19* and early 20* century described 21 species of bivalves from the Red Sea collected by the pioneering expeditions of the vessel “Pola” which took place between 1895 and 1898. We here list and illustrate the type material of these species, provide the original descriptions, a translation into English, and curatorial and taxonomic comments. All species are illustrated in colour and with SEM imaging. To stabilize the nomenclature, we designate lectotypes for Gastrochaena weinkauffi, Cuspidaria brachyrhynchus, and C. dissociata, whose type series contained specimens belonging to other species. This paper concludes the series on the type specimens of marine molluscs described by Sturany from the “Pola” expeditions. Key Words Bivalvia, deep-sea, Indo-Pacific province, Pola expeditions Introduction The Red Sea lies in a key biogeographic position and shows a high level of endemism (Edwards and Head 1987; Janssen and Taviani 2015; DiBattista et al. 2016; Bogorodsky and Randall 2019). Separated by the Indian Ocean by a shallow sill, its biodiversity is hypothesized to be the result of recolonization from the Gulf of Aden after the Last Glacial Maximum (Taviani 1998). At the other end, the Suez Canal connects it to the Mediterra¬ nean Sea, which has been invaded by hundreds of tropi¬ cal species triggering the so-called Lessepsian invasion, one of the largest marine bioinvasions in the marine realm (Por 1978; Galil et al. 2015). Both aspects are of major biogeographic interest because they offer insights into the processes of dispersal, build up, and merging of faunas. Such processes cannot be properly studied without a thorough knowledge of the taxonomy of Red Sea species and the relations with those of nearby areas. It is not uncommon, for example, to encounter clearly non-native organisms in the Mediterranean Sea and not being able to assign them unambiguously to a species (e.g. Steger et al. 2018). Knowledge on type specimens is a fundamental first step for sound taxonomic work and the publication of type catalogues is recommended also by the International Com¬ mission on Zoological Nomenclature (ICZN 1999, 72F.4). Rudolf Sturany, curator of the molluscan collection at the Natural History Museum in Vienna (NHMW) between the late 19* and early 20* century, described 56 species of Red Sea molluscs based on the pioneering expeditions of the vessel “Pola” between 1895 and 1898, the first to ex¬ plore the deep-sea regions of the basin (Schefbeck 1996; Stagl 2012; Janssen and Taviani 2015). However, his work went unnoticed for long, possibly also due to the use of the German language (Huber and Eschner 2011). Copyright Paolo G. Albano et al. This is an open access articie distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the originai author and source are credited. 558 Albano, RG. et al.: Types of Sturany’s Red Sea bivalves The “Pola” samples are also an irreplaceable source of baseline data in this era of global change. Climate warm¬ ing, eutrophication, and coastal development are serious pressures on the Red Sea ecosystem (e.g. Raitsos et al. 2011; Naumann et al. 2015; Hall et al. 2018). Although not always recognized as such, the NHMW and the other insti¬ tutions that preserve similar historical samples are a stra¬ tegic asset in global change research (Johnson et al. 2011; Lister 2011; Albano et al. 2014; Dayan and Galil 2017). We here conclude our work to document the type spec¬ imens of Sturany’s marine taxa, started with the Red Sea gastropods (Albano et al. 2017) and continued with the deep-sea Mediterranean species (Albano et al. 2018), by treating the bivalves described from the Red Sea. We pro¬ vide high-quality optical and SEM images and, to help overcome the language barrier, we translate all original German descriptions into English. The year of publication of the Red Sea bivalves This paper focuses on the 21 bivalve taxa described by Stura- ny in 1899 (Table 1). This work was published in the volume 69 of the journal Denkschriften der kaiserlichen Akademie der Wissenschaften/Mathematisch-NaturwissenschaftUche as part of a separate series on reports of the “Commission fur oceanographische Forschungen” dated 1901. However, preprints (“Besonders abgedruckf’) with double pagination were published in 1899 (date on their frontispiece) and are present in the library of the Natural History Museum of Vi¬ enna; therefore, this should be considered the correct year of publication for all new names. A similar situation occurred for some Red Sea gastropods (Albano et al. 2017). Materials and methods The “Pola” material is entirely stored in the NHMW (Sta- gl et al. 1996). Type series of Sturany’s species were seg¬ regated. Most species were represented by holotypes or very small series. In the latter case, we identified the syn- types best matching the original description and selected lectotypes when appropriate to stabilize the nomenclature (ICZN 1999). A taxon list in alphabetical order with page and figure numbers in this paper is provided in Table 1. For each species, we provide references to the orig¬ inal description and figure, indicate the original local¬ ities, list the type material, reproduce the original de¬ scription, and translate it into English. All the mentioned inventory numbers refer to the Mollusca collection of NHMW. The systematic arrangement follows Bouchet et al. (2010). The reassessment of the current taxonomic status of Sturany’s names is beyond the scope of this paper, and we relied on the published literature to add some comments in this regard. The material studied by Sturany comes from off-shore “stations” (Table 1) and coastal “localities” (Tables 2, 3); we maintained this terminology. In the two tables, we report the collecting sites with their original orthography in German and a modern name between square brackets. The coordinates are those provided by Sturany. Any citation to the In¬ ternational Code of Zoological Nomenclature (ICZN 1999) should be considered to its online version, which includes all recent amendments. Most photographs were shot with a Nikon SMZ25 mi¬ croscope; large shells were photographed with a Canon 350D camera, a 50 mm lens, and extension tubes. SEM images were taken with a JEOE JSM-6610EV using low vacuum without any coating. Specimen measurements have been added for holotypes. Table 1. List of treated taxa in alphabetic order, with original and current name, current family placement, and figure in this paper. Taxon Current name Family Page Figure akabana, Cardita Centrocardita akabana (Sturany, 1899) Carditidae 564 4 bracheon, Raeta Raeta bracheon Sturany, 1899 Anatinellidae 571 8 brachyrhynchus, Cuspidaria Cuspidaria brachyrhynchus Sturany, 1899 Cuspidariidae 585 16 deshayesi, Gastrochaena Gastrochaena deshayesi Sturany, 1899 Gastrochaenidae 579 13 dissociata, Cuspidaria Cuspidaria dissociata Sturany, 1899 Cuspidariidae 587 17 eiachista, Limopsis Limopsis eiachista Sturany, 1899 Limopsidae 559 4 hypopta, Chione Timociea hypopta (Sturany, 1899) Veneridae 578 12 intracta, Lyonsia Poromya intracta (Sturany, 1899) Poromyidae 595 21 pexiphora, Gastrochaena Dufoichaena pexiphora (Sturany, 1899) Gastrochaenidae 582 14 potti, Cuspidaria Cuspidaria potti Sturany, 1899 Cuspidariidae 589 18 raveyensis, Dipiodonta Transkeia raveyensis (Sturany, 1899) Ungulinidae 575 11 siebenrocki, Amussium Parvamussium siebenrocki (Sturany, 1899) Pectinidae 561 2 siebenrocki, Teiiina Jitiada bertini (Jousseaume, 1895) Tellinidae 573 9 squamosina, Tridacna Tridacna squamosina Sturany, 1899 Cardiidae 566 5 steindachneri, Amussium Propeamussium steindachneri (Sturany, 1899) Pectinidae 563 3 steindachneri, Cuspidaria Cuspidaria steindachneri Sturany, 1899 Cuspidariidae 591 19 subcandidus, Soiecurtus Soiecurtus subcandidus Sturany, 1899 Solecurtidae 573 10 suiphurea, Scintiiia Scintiiiuia iutea (Lamarck, 1818) Galeommatidae 566 6 thaumasia, Pseudoneaera Pseudoneaera thaumasia Sturany, 1899 Cuspidariidae 593 20 variabiiis, Scintiiia Scintiiiuia variabiiis (Sturany, 1899) Galeommatidae 569 7 weinkauffi, Gastrochaena Lamychaena weinkauffi (Sturany, 1899) Gastrochaenidae 583 15 zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 557-598 559 Table 2. Off-shore stations of the “Pola” expedition with bivalves (from Sturany 1899). Station Locality Coordinates Depth number [m] 9 “sudlich von Yenbo” [south of Yanbu’ al Bahr, Saudi Arabia] 23°2rN, 3T3TE -791 27 - SS-Al'N, 37°23'E -747 41 “zwischen Mersa Halaib und Jidda” [between Halayeb, Egypt, and Jeddah, Saudi Arabia] 22°4'N, 38°E -2160 44 “vor Jidda” [off Jeddah, Saudi Arabia] 2r36'N, SS-SS'E -902 47 “bei Yenbo” [Yanbu’ al Bahr, Saudi Arabia] 23°4rN, 38°9'E -610 48 “vor Yenbo” [off Yanbu’ al Bahr, Saudi Arabia] SA-S'N, 37°45'E -700 51 “bei Sherm Sheikh” [near Abu Ghusun, Egypt] 24°15'N, SS-SY'E -562 56 “bei Mersa Dhiba” [Mersa Dhiba, Egypt] SS-SG'N, 34°55'E -582 61 “nachst den Hassani-lnsein” [nearAI Hasani, Saudi Arabia] 24°35'N, 36°5rE -828 72 “bei der Insel Shadwan” [Shadwan Island, Egypt] 27°25'N, SrSO'E -1082 76 “sudlich der Insel Senafir” [south of Sanafir Island] SY-AS'N, 34°47'E -900 81 “unweit von Ras Abu Massahrib, Noman Insel” [close to Ras Abu Massahrib, Noman Island, Saudi Arabia] SG-SA'N, 35°33'E -825 87 “bei Ras Mallap im Golfe von Suez” [Ra’s Mal’ab in the Gulf of Suez, Egypt] 29°7.6'H, 32°56'E -50 