Zoosyst. Evol. 96 (1) 2020, 205-216 | DO! 10.3897/zse.96.50360 > PENSUFT. Gye BERLIN A new genus for Pericera septemspinosa Stimpson, 1871 and Pericera heptacantha Bell, 1836 (Crustacea, Brachyura, Majoidea), based on morphology and molecular data Jessica Colavite!*, Amanda M. Windsor’, William Santana! 1 Departamento de Zoologia, Instituto de Biociéncias, Universidade Estadual Paulista “Julio de Mesquita Filho” (UNESP), Botucatu, SP 18618-970, Brazil 2 Laboratory of Systematic Zoology, Centro Universitario Sagrado Coragdo (UNISAGRADO), Rua Irma Arminda, 10-50, Jd. Brazil, Bauru, SP 17011-160, Brazil 3 Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Museum Support Center, 4210 Silver Hill Road, Suitland, MD 20746-2863, USA http://zoobank. org/3 195954C-4A45-4EC4-A E40-20B1I58D75C5E Corresponding author: William Santana (willsantana@gmail.com) Academic editor: Sammy De Grave @ Received 22 January 2020 # Accepted 8 April 2020 Published 20 May 2020 Abstract A new genus of majoid spider crab, Pohleus gen. nov. is established for Pericera septemspinosa Stimpson, 1871 and Pericera hep- tacantha Bell, 1836, based on morphology and molecular data from the partial sequences of the 12S and 16S mitochondrial genes and the 18S small subunit rRNA nuclear locus. The species are re-described and illustrated, based on material from several localities of the western Atlantic and eastern Pacific oceans. The carapace, antennal and pterygostomial spines, male thoracic sternum and first gonopods are distinctive characters, distinguishing Pohleus gen. nov. from species assigned to Macrocoeloma Miers, 1879, where P. septemspinosus and P. heptacanthus are currently included. Key Words Epialtidae, Pisidae, western Atlantic, eastern Pacific, Macrocoeloma heptacanthum, Macrocoeloma septemspinosum Introduction Macrocoeloma heptacanthum (Bell, 1836) and M. septem- spinosum (Stimpson, 1871) were originally described as Pericera Latreille, 1825 and transferred by Miers (1886) to Macrocoeloma Miers, 1879 (see Colavite et al. 2016 for review). As part of an ongoing revision of Macrocoeloma, we identified the necessity to establish a new genus for the above two species. The description of Pericera heptacan- tha was based on two syntypes, one male and one female. This type material was treated as missing in literature for more than 50 years (see Rathbun 1925; Garth 1958) until the female specimen was re-discovered by Di Mauro (1982) in the dry crustacean collection in the University Museum, Oxford (OUM). Pericera septemspinosa, from the western Atlantic, was described by Stimpson (1871) as a geograph- ical analogue to Pericera heptacantha from the eastern Pa- cific and, unfortunately, the type material of P. septemspi- nosa was most likely destroyed in the Great Chicago Fire in 1871 (Evans 1967; Deiss and Manning 1981; Manning 1993: Vasile et al. 2005; Manning and Reed 2006). Pohleus gen. nov., is proposed herein to receive Peri- cera heptacantha and P. septemspinosa and a lectotype and a neotype are designated for each species, respec- tively. The species are re-described, illustrated and the morphological differences between them are detailed be- low. A phylogenetic framework for Poh/eus gen. nov. and allied genera is proposed, based on partial sequences of the 12S and 16S mitochondrial genes and the 18S small subunit rRNA nuclear locus. Copyright Jessica Colavite et al. This is an open access article 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 original author and source are credited. 206 Colavite, J et al.: A new genus for Pericera septemspinosa and Pericera heptacantha Material and methods Specimens examined are deposited in the collections of the Colecao de Invertebrados Aquaticos do Sul da Bahia, Universidade Estadual de Santa Cruz, Brazil (CIASB/ UESC); Grupo de Invetigaciones Carcinologicas, Uni- versidad de Oriente, Venezuela (GIC); Laboratorio de Sistematica Zooldgica, Universidade do Sagrado Coracao, Bauru (LSZ); Muséum National d’ Histoire Na- turelle, Paris (MNHN); Museum of Comparative Zool- ogy, Harvard University (MCZ); Museu de Zoologia da Universidade de Sao Paulo (MZUSP); Natural History Museum of Los Angeles County (NHMLA); Zoological collection of the Oxford University Museum of Natural History (OUM); Senckenberg Museum of Natural Histo- ry, Frankfurt (SMF); University of Louisiana at Lafayette Zoological Collection (ULLZ) and the National Museum of Natural History, Smithsonian Institution (USNM). For comparisons between Pohleus gen. nov. and Macrocoelo- ma s. str., 11 of the 12 known species of Macrocoeloma were examined, including the type species Macrocoelo- ma trispinosum Latreille, 1825 (see Comparative mate- rial section below). Additionally, the monotypic genus Thersandrus Rathbun, 1897 (type species Thersandrus compressus (Desbonne in Desbonne & Schramm, 1867) was included in our comparisons. Abbreviations used are: cl, carapace length (along the dorsal midline, from the base of the rostral sinus to the posterior margin of the carapace); cw, carapace width (taken at the widest point including lateral spines); P2— PS, pereopods 2 to 5 (P1 is the cheliped); G1, first gono- pod; G2, second gonopod; ovig., ovigerous; juv., Juve- nile; RV, research vessel; exped., expedition; stn, station; leg., collector or collected by; det., determined by. DNA extraction, PCR, and sequencing Total genomic DNA was extracted from muscle tissue using either the Qiagen DNeasy Blood and Tissue ex- traction kit or Omega Bio-tek EZNA Tissue DNA Kit. Partial sequences of the 12S, 16S mitochondrial genes were amplified with the following primers, respective- ly: 12SF (Mokady et al. 1994) and 12S1R (Shull et al. 2005), 16SF/16SR (Hultgren and Stachowicz 2008). The nuclear loci of the small subunit 18S rRNA were ampli- fied with the primers A and B (Medlin et al. 1988) and C, Y, O and L of Apakupakul et al. (1999) or D18s1R, D18s2F/R, D18s3F/R, D18s4F/R and D18s5F of Brack- en et al. (2009). Annealing temperatures for PCRs were 58 °C and 54 °C for 12S/18S and 16S, respectively. Re- agent volumes and concentrations followed manufactur- er’s instructions; primer concentrations were 10 uM. Se- quencing reactions were performed using 1 ul of purified PCR product in a 10 ul reaction containing 0.5 pl primer, 1.75 ul BigDye buffer and 0.5 ul BigDye (Life Technol- ogies). Sequencing reactions were purified using Milli- pore Sephadex plates (MAHVN-4550), according to the zse.pensoft.net manufacturer’s instructions and sequenced on the ABI 3730XL automated DNA sequencer. Sequences were assembled, trimmed of primers and checked for quality using Geneious 8.0.5 and 9.1.8. DNA extraction and se- quencing were carried out at the Smithsonian Institution’s Laboratories of Analytical Biology. Molecular data analysis Sequences, generated for this study, were combined with other sequences