94 “bei Nawibi im Golfe von Akabah” [Nuweiba, Gulf of Aqaba, Egypt] 28°58.6'N, 34°43.7'E -314 96 “nordlicher Theil des Golfes von Akabah” [northern part of the Gulf of Aqaba] 29°13.5'N, 34°47.8'E -350 106 “sudlich von Jidda” [south of Jeddah, Saudi Arabia] 2r2'N, 38°41A'E -805 109 “westlich von Jidda” [Jeddah, Saudi Arabia] 2ri9'N, 37°39'E -890 114 “zwischen Suakim und Lith” [between Suakin, Sudan, and Al Lith, Saudi Arabia] 19°38'H, 37°55'E -535 117 “sudlich von Raveya” [south of Raveya, Sudan] 20‘’16.9'N, SY-SS.S'E -638 121 “westlich von Kunfidah” [west of Al Qunfudhah, Saudi Arabia] 18°51.9'N, 39°5.4'E -690 128 “bei Akik Seghir” [Akik Seghir, Eritrea] 18°7.7'N, 39°11.2'E -457 130 “westlich von Kunfidah” [west of Al Qunfudhah, Saudi Arabia] 19°17'N, SG-SY'E -439 138 “Ostlich von Akik Seghir” [east of Akik Seghir, Eritrea] 18°3'N, 40°14.7'E -1308 145 “Ostlich von J. Dahalak” [east of Dahlak Island, Eritrea] 16°2.6'N, 4ri3.5'E -800 156 “nOrdlich von Jidda” [north of Jeddah, Saudi Arabia] 22°5rN, 38‘’2'E -712 170 “bei der Insel Noman” [Noman Island, Saudi Arabia] 27°0.2'N, 35°17.6'E -690 176 “bei Koseir” [El Quseir, Egypt] SS-SY'N, 34°36.VE -612 Table 3. Coastal localities of the “Pola” expedition with bivalves (from Sturany 1899). Locality number Locality Coordinates/region 10 “Nawibi” [Nuweiba, Egypt] Gulf of Aqaba 12 “Dahab (Mersa Dahab)” [Dahab, Egypt] Gulf of Aqaba 16 “Shadwan-lnsel” [Jazirat Shakir, Egypt] Northern Red Sea, 28-2E)°H 25 “Sherm Sheikh (Mersa Sheikh)” [near Abu Ghusun, Egypt] 26-24°N 27 “Port Berenice” [Berenice Troglodytica, Egypt] 24-22°N 30 “Mersa Halaib” [Halayeb, Egypt] 24-22°N 31 “Jidda (Djeddah)” [Jeddah, Saudi Arabia] 22-20°N 32 “Raveiya (Mahommed Ghul)” [Gul Mohammad, Saudi Arabia] 22-20°N 37 “Ras Turfa” [near Jazan, Saudi Arabia] 18-16°N 41 “Massawa (Massaua)” [Massawa, Eritrea] 16-1TN 43 “KamaranJnsel” [Kamaran Island, Yemen] 16-1 rN Systematic list of taxa Family Limopsidae Dali, 1895 Limopsis elachista Sturany, 1899 Figure 1 Sturany 1899; 268, plate IV, figures 1-4. Original localities. Station 48, otf Yanbu’ al Bahr, Saudi Arabia, 24°5'N, 37°45'E, -700 m. Station 106, south of Jed¬ dah, Saudi Arabia, 21°2'N, 38°41.4'E, -805 m. Station 117, south of Raveya, Sudan, 20°16.9'N, 37°33.5'E, -638 m. Type material. Syntypes: NHMW 84346: 2 valves, station 48; NHMW 84347: 3 valves, station 106; NHMW 84348: 2 specimens, station 117 (one specimen in original figure). Original description. Von den Stationen 48, 106, 117 (638-805 m); einige wenige Schalen. Die Schale ist winzig klein, schwach gewolbt, ein wenig schief gewachsen, dock nahezu kreisformig, so hoch wie breit. Der Ob err and ist gerade und wird von den ein wenig aus der Mitte nach vorne geriickten Wirbeln uberragt; Vorder-, Unter- und Hinterrand sind gerundet. Die dussere Sculptur besteht aus einer zarten, aber deutlichen Streifung im Sinne des Wachsthums, sowie aus Radialstreifen, die entweder nur die vordere und mittlere Partie der Schale auszeichnen, Oder, was die Regel ist, bis riickwarts reichen; die davon betrojfenen Stellen zeigen also ein femes Gitterwerk. Die Grundfarbe der Muschel ist schmutzigweiss bis gelblich; dariiber ziehen in der Regel drei radial gestellte, gelbbraune Bander, die jedoch von wechselnder Breite sind und in verschiedener Combination fehlen konnen. zse.pensoft.net 560 Albano, RG. et al.: Types of Sturany’s Red Sea bivalves Figure 1. Limopsis elachista Sturany, 1899, Station 48, south of Raveya, Sudan, -638 m. A, B, H, J Original figures. C-F, G, I, K-P Figured syntype NHMW 84348: right valve interior (C, D), exterior (E, F), and hinge detail (G, I); left valve exterior (K, L), interior (M, N), and hinge detail (O, P). Q Original label. Scale bars; 1 mm (C, I, K, P); 0.2 mm (G, O). Das Innere der Schale ist vor Allem mit einem rela- tiv krdftigen Schloss ausgestattet. Dieses besteht in jed- er Schale aus 7 Zdhnen, und ist diese Zahnreihe in der Wirbelgegend unterbrochen, so dass die Formel 3:4, re¬ spective 4:3 zu verzeichnen ist; mitunter gesellt sich zu den 7 normalen Zdhnen in der rechten Schale noch je ein ganz kleiner Zahn an den beiden dussersten Enden der Reihe. Das Innere der Schale ist ferner noch durch eine stark gekerbte Peripherie und durch eine verwischte Ra- dialstreifung ausgezeichnet. Die Ldnge und Hdhe der Muschel misst 3,5-3,7 mm, die Dicke betrdgt circa 2.2 mm. Es sindnur wenigExemplare, die bei der Abfassung der Diagnose in Betracht kommen konnten. Wie sehr trotzdem die oben angedeuteten wechselnden Charaktere der neu- en Art bei den verschiedenen vorliegenden Schalen sich combiniren, mdgen die folgenden Beispiele zeigen. Eine linke Schale von Station 48 ist bdnderlos und zeigt hauptsdchlich in ihrer hinteren Partie die Gitter- sculptur; eine zweite (rechte) Schale von derselben Sta¬ tion ist allenthalben gegittert und hat ein breites Mittel- band, wdhrend die seitlichen Radialbdnder nur schwach ausgebildet sind. Von der Station 106 liegen zwei rechte, allenthalben gegitterte Schalen vor; bei der einen ist nur das hintere Radialband ausgebildet, die andere ist wied- er bdnderlos. Von Station 117 habe ich das hier abgebil- dete, mit drei Bandern ausgezeichnete Exemplare vor mir (deren Bezahnung sich ausdrucken Idsst mit der Formel; zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 557-598 561 rechts 5 + 4, links 4 + 3), sowie ein solches, bei dem das vordere Band fehlt. Die neue Art ist verwandt mit L. torresi Smith aus der Torresstrasse. Translation. From stations 48, 106, 117 (638-805 m); a few valves. The shell is tiny, slightly arched, a little oblique, but almost circular, as high as long. The dorsal margin is straight and overlapped by the umbo, which is slightly tilted forward; the anterior, ven¬ tral and posterior margins are rounded. The outer sculp¬ ture consists of a delicate but distinct striation consisting of both growth [concentric ridges], as well as of radial costae, which are expressed only over the anterior and central parts of the shell, or, as is the rule, reach poste¬ riorly; the affected areas show a fine reticulate structure. The basic colour of the shell is dirty white to yellow¬ ish; usually there are three radial yellow-brown rays of varying expression across the median area which, how¬ ever, can be missing in different combinations. The interior of the valve is equipped with a relatively strong hinge. This consists in each valve of 7 teeth, and this row of teeth is interrupted in the umbo region, so that the formula 3:4, respectively 4:3 is recorded; sometimes a very small tooth joins the 7 normal teeth in the right shell at the two extreme ends of the row. The interior of the shell is also characterized by a strongly notched periphery and blurred radial incisions. The length and height of the shell measure 3.5-3.7 mm, the thickness is about 2.2 mm. There are only a few specimens that could be consid¬ ered when writing the diagnosis. Nevertheless, the above indicated variable characters of the new species may be combined in the various valves present, as the following examples may show. A left valve of Station 48 lacks colour bands and shows the reticulate sculpture mainly over the posterior area; a second (right) valve from the same station is reticulate on all sides and has a wide central band, while the lateral radial bands are weak. From station 106 there are two right valves, entirely reticulate sculptured; in one, only the rear radial band is formed, the other is again without bands. From sta¬ tion 117,1 have the three-banded specimens pictured here (dentition can be expressed by the formula: right 5 + 4, left 4 + 3), as well as one in which the front band is missing. The new species is related to L. torresi Smith from the Torres Strait. Family Pectinidae Rafinesque, 1815 Amussium siebenrocki Sturany, 1899 Figure 2 Sturany 1899: 269-270, plate IV, figures 5-8. Type locality. Station 72, Shadwan Island, Egypt, 27°25'N, 34°30'E, -1082 m. Type material. Eectotype: NHMW 84355: 1 valve, sta¬ tion 72, designated by Dijkstra and Janssen (2013). Para- lectotypes: NHMW 84177: 1 valve, station 72; NHMW 84353: 1 valve, station 44; NHMW 84354: 1 valve, station 48; NHMW 84356: 5 valves; NHMW 84357: 5 valves; NHMW 84358: 10 valves. Original description. Von den Stationen 44, 48, 72, 91, 106, 109 (700-1082 m); einige wenige Schalen. Die Muschel ist klein, ungleichseitig, sehr wenig gewolbt, von fast kreisformiger Gestalt, schmutzig weiss- er Oder gelber Farbe aussen und milchweisser Farbe in- nen. Der fein zugespitzte Wirbel ist mittelstandig; unter ihm liegt an der Schlossleiste die kleine Ligamentgrube. Die rechte Schale ist kleiner undflacher als die linke, ist aussen gleichfbrmig concentrisch gestreift in ihrer Hauptpartie und mit Radialsculptur versehen auf dem vorderen Ohrchen, indem ndmlich hier dicht aneinan- dergereiht 6-8 beschuppte Rippchen vom inneren Winkel nach dem convexen vorderen Ende des Ohrchens ziehen. Das hintere Ohrchen besitzt gleichsam als Fortsetzung der allgemeinen concentrischen Streifung feine, hier fast senkrecht gestellte Linien. Die Innenseite der rech- ten Schale besitzt 10 weisse Radialrippchen, die knapp vor dem Rande knotig verdickt endigen, und ausser ih- nen lassen sich meist noch die Andeutungen von je ein- er Rippe an der inneren Basis der Ohrchen constatiren. Die 10 Hauptrippen scheinen nach aussen