available from GenBank and previously unpublished sequences generated by AMW, in order to place the target taxa within the context of the superfamily Majoidea. Locality information and GenBank accession numbers for taxa included in the molecular analyses are provided in Suppl. material 1: Table S1. All individual se- quences for each species were initially analysed and as- sembled using the software Geneious 8.0.5 (http:/Awww. geneious.com; Kearse et al. 2012). The alignment amongst all sequences was generated with the algorithm Muscle (Edgar 2004) under default parameters and the perl script PartitionFinder (Lanfear et al. 2017) on CIPRES Portal (Miller et al. 2010) was run to determine the appropriate model of evolution and partitioning scheme. To determine whether the sequence data was appropriate for phyloge- netic studies, the occurrence of substitution saturation was evaluated in the molecular data and identical sequences were excluded, resulting in 23 unique sequences. Num- bers of transition and transversions versus ML-Compos- iteTN93 distance were plotted using the software DAM- BE 7.2.7 (Xia 2013, 2017), with the options of pairwise deletion and genetic distance model F84. The individual datasets were concatenated in RAXML, where we used the ‘-f ae’ option with 1000 bootstrap replicates. Likelihood parameters followed the General Time Reversible (GTR) model with a gamma distribution on the partitioned dataset and RAxML estimated all free parameters. The resulting best tree was used to reflect phylogeny (Fig. 1). To estab- lish the phylogenetic context for Pohleus septemspinosus gen. nov. et comb. nov., we followed the previous phylog- enies of Majoidea (Hultgren and Stachowicz 2008, 2009; Windsor and Felder 2014) to choose the genera used in this study and rooted the tree with the hymenosomatid crab Elamena producta Kirk, 1879. Only nodes with maximum likelihood bootstrap support greater than 50% are shown on the maximum likelihood phylogram (Fig. 1). All analy- ses were run on the Hydra computing cluster at USNM or CIPRES Portal (Miller et al. 2010). Nodes, where maxi- mum likelihood bootstrap support is greater than 50%, are shown on the maximum likelihood phylogram (Fig. 1). The ongoing revision and phylogenetic study of Mac- rocoeloma by the authors (unpubl. data), based on both morphological and molecular data, revealed this genus to be a paraphyletic group. Poh/eus gen. nov. therefore, needs to be established to accommodate P. septemspi- nosus gen. nov. et comb. nov. and P. heptacanthus gen. nov. et comb. nov. Zoosyst. Evol. 96 (1) 2020, 205-216 Results and discussion All three genes (12S, 16S and 18S) were successfully amplified and sequenced only for Pohleus septemspino- sus gen. nov. et comb. nov. The final alignment for this combined analysis of nuclear and mitochondrial genes included a total of 2659 bp (1868 bp 18S, 417 bp 16S and 372 bp 12S). The data were not saturated, considering the R? value for transitions R? (s) = 0.8665 and R? (v) = 0.9856 transversions, demonstrating that the sequences are appropriate for phylogenetic studies at this level. Pohleus septemspinosus gen. nov. et comb. nov. has a high support value (84%) as sister to a clade containing Thersandrus compressus and Macrocoeloma spp. Thersan- drus, presently assigned to the subfamily Majinae sensu Ng et al. (2008), is supported 89% as basal to Macrocoeloma and nested between Macrocoeloma