als weisse Radiallinien schwach durch. Der Oberrand ist dusserst schwach gekerbt. Die linke Schale ist grosser und aussen ganz anders sculptirt. Es findet sich hier ausser der concentrischen Streifung noch eine sehr wechselnde, nichts weniger als constante Anzahl von Radialrippen vor. Einige davon beginnen in kurzer Entfernung vom Wirbel, andere etwa erst in der Mitte der Schalenhdhe; bei alien ist aber an ihren Kreuzungspunkten mit der concentrischen Streifung eine schwache Schuppenbildung zu constatiren. An den Ohrchen sind wieder die Querstreifen der Hauptpartie in senkrechter Richtung fortgesetzt, am vorderen Ohrchen sogar ein paar Radialrippen vorhanden. Das Innere der linken Schale ist glanzend; hier tritt die Berippung in gleicher Anzahl auf wie in der rechten Schale; aber die weissen Rippen reichen hier nicht bis hart an den Rand, sondern endigen mit ihren Verdickungen schon etwas ent- fernter davon. [Tabelle mit Mafiangaben!] Abweichend von dem in der Diagnose erwdhnten re- gelrechten Verhalten erscheint eine rechte Schale von der Station 44 (6,9 : 7,0 mm); hier schieben sich zwischen die vorderen Radialrippen der Innenseite noch 3 ganz kurze Rippchen, welche, da sie unten am Rande stehen, die relativ weiten Abstande von je 2 Rippenendigungen gleichsam ausfullen. Die neue Art ist verwandt mit der Tiefsee-Form des Mittelmeeres, A. hoskynsi Forbes. Translation. From stations 44,48, 72, 91,106,109 (700- 1082 m); a few valves. zse.pensoft.net 562 Albano, RG. et al.: Types of Sturany’s Red Sea bivalves Figure 2. Amussium siebenrocki Sturany, 1899, Station 72, Shadwan Island, Egypt, -1082 m. A, D, G, I Original figures (rotated by 180°). B, C, E, F, H, J, L Lectotype NHMW 84355: right valve exterior (B, C), interior (H), and hinge detail (L). Left valve exterior (E, F) and interior (J). K Original label. Scale bars: 1 mm (B, E), 0.1 mm (L). zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 557-598 563 The clam is small, inequilateral, slightly curved, of al¬ most circular shape, dirty white or yellow outside and milky white inside. The finely pointed beak [umbo] is central; be¬ low there is the small ligament pit at the hinge margin. The right valve is smaller and flatter than the left, exter¬ nally the main part (disc) is uniformly concentrically ridged but with radial sculpture on the anterior auricle, of 6-8 scaly ribs radiating from the inner angle to the convex anterior end of the auricle. The posterior auricle, as a continuation of the general concentric striation, possesses fine lines, here almost vertical. The inside of the right shell has 10 white radial ribs, which terminate with a thick nodule just before the margin, and besides these, the traces of one rib on the inner base of the auricles can usually be ascertained. The 10 main ribs appear on the outside as weak white radial lines. The upper margin is very slightly notched. The left valve is larger and externally sculptured very differently. In addition to the concentric striation, there is also a very variable, nothing less than constant number of radial ribs. Some of them begin at a short distance from the umbo, others only in the middle of the valve; in all, however, faint scales can be observed at their points of intersection with the concentric ridges. At the auricles, the horizontal ridges of the main part are continued in the vertical direction, on the anterior auricle even a few radial ribs are present. The inside of the left shell is shiny; here the ribs occur in the same numbers as in the right valve; but the white ribs do not closely reach the edge, but end with their thickenings a little before. [Table with dimensions] Deviating from the normal behaviour mentioned in the diagnosis, a right shell appears from the station 44 (6.9: 7.0 mm); here 3 very short ribs raise in between the ante¬ rior radial ribs of the internal side, which, since they are at the edge, they almost fill the relatively wide distances of two ridge ends. The new species is related to the deep-sea form of the Mediterranean A. hoskynsi Forbes. Amussium steindachneri Sturany, 1899 Figure 3 Sturany 1899: 269, plate IV, figures 9-12. Type locality. Akik Seghir, Eritrea, 18°7.7'N, 39°11.2'E, -457 m. Type material. Eectotype: NHMW 84350: 1 valve, station 128, designated by Dijkstra and Janssen (2013). Paralectotypes: NHMW 84351: 5 valves, station 128; NHMW 84352: 2 valves, station 130; NHMW 92602: 3 valves, station 128. Original description. Von den Stationen 128 und 130 (439-457 m); einzelne Schalen. Die Muschel ist klein, ungleichschalig, fast kreisfor- mig, schwach gewolbt und glanzend, besitzt zarte, fein zugespitzte Wirbel und darunter in der Schlossleiste eine dreieckige Ligamentgrube. Die rechte Schale ist kleiner, aber dicker als die linke, ist milchweiss und durchscheinend. An ihrer ziemlich glatten, nur von ein paar stdrkeren, aber unregelmdssig auftretenden Anwachslinien durchzogenen, sehr stark glanzenden Aussenseite scheinen 6 weisse Rippen durch, die an der Innenseite radial angeordnet sind und bis an den Rand reichen, wo sie mit schwachen, knopjfbrmigen Verdickungen endigen. Uberdies verlduft iiber die innere Basis eines jeden Ohrchens noch eine Rippe, die allerdings nur schwach entwickelt ist und nach aussen kaum durchzuscheinen vermag. Das vordere Ohrchen ist vorne abgerundet (convex) und schwach quergestreift, d. i. concentrisch mit dem Vorderrande; das hintere istfast rechtwinkelig abgestutzt. Der Oberrand der rechten Schale ist zart gekerbt. Die linke Schale ist grosser, aber dunner als die rechte, ist glatt bis auf mikroskopisch feine Spuren von Quer- und Radialstreifen und stark durchscheinend. Durch zahlreiche uber die ganze Aussenseite verbreitete Flecken von weisser Farbe und hauptsdchlich gegen den Rand zu auftretende Flecken oder Streifchen von gelber oder orangerother Farbe gewinnt die etwas mehr gewolbte Schale ein charakteristisches Aussehen, das noch erhoht wird durch die krdftig Orangeroth oder gelb durchblickenden Radialrippen der Innenseite. So wie in der rechten Schale sitzen auch in der linken Schale 6 knotig verdickte Hauptrippen, welche in der Wirbelgegend, nicht weit vom Schlossrande entfernt, ihren Ursprung nehmen und, radial verlaufend, in einiger Entfernung vom convexen Rande endigen: ferner kommen auch hier noch zwei kleinere Rippchen an der inneren Basis der Ohrchen hinzu. Das vordere Ohrchen ist hier ein wenig concav, das hintere rechtwinkelig abgestutzt. Beide sind zum Unterschiede vom Haupttheile der Schale etwas deutlicher senkrecht gestreift. [Tabelle mit Mafiangabenl] Angesichts der bunten linken Schale dieser neuen Art wird man an die gefleckte Form erinnert, die Dali von seinem A. pourtalesianum erwdhnt. Dass im Gegensatze zu den Verhdltnissen der rech¬ ten Schale die inneren Radialrippen der linken Schale weit entfernt vom Unterrande endigen, steht im Zusam- menhange mit der grosseren Ausdehnung, dem grosser- en Umfange der linken Schale. Sind ndmlich die Scha¬ len zusammengeklappt, so decken sich gewissermassen Anordnung und Lange der beiderseitigen Radialrippen vollstandig und ragt der glatte, radienlose Unterrand der linken Schale um so viel hervor, als diese Schale eben grosser ist. Diese Thatsache ist sowohl bei A. steindach¬ neri m. wie bei der folgenden Art zu constatiren. Translation. From stations 128 and 130 (439-457 m); single valves. The clam is small, inequilateral, almost circular, slight¬ ly arched and shiny, and has delicate, finely pointed beaks and a triangular ligament pit beneath in the hinge margin. zse.pensoft.net 564 Albano, RG. et al.: Types of Sturany’s Red Sea bivalves Figure 3. Amussium steindachneri Sturany, 1899, Station 128, Akik Seghir, Eritrea, -457 m. A, D Original figures (rotated by 180°). B, C, E, F Lectotype NHMW 84350: right valve exterior (B), interior (C, E), and hinge detail (F). G Original label. Scale bars: 1 mm. The right valve is smaller, but thicker than the left, and is milky white and translucent. Its fairly smooth, very strongly glossy exterior, is crossed only by a few thick¬ er but irregular growth lines. Six white ribs are arranged radially on the inside and reach the edge where they end with weak, button-shaped thickenings. Moreover, above the inner base of each auricle there is a rib which is only weakly developed and is hardly visible on the outside. The anterior ear is anteriorly rounded (convex) and slightly striated, i.e. concentric with the margin; the rear is truncated almost at right angles. The upper margin of the right shell is delicately incised. The left shell is larger, but thinner than the right, and is smooth, except for microscopic traces of transverse and radial ridges, and highly translucent. Through numerous white patches spread all over the outside and marks or lines of yellow or orange-red colour appearing towards the edge, the slightly more rounded shell acquires a char¬ acteristic appearance, which is enhanced by the strong orange-red or yellow-looking radial ribs on the inside. There are six nodular thickened main ribs in the right and the left valve, which originate in the umbo region, not far from the edge of the hinge, and terminate radially, in some distance from the convex margin; additionally, there are two smaller ribs to the inner base of the auricles. The anterior auricle is a little concave, the posterior rectangular. In contrast to the main part of the valve both are slightly more vertically striped. [Table with dimensions] Considering the colourful left valve of this new spe¬ cies, one is reminded of the spotted shape which Dali mentioned in A. pourtalesianum. In contrast to the proportions of the right shell, the inner radial ribs of the left valve are ending far away from the ventral margin, which is related to the greater elongation and size of the left valve. If the valves are paired, the arrangement and length of the radial ribs on both sides completely coincide, and the smooth, low¬ er edge of the left shell, without radial sculpture, pro¬ trudes as much as this valve size. This fact can be stated for both A. steindachneri m. and the following species \A. siebenrocki\. Family Carditidae Femssac, 1822 Cardita akabana Sturany, 1899 Figure 4 Sturany 1899: 267-268, plate III, figures 10-12. zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 557-598 565 Figure 4. Cardita akabana Sturany, 1899, northern part of the Gulf of Aqaba, -350 m. A, D, F Original figures. B, E, C Figured syn- type NHMW 84343; right valve exterior (B), interior (E), and hinge detail (C). G Original label. Scale bars: 3 mm (B); 2 mm (C). Type locality. Station 96, northern part of the Gulf of Aqaba, 29°13.5'N, 34°47.8'E, -350 m. Type material. Syntypes: NHMW 84343: 2 valves, sta¬ tion 96 (one specimen in original figure). Original description. Von Station 96 (350 m); zwei einzelne linke Schalen. Die Schale ist gross, abgerundet, aufgeblasen und un- gleichseitig, aussen ockergelb mit hellen, unregelmdssig vertheilten Flecken, innen rein weiss. Der Wirbel ist nach innen und vorne gedreht und uber- ragt den Schlossrand um ein Bedeutendes; vor ihm liegt vertieft eine herzfdrmige, gestreifte Lunula. Vom Wirbel ziehen radial angeordnet 23 Rippen zum Rande. Sie sind meist gleich breit und breiter als Hire vertieft liegenden und undeutlich oder schwach quergestreiften Zwischen- rdume; in der hinteren Partie der Schale allerdings konnen mitunter die Rippen (etwa 5-6 an Zahl) weniger breit sein und dafur die Zwischenrdume relativ weiter von einander abstehen. Die Rippen sind dicht mit geldrollenartig an- geordneten Querwulsten oder Scheiben besetzt, die umso grosser sind, je weiter sie vom Wirbel entfernt liegen. Der Rand der geojfneten Muschel ist kreisformig, und nur am Ubergange des Hinterrandes in den Unterrand ist eine schwache Winkelung zu verzeichnen. Entsprech- end den Endigungen der Radialrippen sind die Pander stark crenelirt. Das Schloss der linken Schale besteht aus einem stumpfen Zahn, der direct unter dem Wirbel aus einer Schlossleiste hervortritt, und aus einer kleinen zahnartigen Erhebung vor demselben am Oberrande dort, wo die erste kurze Radialrippe endigt. Hinter dem Mittelzahn liegt eine lange und tiefe, dreieckige Grube, und auf diese folgt ein longer, dicker, bogiger und lamellenartiger Hinterzahn, der von dem das dussere Ligament tragenden hinteren Oberrande noch durch eine Vertiefung getrennt und oben wie unten fein quergestreift ist. Unter der Schlossleiste liegt die tiefe Aushohlung der Wirbelgegend. Die eine linke Schale ist 26,2 mm lang, 21,1 mm hoch und 12,2 mm dick; die andere (ebenfalls linke) misst 28,7, respective 30,2 und 14,1 mm. Die neue Art, von der mangels rechter Schalen das Schloss leider nur unvollstandig beschrieben werden konnte, erinnert einigermassen an Cardita cardioides Rve. Translation. From Station 96 (350 m); two single left valves. The shell is large, rounded, inflated and inequilateral, ochre coloured outside with bright, irregularly distributed spots, inside pure white. The umbo turns inward and forward and clearly pro¬ trudes the edge of the hinge, it is reclined with a heart- zse.pensoft.net 566 Albano, RG. et al.: Types of Sturany’s Red Sea bivalves shaped, striated lunula. Twenty-three radial ribs run from the beak to the margins. They are usually the same width or wider than the interspaces which are indistinctly or weakly radially striate; over the posterior part of the shell, however, sometimes the ribs (about 5-6 in num¬ ber) may be less wide and the interspaces stand farther apart. The ribs are densely covered by many bar-shaped varices or discs, which get larger the farther they are away from the beak. The margin of the open bivalve is circular, and only at the transition from the posterior to the ventral edge is a weak angulation recorded. Coinciding with the ends of the radial ribs the edges are strongly crenulate. The hinge of the left valve consists of an obtuse tooth, which emerges directly underneath the beak from the hinge margin, and of a small tooth-like bump at the dor¬ sal margin where the first short radial rib ends. Behind the middle tooth lies a long and deep triangular pit, followed by a long, thick, curved and lamellar posterior tooth, which is separated from the posterior upper margin, sup¬ porting the outer ligament by a depression, and is finely striated above and below. Below the hinge margin, the deep cavity of the umbo area lies. The first left valve is 26.2 mm long, 21.1 mm high and 12.2 mm thick; the other (also left) measures 28.7, respectively 30.2 and 14.1 mm. The new species, which unfortunately could not be described completely due to the lack of right valves, is reminiscent of Cardita cardioides Reeve. Family Cardiidae Lamarck, 1809 Tridacna elongata var. squamosina Sturany, 1899 Figure 5 Sturany 1899: 283-284, not figured. Type locality. Yemen, Red Sea, Kamaran IsL, ca 15°17'N; 42°37'E, shallow water. Type material. Lectotype: NHMW 107075: 2 valves (1 specimen), station 43 designated by Huber and Eschner (2011). Paralectotypes: NHMW 107076: 8 valves (4 specimens), station 12; NHMW 107077: 4 valves (2 specimens), station 10. Original description. Die stattliche Reihe der vorliegenden Schalen veranlasst mich, die in der Reeve’schen Monographie abgebildete Tr. compressa als Jugendform der elongata Lm. aufzufassen und ihren Namen einzuziehen, hingegen eine Varietdt besonders hervorzuheben, die systematisch zur Tr. squamosa Lm. hinuberfuhrt. Diese mit squamosina nov. var. zu bezeichnende Form liegt von den Localitdten 12, 14 und 43 in mehreren Exemplaren vor und ist durch die hauptsdchlichgegen den Unterrand bldttrig aufgestellten Querschuppen der Rippen ausgezeichnet. Translation. The impressive series of shells induces me to consider the Tridacna compressa depicted in Reeve’s monograph as a young form of T. elongata Eamarck, and to cancel this name, and to highlight a variety that is sys¬ tematically close to Tr. squamosa Eamarck. This form should be designated as squamosina nov. var. and several specimens are present from the localities 12, 14 and 43 and it is distinguished by the foliated transverse scales of the ribs mainly towards the lower margin. Comments. Sturany introduced the name squamosina for a variety of Tridacna elongata Eamarck, 1819; it has a subspecific rank according to the Art. 45.6.4 of the ICZN. This species had been poorly known for a long time but was recognised as a distinct entity on the basis of a mo¬ lecular phylogeny with the mitochondrial gene 16s rRNA (Richter et al. 2008). However, Richter et al. (2008) de¬ scribed this entity with the new name Tridacna costata which was recognised as a junior synonym of T. squamo¬ sina by Huber and Eschner (2011). Family Galeommatidae Gray, 1840 Scintilla sulphurea Stnrany, 1899 Figure 6 Sturany 1899: 286, plate VII, figures 6-9. Type locality. Eocality 25, near Abu Ghusun, Egypt, 26-24°N. Type material. Holotype: NHMW 38098, fixed by monotypy. Original description. Von der Localitdt 25; ein einziges Exemplar. Die Muschel ist elliptisch gestaltet, massig gewolbt, an den Randern ganz schliessend, ziemlich dickschalig, durch- scheinend, dicht und ziemlich stark concentrisch gestreift, aussen und innen schwefelgelb gefdrbt und glanzend. Der Wirbel uberragt den Schlossrand nur wenig und en- digt mit einem winzigen, glashell glitzernden Bldschen vor der Mitte der Schale am Schlossrande. Die Schlossleiste trdgt, entsprechend dem Gattungscharakter, ausser einem nahe herangeriickten Nebenzahn noch einen krdftigen Hauptzahn in der rechten Schale und 2 Hauptzdhne in der linken Schale, von denen der vordere starker als der hintere ist. Das braungefdrbte Ligament liegt in dem grubenformi- gen Zwischenraume zwischen Haupt- und Nebenzahn (fig. 7), welchen es jedoch nicht ganz ausfullt, und ist bei der geschlossenen Muschel von aussen nur wenig sichtbar. Der vordere Oberrandfdllt sanft nach vorne und bildet mit dem nahezu senkrecht gestellten, also ziemlich gerade (sogar wenig nach ruckwdrts) abgestutzten Vorderrand fast einen Winkel (eine sogenannte »runde Ecke«). Eben- so gestaltet ist der Ubergang von Vorder- in Unterrand; der letztere verldufi nicht ganz gerade, sondern zeigt eine zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 557-598 567 Figure 5. Tridacna elongata var. squamosina Sturany, 1899. A-F. Lectotype NHMW 107075; right valve exterior (A) and interior (B), left valve exterior (C) and interior (D), back view (E), front view (F). Scale bar: 2 cm. geringe Concavitdt und verhindet sich in rundem Bogen mit dem convexen Hinterrande, der andererseits auch mit dem etwas aufwdrts ziehenden hinteren Oberrande im Bogen verbunden ist. Lange der Muschel 9,0, Hohe 6,3, Dicke 