and Pohleus gen. nov. in all scenarios analysed and with high support (Fig. 1). Thus, we believe that Thersandrus could be better fitted in Pisinae, based on the molecular results. Pohleus gen. nov. nested amongst other Pisinae as expected (Fig. 1). Systematics Family Epialtidae MacLeay, 1838 Subfamily Pisinae Dana, 1851 Pohleus gen. nov. http://zoobank.org/6FB44605-6302-44ED-8667-F4B321C21C11 Type species. Pericera septemspinosa Stimpson, 1871, by present designation. Included species. Pohleus septemspinosus (Stimpson, 1871) gen. nov. et comb. nov. and Pohleus heptacanthus (Bell, 1836) gen. nov. et comb. nov. 207 Diagnosis. Carapace subglobose, covered by short velvet pubescence interspaced by dense rows of long, hooked and simple setae in all carapace regions. Carapace armed with strong spines, including seven sharp spines on posterior half: one short mesial metagastric, four long, strong, conical lateral spines (two in each branchial re- gion) aligned with one cardiac spine and one intestinal spine; lateral spines longest, slightly directed upwards. Pterygostomial region with strong spines visible in dor- sal view. Rostrum bifurcated, base elongated, fused, di- verging abruptly forming a Y-shape, ending in acute tips. Pre-orbital spine strong, acute, directed upwards; post-or- bital spine short, acute. Basal article of antenna with three spines, one tubercle, not visible in dorsal view. Cheliped long, merus armed with short spines or tubercles, gran- ulated. P2 shorter than cheliped, dactylus much shorter than propodus. Thoracic sternal somite IV with later- al margins straight. Sternite VHI concealed by pleon in males. Male and female pleon with six somites not fused plus telson. Male telson tight-fitting into sterno-pleonal cavity, distinctly triangular. G1 slender, straight, with bi- lobed apex. G2 slender, straight, tapering distally, short about one fifth of G1 length. Comparative material. Libinia spinosa H. Milne Ed- wards, 1834 — Brazix * 1 male, | female; Macaé, near Santana Archipelago, PITA stn 12 HI (MZUSP 20271). Macrocoeloma camptocerum (Stimpson ,1871) — USA ¢ 13 males, 7 females, 7 ovig. females; Sanibel Island, 26.440359N, 82.113705W, 0.54—-11 m depth; Mar 1938; F A Chace Jr. leg. det. (MCZ 10191). Macrocoeloma concavum Miers, 1886 — VENEZUELA * | female; Cos- ta de Falcon, UTM 378365 and 1358259; 27 Apr 2007 (GIC040). Brazit *« 1 male; Paraiba, Projeto Algas, stn 85-B; 04 Jun 1981; Apr 2008, L E A Bezerra det. (MZUSP 5937). Macrocoeloma diplacanthum (Stimp- son, 1860) — US VirGIN ISLANDS * | male; Saint Thom- “ EE Vacrocoeloma trispinosum a7 82 83 | 89 91 34 7 ns Vacrocoeloma camptocerum amas \acrocoeloma eutheca Macrocoeloma concavum —nes Vcr ocoe/oma laevigatium Macrocoeloma villosum aes Vcr ocoe/oma diplacanthum ns Vicrococioma subparallelum es TET SONAT US COMPIeSSUS —— es Poles septemspinosus gen. nov. comb. nov. Leptopisa setirostris —_—_—_—_—— a a Stenocionops tS (177!) -———_—_———_| 0.020 Elamena producta Figure 1. Molecular phylogenetic tree represented as maximum likelihood topology of two mitochondrial and one nuclear loci (12S, 16S and 18S) to place Pohleus septemspinosus (Stimpson, 1871) gen. nov. et comb. nov. based on six close genera. Nodal support values represent the frequencies observed, using 1000 bootstrap pseudo-replicates. Values below 50% are not represented. zse.pensoft.net 208 as, R/V Albatross; 17-24 Jan 1884 (USNM 16182). GUADELOUPE °* 1 ovig. female; 16°13'37.3188"N, 61°32'23.0388"W, KARUBENTHOS 2012, stn GD49; 21 May 2012 (MNHN_ IU-2013-6755). Macrocoeloma eutheca. (Stimpson,1871) — USA « 1 ovig. female; off North Carolina, 33°48'06"N, 76°34'42"W, 77 m depth; 03 Apr 1981; Duke University for MMS 0S05 exped., 1981, P Krikorian det. (USNM 220812). Macrocoelo- ma intermedium Rathbun, 1901 — CusBa ¢ male holo- type; off Havana, R/V Albatross, stn 2323, 23°10'51"N, 82°19'03"W, 298 m depth; 17 Jan 1885; M J Rathbun det. (USNM 9492). CoLomsia * | female; Santa Marta; 29 Jun 1975; M Vélez det. (SMF 9093). Macrocoeloma laevigatum (Stimpson, 1860) — USA « 1 male; Florida, Hawk Channel, R/V Fish Hawk, stn 7429, 4.2 m depth; 27 Jan 1903 (USNM 46933). BAHAMAS * | female; off Whale Cay, 23.7 m depth, 9 Jul 1904, F A Chace Jr. det. (MCZ 8927). Macrocoeloma maccullochae Garth, 1940 — Mexico * 1 male, | female; Isabel Island, Allan Han- cock Pacific exped., R/V Velero III, stn 747-37, 18-32 m depth; 2 Apr 1937; W Schmitt leg., 1940; J S Garth det. (NHMLAC-AHF 372). Macrocoeloma nodipes (Desbonne in Desbonne & Schramm, 1867) — USA» 3 males, 2 females; Florida, Off Cape Sable, R/V Fish Hawk, stn 7351, 25°09'45"N, 81°18'35"W, 17 Dec 1902, 5.9 m depth (USNM 46922). GRENADA * 1 male; Grand Anse Bay, 12°01'45.19"N, 61°45'21.29"W, 11 Nov 2012, L RL Simone, A P Dornellas, V S Amaral leg., 27 Nov 2019, J Colavite det. (MZUSP 40162). Macrocoeloma subparallelum (Stimpson, 1860) — US VirGIN ISLANDS ¢ male lectotype; soft shell, cl. 13.98 mm, cw. 9.90 mm; St. Thomas, 18.345591N, 64.923613W, no date, A H Ri- ise leg., 1860, W Stimpson det. (MCZ 1243). BRAziIL * 3 females; Porto da Barra, Salvador, left side, 24 Apr 2006, R Bispo, R Jhonsson, W Santana, F Faria leg., Apr 2008, G Melo det. (MZUSP 18626). Macrocoeloma trispino- sum (Latreille, 1825) — USA « 1 ovig. female; Kings- ton Harbour; 1893; R P Bigelow leg., M J Rathbun det. (USNM 17959). ANTiIGUA * 1 male; English Harbour, Antigua-Barbados exped., 1918; University of Iowa State exped., M J Rathbun det. (USNM 72956). Mac- rocoeloma villosum (Bell, 1836) — EcuADoR *¢ 3 males, 1 female; Salinas, Walter Rathbone Bacon travelling Scholarship exped., stn 1, 2, 3; 12-14 Sep 1926; W L Schmitt leg., M J Rathbun det. (USNM 70942). Thersan- drus compressus (Desbonne in Desbonne & Schramm, 1867) — BELIzE * 1 male; west Bay, 1.3 m depth; 09 Jun 1985 (USNM 1526077). Remarks. Macrocoeloma Miers, 1879 s. str. is an am- phi-American genus with 12 species. This genus 1s char- acterised by the pyriform or triangular carapace, densely covered by short, velvet-like setae; with well-developed bifurcated or parallel rostral spines; the eyes completely protected by the orbits when retracted; orbits composed by the pre-orbital and the post-orbital spines and one or zse.pensoft.net Colavite, J et al.: A new genus for Pericera septemspinosa and Pericera heptacantha two projections of the basal article of antenna forming a functional, laterally projected protective hood. Although some of these characters can be observed in Pohleus gen. nov., the new genus can be easily distinguished from Macrocoeloma s. str. by a unique combination of characters, which include: (i) carapace relatively more globose in Pohleus gen. nov. (Figs 2A—C, 3, 4) (vs. car- apace subtriangular or pyriform in Macrocoeloma, Fig. 2E, F); (11) carapace covered by short velvet pubescence interspaced by dense rows of long hooked and