4,7 mm. Es wollte mir nicht gelingen, die hier beschriebene Form mit einer der zahlreichen bestehenden Arten zu identificiren, von denen als die ndchst verwandten genanntseien: Sc. tQmxis Desk., semiclausaS'ow., oblonga Sow., pisumS'ow. hydrophana Translation. From locality 25; one single specimen. The clam is elliptical in shape, massively arched, completely closing at the edges, rather thick, translucent, densely and rather strongly concentrically Urate, outside and inside sulphurous in colour and shiny. zse.pensoft.net 568 Albano, RG. et al.: Types of Sturany’s Red Sea bivalves Figure 6. Scintilla sulphurea Sturany, 1899, near Abu Ghusun, Egypt. A-D, F, G, I, J, 1^0 Holotype NHMW 38098: left valve exterior (A, B) interior (F, G), and hinge detail (C, D); right valve exterior (I, J), interior (L, M), and hinge detail (N, O). E, H, K, P Original figures. Q Original label. Scale bars: 0.5 mm (C, D, N, O); 2 mm (A, I). zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 557-598 569 The umbo slightly protrudes over the hinge edge and ends with a tiny glassy glittering blister just before the centre of the shell at the hinge edge. The hinge margin, according to its generic character, has, in addition to a closely adjacent lateral tooth, a strong cardinal tooth in the right valve and two cardinal teeth in the left valve, of which the anterior is stronger than the posterior. The brown-coloured ligament lies in the pit-like spaces be¬ tween the cardinal and lateral tooth (Fig. 7), which it does not completely fill, and is only slightly visible from the outside in the closed shell. The anterior upper margin gently declines and almost forms an angle (a so-called “round corner”) with the al¬ most vertical (even slightly backwards facing) truncat¬ ed front edge. Equally designed is the transition from anterior to lower margin; the latter does not run quite straight but is slightly concave and connects in a circu¬ lar arc with the convex posterior margin, which on the other hand is also connected with the slightly ascending posterior margin. Length of the shell 9.0, height 6.3, thickness 4.7 mm. I did not succeed in identifying the form described here with one of the numerous existing species, of which the most related are: Sc. tenuis Deshayes, semiclausa Sowerby, oblonga Sowerby, pi sum Sowerby and hydro- phana Deshayes. Comments. This taxon is currently considered a junior synonym of Scintillula lutea (Lamarck, 1818) (Huber 2015). Scintilla variabilis Sturany, 1899 Figure 7 Sturany 1899: 287, plate VII, figures 1-5. Original localities. Locality 27, Berenice Troglodytica, Egypt, 24-22°N; locality 30, Halayeb, Egypt, 24-22°N; locality 41, Massawa, Eritrea, 16-11°N. Type material. Syntypes: NHMW 38099: 1 specimen, locality 27; NHMW 37416: 1 specimen (in alcohol), lo¬ cality 41; NHMW 37417: 2 specimens (in alcohol), lo¬ cality 30. Original description. Von den Localitdten 27, 30 und 41. Die Muschel ist von elliptischer Gestalt, massig gewolbt, dickschalig, an den Randern sich vollstandig schliessend, dicht und ziemlich stark concentrisch gestreift, schwach durchscheinend, innen und aussen milchweiss und glanzend. Der Wirbel uberragt den Schlossrand nur mit seinem bldschenfdrmigen Ende und steht in der vorderen Hdlfte der Schale. Das Schloss besteht in der rechten Schale aus einem krdftigen Hauptzahn 1 und einem nahe herangeriickten Seitenzahn; in der linken Schale aus 2 schwdcheren, un- gleich starken Hauptzdhnen und einem Nebenzahne. Das dunkelgefarbte Ligament ist von aussen schwach zu se- hen, ist auch hauptsdchlich erst unter dem Schlossrande, wo fur seine Aufnahme ein Ausschnitt der Schlossleiste zwischen Haupt- und Seitenzahn besteht, starker, und zwar etwa kugelig entwickelt (figs 2 und 4). Bei dlteren Exemplaren ist der Umriss der Muschel fast der einer Ellipse; nirgend sind sogenannte »Ecken« gebildet, sondern alle Ubergdnge (von Ober- und Un¬ ter- in Vorder- und Hinterrand) sind abgerundet. Bei jungeren Schalen jedoch grenzen sich die verschiedenen Bander etwas schdrfer von einander ah und ist hier und dort eine »stumpfe Ecke« oder ein Winkel gebildet. Auch ist hier zu bemerken, dass Ober- und Unterrand nicht streng parallel zu einander verlaufen mils sen, sondern dass sich die hochste Stelle der Muschel in der Regel riickwarts befindet, indem die Muschel vorne etwas nie- driger gebaut ist. [Tabelle mit Mafiangaben!] Leider habe ich mich veranlasst gesehen, der stattli- chen Artenreihe der Gattung Scintilla einen neuen Na- men hinzuzufugen, da sich die vorliegende, in verschiede¬ nen Altersstufen verschieden aussehende Form mit keiner der zahlreichen bisher bekannt gewordenen Scintillen mit Sicherheit identificiren Idsst. Als die ndchsten Verwand- ten mochte ich u. A. Sc. cumingii Desk von Panama und Sc. Candida Desk von den Philippinen bezeichnen. Translation. From localities 27, 30 and 41. The clam is elliptical in outline, massively arched, thick, completely closing at the edges, dense and with rather strong concentric ridges, slightly translucent, milk- white and shiny inside and outside. The umbo protrudes the hinge edge only with its vesicular end and stands in the anterior half of the shell. In the right valve, the hinge consists of a strong central tooth and a close lateral tooth; in the left valve, of two weaker, unequally strong main teeth and one lateral tooth. The dark-coloured ligament is hard to see from outside, it is mainly under the hinge margin, where there is a nearly spherical area between the central and posterior teeth for its inclusion (figs 2 and 4). In older specimens, the outline of the shell is almost elliptical; the so-called “corners” are not formed, but all margins (from dorsal and ventral to anterior and posteri¬ or) are rounded. In younger shells, the different margin edges are a little sharper, and here and there an “obtuse comer” or an angle is formed. It should also be noted that the dorsal and ventral margin do not have to be strictly parallel, so that the highest point of the shell is usually posterior and positioned slightly lower in the front. [Table with dimensions] Unfortunately, it seemed necessary to add a new name to the impressive series of species in the genus Scintilla, since the present form, which looks different at different ages, cannot be identified with any of the numerous es¬ tablished Scintilla. As closest relatives, I would like to name S. cumingii Deshayes from Panama and S. Candida Deshayes of the Philippines. zse.pensoft.net 570 Albano, RG. et al.: Types of Sturany’s Red Sea bivalves Figure 7. Scintilla variabilis Sturany, 1899. A-D, F, G, I, J, L-O Syntype NHMW 38099; left valve exterior (A, B), interior (F-G) and hinge detail (C, D). Right valve exterior (I, J), interior (L, M) and hinge detail (N, O). E, H, K, P Original figures. Q Original label. Scale bars: 0.5 mm (C, D, N, O); 2 mm (A, I). zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 557-598 571 Family Anatinellidae Deshayes, 1853 Raeta bracheon Sturany, 1899 Figure 8 Sturany 1899: 266, plate III, figures 1-6. Type locality. Station 87, Ra’s Mal’ab in the Gulf of Suez, Egypt, 29°7.6'N, 32°56'E, -50 m. Type material. Syntypes: NHMW 84335: 2 valves, sta¬ tion 87 (in original figure). Original description. Von Station 87 (50 m), eine rechte und eine linke Schale, die jedoch nicht zueinander gehoren. Die Muschel ist gross, Cm^idmm-fdrmig, mdssig gewolbt, dunn, durchscheinend, aussen milchweiss, matt, concentrisch gefaltet, innen glanzend. Der Wirbel liegt ein wenig vor der Mitte der Schale und uberragt den Schlossrand nicht besonders stark. Die vordere Hdlfte der Schale ist gewolbt, die hintere abge- flacht und schnabelformig ausgezogen. Der vordere Oberrand geht im Bogen in den gewolbten Vorderrand und dieser ebenso in den convexen Unterrand uber Der hintere Oberrand fdllt schief ab zum abgerundeten Hinterende des Schnabels, die untere Begrenzung des Schnabels ist ebenfalls von einer schiefen Linie gebildet; der Winkel des Rostrums ist circa 60°. Zwischen den concentrisch angeordneten Falten der Oberfldche, welche nach innen vollstandig durchgepragt sind, liegen noch mikroskopisch feine concentrische Streifen (in der Regel 5-6 Streifen zwischen 2 Falten). Gegen den Unterrand zu werden die Zwischenrdume der Faltung enger; die Falten selbst sind, entsprechend der Form der Schale, in ihrem Verlaufe mehrfach geknickt, besonders am Oberrande. Von Muskeleindriicken sind im Inneren der Schale zu sehen: ein langgestreckter, fast senkrecht stehender, nur wenig gekrummter vorne ndchst dem Vorderrande und ein etwa kreisfbrmiger riickwarts am hinteren Ober¬ rande, wo das Rostrum entspringt. Von oben betrachtet, Idsst die Muschel ein undeutlich begrenztes, schmales und Idngliches Feld vor dem Wirbel erkennen (lunula). Das Schloss besitzt ein inneres Ligament, welches in einer Idnglichen, etwa dreieckigen Grube liegt; unmittelbar davor stehen in der rechten Schale zwei senkrecht gestellte Mittelzdhne parallel zu einander, uber demselben, also am Schlossrande undgewissermassen als obere Begrenzung der Ligamentgrube, liegt ein ziemlich starker Zahn von gleicher Lange wie die Ligamentgrube; ferner sind leistenfbrmige Seitenzdhne, vorne und riickwarts je einer, zu constatiren; dieselben sind vom Oberrande durch Vertiefungen getrennt. In der linken Schale ist nur ein senkrechter Mittelzahn