simple setae in all carapace regions in Pohleus gen. nov. (Fig. 2C) (vs. carapace densely covered by velvet pubescence with hooked setae in specific regions of the carapace in Macrocoeloma), (111) basal article of antenna with small spines ventrally directed, not visible in dorsal view in Pohleus gen. nov. (Fig. 2B) (vs. with a long laterally-di- rected spine, between the rostral and pre-orbital spine, visible in dorsal view of Macrocoeloma, except in Mac- rocoeloma diplacanthum, Fig. 2F); (iv) pterygostomial region with a strong spine, laterally projected, visible in dorsal view in Pohleus gen. nov. (Fig. 2B) (vs. ptery- gostomial spines short, reduced or not visible in dorsal view in Macrocoeloma, Fig. 2F); (v) male sterno-pleo- nal cavity with no crest anteriorly in Pohleus gen. nov. (Fig. 2B); (vs. male sterno-pleonal cavity with a distinct crest anteriorly on thoracic sternite IV in Macrocoeloma, Fig. 2E); (vi) male sternite IV almost straight laterally in Pohleus gen. nov. (Fig. 2B) (vs. sternite IV deeply con- cave laterally in Macrocoeloma; Fig. 2E); (vii) episterni- tes IV and V forming a continuous line with the sternite, slightly downward directed in Pohleus gen. nov. (Fig. 2B) (vs. episternites IV, V and VI upward directed in Macrocoeloma, Fig. 2F); and (viii) sternal sutures shal- low in Pohleus gen. nov. (Fig. 2B) (vs. sternal sutures deep sculpted in Macrocoeloma;, Fig. 2F). In Macrocoeloma, the gonopods are highly variable amongst species, but it is possible to recognise a general pattern with G1 being longer than the thoracic sternal su- ture V/V, parallel and usually with a bilobed apex (except in M. concavum, M. intermedium and M. laevigatum that have a unilobed apex). Although Pohleus septemspinosus gen. nov. et comb. nov. (G1 of Pohleus heptacanthus gen. nov. et comb. nov. not examined) can be fitted in this gen- eral pattern, the G1 apex is notably more similar to the G1 apex of Libinia Leach, 1815 (Fig. 2D and see Tavares and Santana 2011: 63, fig. 2D for Libinia spinosa). Thersandrus Rathbun, 1897, is a monotypic genus exhibiting extremely efficient camouflage behaviour as Macrocoeloma and Pohleus gen. nov. However, Ther- sandrus does not actively decorate itself, presenting crypsis behaviour consisting of matching the body to the environment in shape and colour, being morphologically adapted to live on green algae fronds. For instance: (1) Thersandrus has a carapace covered by long setae giv- ing a felt-like texture (vs. velvet-like and hooked setae in Macrocoeloma and Pohleus gen. nov.); (i1) the carapace Zoosyst. Evol. 96 (1) 2020, 205-216 209 Figure 2. Pohleus septemspinosus (Stimpson, 1871) gen. nov. et comb. nov. A, B. male, (USNM 1256361); C. male (USNM 241030); D. sterno-pleonal cavity with first (G1) and second (G2) gonopods in place (USNM 1256361). Macrocoeloma trispino- sum (Latreille, 1825); E, F. male (USNM 17959). A, E. habitus; B, F. ventral view; C. frontal view; D. pleonal view in detail. Note the spine of the basal article of antenna (white arrow); pterygostomial spine (grey arrow); margin of the IV sternite thoracic (black arrow). Scale bars: 10 mm. zse.pensoft.net 210 and pereopods are flattened in Thersandrus (vs. carapace subtriangular or piriform, not flattened and with cylindri- cal pereopods in Macrocoeloma and subglobose carapace and cylindrical pereopods in Pohleus gen. nov.); (iii) the orbital spines are reduced, not forming a hood in Ther- sandrus (vs. orbital spines long, blunt, forming a hood in Macrocoeloma and long, acute and forming a hood in Pohleus gen. nov.); all characters that prevented us from synonymising Thersandrus to Macrocoeloma. However, it 1s important to note that, based on the molecular re- sults, Thersandrus should be transferred from Mayidae to Pisinae as mentioned above. Etymology. Generic name in honour of the renowned marine biologist and carcinologist Gerhard Werner Pohle (Atlantic Reference Centre, Huntsman Marine Science Centre). Gender masculine. Pohleus septemspinosus (Stimpson, 1871) gen. nov. et comb. nov. Figs 2A—D, 3, 5A, B Pericera septemspinosa Stimpson, 1870 (1871): 113 [type locali- ty: West of Tortugas; 65 m depth, type material: non-extant]. — A. Milne-Edwards 1873: 59, 200, pl. 15A, fig. 2; Gundlach and Torral- bas 1900: 365, fig. 366G. Macrocoeloma septemspinosa — Miers, 1886: 80; Rathbun, 1892: 250; 1898b: 257; 1899: 576. Macrocoeloma septemspinosum — Moreira 1901: 64, 136; Rathbun 1925: A477, pl. 173, figs 2, 3; Coelho 1971:142; Coelho and Ramos 1972: 218; Soto 1980 (digital document); Powers 1977: 52; Takeda and Okutani 1983: 141; Abele and Kim 1986: 45, fig. 521A; Melo 1996: 219, fig. 1; 1998: 464; Coelho-Filho 2006: 19; Almeida et al. 2007: 15; Alves et al. 2008: 58; Ng et al. 2008: 119; Alves et al. 2012: 54. Neotype (Here designated). USA « male neotype, cl 31 mm, cw 35 mm: Florida, West of Sarasota, R/V Oregon, stn 4088, 27°44'N, 83°45'W; 04 Dec 1962, National Marine Fisheries Service exped.; 27 Oct 2014, W Santana det. (USNM 1256361) (Fig. 2A, B, D). Material examined. USA « 1 male; North Carolina, 33°48'06"N, 76°34'42"W, 77 m depth; 03 Apr 1981, Duke Univ. for MMS leg.; 1981, P Krikorian det. (USNM 220811). * 1 male, 1 juv. female; South Carolina, East of Cape Romain, R/V Albatross, stn 2311, 32°55'00"N, 77°54'00"W, 59.1°F, 114.4 m depth; 05 Jan 1885, U.S. fish commission leg. (USNM 15127). * 1 juv.; Florida, between Cedar Keys and Delta of Mississippi, R/ V Albatross, stn 2369-2374, 46-48 m depth; Feb 1885 (USNM 46957). ¢ 1 juv. female; Gulf of Mexico, Southwest of Cape San Blas, R/V Albatross, stn 2373, 29°14'00"N, 85°29'15"W, 45.7 m depth; 07 Feb 1886 (USNM 15132). « 1 juv. fe- male, 1 male; same collection data as preceding (USNM 15130). * 2 females; same collection data as preceding (USNM 15131). * 1 male; Suez of Mexico, R/V Oregon, stn 892, 28°55'N, 85°07'W, 53 m depth; 07 Mar 1954, Fish and Wildlife leg.; 21 Apr 1954, F A Chace Jr det. (USNM zse.pensoft.net Colavite, J et al.: A new genus for Pericera septemspinosa and Pericera heptacantha 96389). * 1 male; off Apalachicola Bay, R/V Indian Seal, 28°49'59"N, 85°37'02"W, 177 m depth; 31 Jan 1978; R Lemaitre det. (USNM 1085620). * 1 male; South of Dog Island, R/V Albatross, stn 2407, 28°47'30"N, 84°37'00"W, 43.8 m depth; 07 Feb 1886, U.S. fish commission exped.; M J Rathbun det. (USNM 15135). * 3 juv. females; South of stn George Island, R/V Albatross, stn 2406, 28°46'00"N, 84°49'00"W, 47.5 m depth; 15 Mar 1885 (USNM 15134). ° 1 female; 26°45'52"N, 83°21'26"W, 50.2 m depth; 18 Jul 1981, Continental Shelf Associates exped.; R Lemaitre det. (USNM 273379). * 1 male; same collection data as pre- ceding (USNM 241027). ¢1 juv. female; same collection data as preceding (USNM 236995). « 1 male; 26°16'S0"N, 83°23'49"W, 55.5 m depth; 05 Feb 1982, Continental Shelf Associates exped.; R Lemaitre det., (USNM 