wahrnehmbar, welcher vor der Ligamentgrube steht; im ubrigen liegen hier die Schlossverhdltnisse wie in der rechten Schale. Die vorliegende rechte Schale ist 30,5 mm lang und 20 mm hoch, die linke Schale 29 mm lang und 19,5 mm hock Die neue, anscheinend nur geringe Tiefen des Roth- en Meeres bewohnende Raeta-ylrr ist nun die erste fur das eigentliche erythrdische Seebecken bisher bekannt gewordene aus dieser Gattung. In Aden kommt nach Shopland R. abercrombiei Melvill vor, deren Origi- nalfundort Bombay ist, und mit der meine Art nicht zu verwechseln ist. Translation. From station 87 (50 m), one right and one left valve, which do not belong to the same individual. The clam is large, Cuspidaria-shwpQd, moderately arched, thin, translucent, outside milky white, dull, con¬ centrically folded, shiny inside. The umbo is slightly anterior to the centre of the shell and does not much protrude the hinge edge. The anterior part of the valve is arched, the posterior on the same plane and subrostrate. The anterior upper margin shades off in a curve to the arched anterior margin and continues into the convex lower margin. The posterior upper margin slopes off to the rounded rear end of the beak, while the lower marking of the beak is formed by an oblique line; the angle of the rostrum is about 60°. Between the concentrically arranged folds, which are completely imprinted on the inside, microscopically fine concentric striae are present (usually 5-6 striae between two folds). Towards the lower margin, the spaces be¬ tween the folds become narrower; the folds themselves, according to the shape of the shell, are repeatedly flexed, especially at the upper margin. Muscle scars can be seen in the interior of the shell: one elongated, almost vertical, slightly curved near the anterior margin and another almost circular at the posteri¬ or margin, where the rostrum originates. Viewed from above, the clam shows an indistinct, nar¬ row and elongated area in front of the umbo (lunula). The hinge has an inner ligament, which lies in an elongated, approximately triangular pit; directly in front of it, in the right shell, two perpendicular medi¬ an teeth are parallel to each other. At the edge of the hinge, and as a kind of upper restriction of the ligament pit, a fairly strong tooth lies and is of the same length as the ligament pit; in addition, bar-like lateral teeth, one in front and one on the back are separated from the upper margin by depressions. In the left valve, only one vertical median tooth is perceptible, in front of the ligament pit; otherwise, hinge conditions are the same as in the right shell. The present right valve is 30.5 mm long and 20 mm high, the left valve 29 mm long and 19.5 mm high. This new species is the first for the Red Sea basin known from this genus inhabiting apparently only shal¬ low waters. According to Shopland, R. abercrombiei Melvill occurs in Aden, its original location is Bombay and should not be confused with my species. zse.pensoft.net 572 Albano, RG. et al.: Types of Sturany’s Red Sea bivalves Figure 8. Raeta bracheon Sturany, 1899, Station 87, Ra’s Mal’ab in the Gulf of Suez, Egypt, 50 m. A, B, J-M Original figures. C-F, H, I Figured syntypes NHMW 84335: right valve exterior (C) interior (E), and hinge detail (H); left valve exterior (D), interior (F), and hinge detail (I). G Original label. Scale bars: 5 mm (C, D); 2 mm (FI, I). zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 557-598 573 Family Tellinidae Blainville, 1814 Tellina siebenrocki Sturany, 1899 Figure 9 Sturany 1899: 278, plate VI, figures 4-7. Type locality. Locality 45, Ras Mujamila, Yemen, 16-11 °N. Type material. Holotype: NHMW 38016, fixed by monotypy. Original description. Von der Localitdt 45; ein einziges, aber schon erhaltenes Exemplar. Die Muschel ist dickschalig, ungleichseitig und fast gleichschalig, wenig gewolbt, eiformig mit schnabelig vortretendem Hinterende; aussen stark glanzend und et- was opalisirend, rosafarbig im Grundtone und purpur- roth gefdrbt in der Wirbelgegend, innen glanzend und mehr minder orangegelb, in der Wirbelgegend schwach durchscheinend. Sie ist concentrisch gestreift, und zwar ziemlich dicht und unregelmdssig (etwas grober am Un- terrande) und ist durch Spuren von radialer Streifung aussen sowohl wie innen ausgezeichnet. Die Wirbel sind mittelstandig und stehen einander am Schlossrande dicht gegenuber. Vor dem Wirbel fdllt der Rand der Schale in schwachem, etwas herausgekriim- mten Bogen schief herab; er verbindet sich bogig mit Vorder- und Unterrand; hinter dem Wirbel senkt sich der Rand schief und etwas convex herab zu dem kurzen, und abgeschlossenen Rostrum, das unten, am Ubergange in den schwach convexen Unterrand, eine schwache Con- cavitdt aufweist. Das Schloss der rechten Schale besitzt direct unter dem Wirbel zwei divergirende, freistehende Cardinalzahne, wovon derhintere gespalten ist, einen nahe herangeriickten vorderen und einen Idngeren, ebenfalls nicht weit entfernt stehenden Lateralzahn; iiber diese Lateralzdhne ist der Rand leistenfbrmig hervorgezogen. In der linken Schale ist ein kaum gespaltener Cardinalzahn zu verzeichnen, eingefasst von Gruben fur die gegenuberstehenden Zdhne der rechten Schale. Die Seitenzdhne werden hier vertreten durch zahnartig vorgezogene Rander vorne und riickwarts, welche in die entsprechenden Vertiefungen zwischen den Lateralzdhnen und Randern der rechten Schale passen. Das braune Ligament der Muschel liegt aussen hinter dem Wirbel. Vom Wirbel zieht in jeder Schale eine schwache kielfbr- mige Erhebung zum schnabelfdrmigen Ende der Schale, wodurch also riickwarts eine lanzettfbrmige Area entsteht. Die Muskeleindriicke sind deutlich; der vordere ist auf- rechtstehend oval, der hintere rund. Die Mantelbucht reicht bis zum vorderen Muskeleindruck, steigt unter dem Wirbel ziemlich hoch hinauf und endigt vorne ziemlich spitz. Lange der Muschel 15,7, Hohe 11,3 und Dicke 6,2 mm. Die neueArt hat die Gestalt einer T. producta Sow. ein- er T. culter Hanl. (d. i. eine Form von den Philippinen, die auch eine dhnliche Farbe besitzt), einer T. cuspis Hanl. einer T. brevicostata Sow. etc.; die beiden letzteren sind. abgesehen von anderen viel wichtigeren Unterschieden, auch viel grosser in ihren Umrissen. Translation. From locality 45; a single, but well-pre¬ served specimen. The clam is thick, inequilateral and is almost equiv- alve, little arched, egg-shaped with a protruding beaked posterior end; outside strongly glossy and somewhat opalescent, with a pinkish background and purple colour¬ ed in the umbonal region; on the inside, glossy and more or less orange-yellow, slightly translucent in the umbonal region. It has concentric striae, rather dense and irregular (somewhat coarser at the lower margin) and is marked by traces of radial striation both externally and internally. The beaks are centred and stand close to each other at the hinge edge. In front of the beak, the valve margin is sloping down in a faint, slightly arched curve; and curved again when connecting the anterior and ventral margins; behind the beak, the edge slopes convexly down to the short and closed rostrum, which is weakly concave below and then merges into the weakly convex lower margin. The hinge of the right valve has directly below the umbo, two diverging free-standing cardinal teeth, of which the posterior is split, closely adjacent is an anterior lateral tooth; over these lateral teeth the edge is extended and ridge¬ shaped. The left shell has a weakly split cardinal tooth, sur¬ rounded by pits for the opposing teeth of the right shell. The lateral teeth are represented by anterior and posterior tooth- like margins which fit into the corresponding pits between the lateral teeth and the edges of the right valve. The brown ligament of the clam lies externally behind the umbo. From the umbo, a weak keel-like elevation in each valve runs to the beak-shaped end of the valve, thus cre¬ ating a lancet-shaped area backwards. The muscle scars are light; the anterior is upright oval, the posterior round. The pallial sinus extends to the ante¬ rior muscle scar, rises quite high under the umbo and ends quite pointedly. Length of the shell 15.7, height 11.3 and thickness 6.2 mm. The new species has the shape of T. producta Sowerby, T. culter Hanley (a form of the Philippines, which has a similar colour), T. cuspis Hanley, T. brevicostata Sowerby, etc.; the latter two, apart from other much more important differences, are also much broader in their outlines. Comments. It is considered a junior synonym of Tellina bertini (Jousseaume, 1895) (Oliver 1992). Family Solecurtidae d’Orbigny, 1846 Solecurtus subcandidus Sturany, 1899 Figure 10 Sturany 1899: 260-261, plate I, figures 1-4. Type locality. Station 94, Nuweiba, Gulf of Aqaba, Egypt, 28°58.6'N, 34°43.7'E, -314 m. zse.pensoft.net 574 Albano, RG. et al.: Types of Sturany’s Red Sea bivalves iiiifil Figure 9. Tellina siebenrocki Sturany, 1899, locality 45, Ras Mujamila, Yemen. A, B, D, F, H, I Holotype NHMW 38016; right valve exterior (A), interior (B), and hinge detail (D); left valve exterior (F), interior (H), and hinge detail (I). C, E, G, J Original figures. K Original label. Scale bars; 1 mm (D, I); 2 mm (A, F). I o « V Vi : zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 557-598 575 Type material. Holotype: NHMW 84296, fixed by monotypy. Original description. Ein einziges Exemplar von Station 94 (314 m). Die Muschel klafft an beiden Enden, ist wenig ge- wdlbt, langgestreckt oval, ziemlich festschalig, schwach durchscheinend und ein wenig glanzend, aussen weiss mit Spuren gelber Zeichnung, innen rein weiss. Die Sculptur derselben ist im Allgemeinen fein und un- deutlich. A us der dichten concentrischen Streifung treten gegen die Rander der Schalen zu meh- rere Anwachsstreifen krdftig hervor; eine Radiarstreifung fehlt nur in der vorderen Schalenpartie, in der Mitte treten sehr schief gestellte Streifen auf, in der hinteren Schalen¬ partie stehen aufrechte Streifen, die jedoch ein- bis zwei- mal gekrummt sind, und zwar oben mit dem Bogen nach vorne, unten mit dem Bogen nach hinten (vide fig. 1). Der Oberrand der Muschel weicht wenig von einer Geraden ab, vorne ist er schwach abfallend, hinter dem Wirbel minimal eingebogen; der Unterrand ist ganz ge- rade; Vorder- und Hinterrand gehen oben und unten mit »runden Ecken« in Ober- und Unterrand iiber. Der Wirbel steht vor der Mitte, ist schwach zugespitzt und uberragt den Schlossrand wenig. Aus der Schlossleiste ragen in der rechten Schale 2 spatenfbrmige Zdhne hervor, von denen der hintere be- deutend krdftiger entwickelt und longer ist; in der linken Schale befindet sich nur 1 schwdcherer Hauptzahn, der vorne und riickwarts von einer Grube umstellt wird. Hin¬ ter den genannten Zdhnen liegt auf vorgezogenem Rande das Ligament. Die Mantelbucht ist zungenformig und reicht bis iiber die Wirbelregion hinaus in die vordere Schalenpartie. Die Lange der Muschel betrdgt 30, die Breite 12,4, die Dicke circa 7 mm. Die nachstverwandten Arten sind S. divaricatus Lischke aus Japan und S. Candidas Renier aus dem Mittelmeer und demAtlantischen Ocean. Die erstgenannteArt unterscheidet sich hauptsdchlich dadurch, dass die Querlinien vorne nicht so schief gestellt sind wie bei der neuaufgestellten Form, und dass sie riickwarts, respective oben runzelig werden, auch anders geknickt erscheinen. Bei S. candidus Renier ist die Quer- (oder Radial-) streifung dhnlich ausgebildet wie bei S. subcandidus m., doch ist jene Muschel gewolbter und relativ hoher. Beiden in Vergleich gezogenen Arten gegeniiber ist die neue Art iiberdies durch die besonders stark entwickelte Bezahnung ausgezeichnet. Translation. One single specimen from station 94 (314 m). The clam gapes at both ends, is slightly arched, elon¬ gated oval, quite hard-shelled, weakly transparent and a little shiny, white with slight yellowish shading on the outside, pure white on the inside. The sculpture is generally fine and indistinct. From the dense concentric striation, a number of growth lines emerge most prominently at the margins of the valves; a radial striation is missing only over the anterior part of the shell, in the middle part very oblique incisions are pres¬ ent, over the posterior part there are upright ridges, which are once or twice sinuous, in fact they have the concavity to the front dorsally, and to the back ventrally (see fig. 1). The upper margin of the clam deviates little from a straight line; it is slightly sloping, minimally sloping be¬ hind the umbo; the lower margin is very straight; above and below, the anterior and posterior margin change with “rounded corners” into the upper and lower margins. The weakly pointed umbo is located anteriorly and pro¬ trudes little beyond the hinge edge. Two fiat teeth protrude from the hinge edge in the right valve, the most posterior of which is much more prominent and long; in the left valve there is only one weaker central tooth, which is en¬ closed by a cavity in the front and back. The ligament lies behind the aforementioned teeth on an elongated edge. The pallial sinus is tongue-shaped and reaches beyond the umbo region into the anterior part of the shell. The length of the shell is 30 mm, height 12.4, thick¬ ness 7 mm. The closest related species is S. divaricatus Lischke from Japan and S. candidus Renier from the Mediterrane¬ an Sea and the Atlantic Ocean. The first species is mainly distinguished by less tilted transverse lines compared to the newly described form, and by a rugose structure on the back, with a differently tilted appearance. In S. candi¬ dus Renier, the lateral (or radial) striation is developed in a similar way as in S. subcandidus m., however, the shell is more rounded and relatively higher. Furthermore, the new species is distinguished by the strongly developed dentition compared to both species. Family Ungulinidae Gray, 1854 Diplodonta raveyensis Sturany, 1899 Figure 11 Sturany 1899: 285-286, plate VI, figures 8-11. Type locality. Locality 32, Gul Mohammad, Saudi Ara¬ bia, 22-20°N. Type material. Holotype: NHMW 38097 lost; see comments. Original description. Von der Localitdt 32; ein tadello- ses Exemplar. Die Muschel ist fast kreisfbrmig im Durchschnitte, ziemlich festschalig, stark gewolbt; aussen etwas gldn- zend, mit feiner, dicht stehender, concentrischer Punkt- streifung ausgestattet, in der Farbe schmutzigweiss bis gelblich mit einigen hellgrauen, nach innen durchschim- mernden Zonen; innen reinweiss, glatt und glanzend am Rande, rauh und matt in der Mitte. Die Wirbel sind stark ausgehohlt, stehen vor der Mitte, sind mit ihren stumpfen Spitzen nach innen und vorne gekehrt und stehen sich an dem Schlossrande gegeniiber, den sie nicht viel iiberragen. Eine Lunula ist kaum ausgebildet. zse.pensoft.net 576 Albano, RG. et al.: Types of Sturany’s Red Sea bivalves Figure 10. Solecurtus suhcandidus Sturany, 1899, Station 94, Nuweiba, Gulf of Aqaba, Egypt, -314 m. A, C, G, I Original figures. B, D-F, FI, J Holotype NHMW 84296: left valve exterior (B), interior (D), and hinge detail (J), right valve exterior (E), interior (F), and hinge detail (H). K Original label. Scale bars: 5 mm (B, E); 2 mm (H, J). Der vordere Oberrand fdllt schief ab und geht im Bo- gen in den Vorderrand uber; ebenso ist der Ubergang von Vorder- in Unterrand und von Unter- in Hinterrand schon gerundet, nur der vom Wirbel schief abfallende hintere Oberrand bildet an seinem Ubergange in den Hinterrand einen schwach ausgeprdgten Winkel, der nicht viel mehr als 100-110°betrdgt. Das Schloss besteht aus einem inneren Ligament di¬ rect unter dem Rande und aus einer aujfallenden Bezah- nung. Die letztere besteht in der rechten Schale in 2 di- vergirenden Mittelzdhnen unter dem Wirbel, von denen der hintere gegabelt ist und die voneinander durch eine dreieckige Grube getrennt sind. In der linken Schale sind ebenfalls zwei Mittelzdhne zu sehen, von denen aber der vordere gespalten ist und der hintere einfach bleibt. Auch hier sind dieselben voneinander durch eine dreieckige Grube in der breiten Schlossleiste getrennt, und hier wie dort liegt vor dem vorderen Mittelzahne eine schwache Vertiefung, die nach vorne rinnenformig verlduft, und hier wie dort liegt das Ligament gleich hinter dem hinteren Mittelzahn, schief vom Wirbel herab nach hinten ziehend. zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 557-598 577 Figure 11. Diplodonta raveyensis Sturany, 1899, locality 32, Gul Mohammad, Saudi Arabia. A, C, D, F, H Original figures. B, E, G, I Holotype NHMW 38097 left valve exterior (B) and interior (E); right valve exterior (G) and interior (I). Photo courtesy Henk Dekker. Scale bar: 3 mm. Der Mantelrand verlduft parallel dem unteren Rande der Schale und endigt vorne und riickwarts an den Mus- keleindrucken. Lange der Muschel 10,4, Hohe 9,8, Dicke 7,3 mm In der Gestalt ist diese neue Art wohl dhnlich der Lu- cina globularis Lam., welche nach Jousseaume auch im Rothen Meere vorkommen soil; ferner der nunmehr in die GattungT>\^\odtmtdi verwiesenen Lucina rotundata Turton, fur welche die Bezahnung ganz so beschrieben wird, wie wir sie bei der neuenArt gesehen haben und die von Reeve fur »Mediterranean and southern shores of Britain«, von Smith und Shopland fur Aden, von Caramagna sogar fur Assab im sudlichsten Theile des Rothen Meeres angegeben wird. Von dieser L. rotundata Turton ist aber meine Art schon durch die geringe Grosse geniigend verschieden. Mysis tumida A. Ad., in der Gestalt der neuenArt eben- falls dhnlich, ist nur wenig grosser, jedoch durch das Merk- mal »striolis confertis radiantibus et concentricis obsolete decussata« hinreichend als verschieden gekennzeichnet. Translation. From locality 32, one perfect specimen. The clam is nearly circular, quite hard-shelled, strongly arched; slightly shiny on the outside, with fine, compact, concentric dotted striation, of dirty white to yellow colour with some light grey, translucent to the inside zones; pure white on the inside, smooth and shiny on the edge, rough and matt in the middle. The beaks are strongly extended, positioned anteriorly, tilted to the inside and to the front with obtuse tips and touch each other at the hinge edge, which they do not protrude by much. The lunula is hardly developed. The anterior upper margin goes down and merges with the frontal margin in a curve; also, the transition from the anterior to the lower margin and from the lower to zse.pensoft.net 578 Albano, RG. et al.: Types of Sturany’s Red Sea bivalves the posterior margin is nicely rounded; only the uneven¬ ly sloping posterior upper margin forms at its junction a weakly developed angle of no more than 100-110°. The hinge is formed by an inner ligament directly be¬ neath the margin and by a prominent dentition. The latter consists of two divergent middle teeth be¬ low the umbo, which are separated from each other by a triangular cavity and the posterior one is bifurcate. In the left shell, there are also two middle teeth; however, the anterior one is bifurcate and the posterior is ordinary. Also in this case, they are separated by a triangular cavity within the broad hinge margin and in both cases there is a shallow deepening in front of the central tooth, which proceeds towards the front in a groove, and in both cases the ligament lies right behind the posterior central tooth. The pallial line proceeds parallel to the lower margin of the shell and ends in the front and posteriorly at the muscle scars. Length of the shell 10.4, height 9.8, thickness 7.3 mm. In its shape, this new species apparently resembles Lucina globularis Lamarck which according to Jous- saume occurs also in the Red Sea. Furthermore, there is a similarity to Lucina rotundata Turton, now placed in the genus Diplodonta, for which the dentition is described just as we have seen it in the new species and which was named by Reeve for “Mediterranean and southern shores of Britain”, by Smith and Shopland for Aden, and by Caramagna even for Assab in the southernmost part of the Red Sea. My species, however, is different enough because of the small size from this L. rotundata Turton. Mysis tumida A. Adams, also similar in shape to the new species, is only slightly smaller, but adequately differ¬ entiated by the characterization '"striolis confertis radian- tibus et concentricis obsolete decussatd" [reticulate sculp¬ ture of compact radial and obsolete concentric striae]. Comments. The holotype was lost while on loan. Family Veneridae Rafinesque, 1815 Chione hypopta Sturany, 1899 Figure 12 Sturany 1899: 281-282, plate VII, figures 10-14. Original localities. Locality 10, Nuweiba, Egypt, Guff of Aqaba; locality 16, Jazirat Shakir, Egypt, northern Red Sea, 28-26°N. Type material. Syntypes: NHMW 38049: locality 16, 5 valves (one specimen in original figure). Original description. Von den Localitdten 10 und 16. Die Muschel ist oval bis dreieckig, dickschalig, wenig gewolbt, aussen weiss bis gelb mit unregelmdssig in grosseren oder kleineren braunen Flecken vertheilter Zeichnung, innen violett oder weiss. Die Schale ist an Hirer Oberfldche radial und der Lan¬ ge nach von Furchen durchzogen, die tief einschneiden und eine bemerkenswerthe Felderung hervorrufen. So stehen mehr als dreissig derbe Radialrippen dicht anein- ander, die am Wirbel schwach entspringen und gegen den Rand zu stark werden, und welche eben durch die Quer- furchen eine Gitterung erhalten. In der hinteren Schalen- partie sind die Felder schuppig oder dornig ausgebildet, dock ist dies nur bei jungen Exemplaren gut zu sehen. Die an der Spitze violett oder roth gefdrbten Wirbel stehen etwas vor der Mitte der Schale und uberragen den Schlos- srand nur wenig. Der vordere Oberrand fdllt vom Wirbel schief und etwas bogig herab in den gerundeten Vorderrand, welcher auch mit dem Unterrand bogig verbunden ist. Der hintere Oberrand verlduft etwas schief nach riick- wdrts und hinab zum Hinterrand, mit dem er unter ei- nem kaum merklichen, stumpfen Winkel sich verbindet, wahrend wieder Hinter- und Unterrand an dem im Alter etwas ausgezogenen Hinterende der Muschel bogig ver¬ bunden sind. Eine Kerbung der Rander, entsprechend den dusseren Endigungen der Radialrippen, ist nur bei jun¬ gen Exemplaren aujfallend entwickelt; bei diesen ist dann innerhalb der Kerbung auch jene alien Chionen zukom- mende Strichelung besonders gut zu sehen, die bereits an den Oberrandern beginnt und ringsum zieht. Vor den Wirbeln liegt eine deutlich begrenzte, lanzett- fbrmige Lunula, hinter derselben das dussere Ligament. Die Schlossleiste trdgt im Allgemeinen 2 divergirende Zdhne und 3 Gruben in der rechten Schale sowohl wie in der linken. Bei jungen Exemplaren ist des Ndheren zu se¬ hen, dass die Grube vor dem vorderen Zahn der rechten Schale noch von einem schwachen Zdhnchen uberstellt ist, femer dass der hintere Hauptzahn der linken Schale vorne etwas gespalten ist und darauf noch ein schwacher, leis- tenfdrmiger Zahn folgt, der schief nach ruckwdrts Iduft. Der Mantelrand ist ruckwdrts kurz zungenfdrmig ein- gebuchtet. [Tabelle mit Majlangabenl] Translation. From locality 10 and 16. The clam is oval to triangular, thick-shelled, poorly arched, white to yellow on the outside with a pattern of ir¬ regularly sized, brown spots, purple or white on the inside. The shell is radially and longitudinally carved by deep grooves on its surface, causing a notable sculpture. Over thirty compact radial ribs are densely arranged, they are initially weak at the umbo and become stronger towards the margin and become cancellate when crossing the concentric grooves. In the posterior part of the shell, the sculpture is scaly or thorny; however, this is clearly visi¬ ble only in young specimens. The umbos are violet or red at their tips, they are po¬ sitioned anteriorly and protrude the hinge margin only slightly. The anterior dorsal margin declines from the umbo with a slight curve into the rounded anterior mar¬ gin, which is also connected to the ventral margin. The posterior dorsal margin slopes slightly backwards and downwards to the posterior margin, with which it zse.pensoft.net Zoosyst. Evol. 95 (2) 2019, 557-598 579 connects in a hardly noticeable obtuse angle, while poste¬ rior and upper margins are connected again in a curve at the posterior end of the bivalve, which is slightly extend¬ ed in older specimens. A crenulation of the margin, due to the endings of the radial ribs, is noticeably developed only in young specimens. Here, within the crenulations, the finer sculpture typical for all Chione is especially vis¬ ible, it already starts at the upper margins and continues all around the shell. A clearly confined, elongate lunula lies in front of the umbos, located behind them there is the external liga¬ ment. The hinge margin generally shows two diverging teeth and three pits in both the right and left valves. In young specimens, small teeth surround the pit in front of the central tooth of the right valve; furthermore, the pos¬ terior central tooth of the left valve is slightly split at the tip and is followed by a weak, elongated oblique tooth. The pallial sinus is slightly indented posteriorly in a tongue-shape. [Table with dimensions] Family Gastrochaenidae Gray, 1840 Gastrochaena deshayesi Sturany, 1899 Figure 13 Sturany 1899: 273-274, plate V, figures 1-7. Type locality. Locality 37, near Jazan, Saudi Arabia, 18-16°N. Type material. Syntypes: NHMW 37982: locality 37, 4 specimens (one specimen in original figure). Original description. Von der Localitdt 37 (Ras Turfa); einige wenige abgestorbene Exemplare. Die Muschelliegteingeschlossen in einem Kalkgehduse, das aus 6-7 aneinander gegliederten Ringen besteht. Der vorderste ist kopjfdrmig oder kugelig aufgeblasen und am grossten, nach hinten zu verschmdlern sich die Ringe, und der letzte, kleinste besitzt die Ojfnungfur denAustritt der Siphonen. Das Kalkgehduse sitzt meist einer fremden Molluskenschale auf oder ist mit dem Gehduse eines zweiten Individuums verklebt. Die Muschel ist Idnglich oder nahezu viereckig, stark gewolbt und gedreht, ziemlich dickschalig, ventral weit gedjfnet, so dass ein lang herzfbrmiger Hiatus entsteht, und hat ihre wenig eingedrehten Wirbel fast am vordersten Rande stehen, wo die Muschel am hochsten ist. Ober- und Unterrand sind mitunter parallel; erstere verlduft vom Wirbel nach hinten zuerst aufwdrts, dann ein wenig concav, letzterer ist stark nach aussen gebogen und verlduft uberdies etwas concav. Der Vorderrand fdllt nahezu senkrecht vom Wirbel herab, der Hinterrand ist ein convexer, aufrecht stehender Bogen. Die Schalen sind schmutzigweiss bis gelb und werden diagonal, d. i. vom Wirbel zum Unterrande, von einer Depression durchzogen, wodurch sie sich hier abflachen und einander ndhern. In der vorderen, stark gewdlbten Hdlfte der Schale, also vor der Depression, ist eine grobe Ldngsstreifung bemerkbar, in der hinteren und oberen Partie (hinter der Depression) treten aus der hier aufrecht stehenden Streifung in der Regel 5 mit dem Hinterrande gewissermassen concentrisch gestellte Wachsthumslinien auf. Dieselben sind auch im Inneren der Schale markirt und durften mit der Articulation der dusseren Kalkhulle mehr minder correspondiren, d. h. gleichzeitig mit der Anlage eines neuen Ringes aussen durfte innen ein Wachsthum der Schale stattftnden. Im Inneren der Schale liegt hinter der Mitte ein grosser, runder Muskeleindruck, vorne am Vorderrande liegen ein paar ganz kleine, undeutliche Eindrilcke unter einander. Die Bezahnung des Schlossrandes ist in der Regel gleich Null; nur ausnahmsweise tritt rechts ein zahnfbrmiger Stumpf auf, dem dann links eine kleine Grube entspricht. [Tabelle mit Mafiangaben!] Reeve bildet in seiner Monographie der Gattung Gastrochaena einige Formen ab, die mit der vorliegenden als neu beschriebenen Art zweifellos grosse Ahnlichkeit besitzen. Es ist dies vor Allen die Sowerby ’sche G. ovata von Panama. Wahrenddiefig. 16a bei Reeve zwei mit einander verklebte Kalkgehduse darstellt, ganz dhnlich einem mir vorliegenden