241026). « 1 male, 2 females; same collection data as preceding (USNM 273381). ¢ 1 female, 1 male; same collection data as pre- ceding (USNM 241024). « 1 male, 1 ovig. female; same collection data as preceding (USNM 229838). * 1 male; 26°16'44"N, 83°42'49"W, 71.3 m depth; 03 Nov 1980, Continental Shelf Associates exped.; R Lemaitre det. (USNM 236994). * 1 juv. female; same collection data as preceding (USNM 241007). « 1 male; same collection data as preceding (USNM 236987). * 2 males; same collection data as preceding (USNM 236986). * 1 male; 26°16'43"N, 83°46'49"W, 77 m depth; 24 Jun 1981, Continental Shelf Associates exped.; R Lemaitre det. (USNM 241029). ¢ 1 female; same collection data as preceding (USNM 273382). © 1 male; 25°45'35"N, 83°20'14"W, 58.5 m depth; 24 Apr 1981, Continental Shelf Associates exped.; R Le- maitre det. (USNM 242947). « 1 male; R/V Silver Bay, 25°32'N, 80°04'W, 65.8 m depth; 24 Oct 1960; Oct.1970, D J G Griffin det. (USNM 1278767). * 1 female; off Dry Tortugas, 24°34'N, 83°16'W, 65.8 m depth; Dec 1877- Jan 1878, USCSS Blake exped.; A. Milne-Edwards det. (MCZ 8206). * 1 male, 1 female; off Key West, R/V Albatross, 24°25'45"N, 81°46'45"W, stn 2317, 75 °F, 82.3 m depth; 15 Jan 1885, U.S. fish commission exped. (USNM 15128). ¢ 1 male, 1 female; same collection data as preceding (USNM 15129). * 1 male; Off Key West, Sand key Light bearing West Northwest, Key West Light bearing North, State Univ. Iowa exped., stn 24, 109.7 m depth; 19 Jun 1893, M J Rathbun det. (USNM 75724). * 1 male; same collection data as preceding (USNM 72863). ¢ 1 male, 1 juv. female; Sand Key Light bearing Northwest by North, Key West Light North 0.5mile East, 91.4-109.7 m depth (USNM 68913). ¢ 1 male; Florida, Pompano; 23 May 1949 (AMNH 10961). Bananas ¢ 1 juv. female; Bahamas Bank; 18 May 1893, State Univ. Iowa Bahamas exped. (USNM 72862). COLOMBIA, * | male; 2 miles Southwest of Cape La Vela; 8 Apr 1939, J Garth leg.; 17 Jan 2018, J Colavite det. (AHF 39295). * 1 juv. female; 2 miles off Bahia Hon- da, R/V Velero II, stn A15-39, 9-18.28 m depth; 08 Apr 1939, J Garth det. (AHF 39292). VENEZUELA * | juv. male; 7 miles of Tortugas Island, R/V Velero HI, stn A43-39, 73- 75 m depth; 21 Apr 1939, J Garth det. (AHF 39293). « 1 male; 125 miles northeast of Macaibo, 12°37'N, 70°45'W, R/V Oregon, stn 4400, 97 m depth; 26 Sep 1963; 27 Oct 2014, W Santana det. (USNM 1256370). * 1 male; 50 miles Zoosyst. Evol. 96 (1) 2020, 205-216 northeast of Caracas, 10°44'N, 66°09'W, R/V Oregon, stn 4466, 73 m depth; 17 Oct 1963; 23 Oct 2014, W Santana det. (USNM 1256347). FRENCH GUIANA * | ovig. female; Guiana 2014 exped., R/V Hermano Gines, 6°17'58.2"N, 52°13'18.5952"W, 95-97 m depth; 08 Aug 2014 (MNHN IU 2013-2682). BRAziL * 1 ovig. female; Recife, dredge 2; J Colavite det. (R2 unnumbered). * 1 ovig. female; Bahia, Ilhéus, Costa de Ilhéus, 14°43'33"S, 38°57'20"W, 41-42 m depth; 28 Nov 2004, A O Almeida det.; old number MZU- ESC 406 (CIASB M. 2017.0084. UESC). Diagnosis. Rostrum width half of interorbital length bi- furcated, base elongated, fused, diverging abruptly form- ing a Y-shape. Pleonal somite IT smooth; merus of second pereopod armed with a spine. Description. Cephalothorax and appendages sparsely cov- ered with short, velvet-like pubescence. Carapace subglo- bose wider than long, convex, with long lines of hooked and simple setae in all carapace regions, denser in rostral and lateral spines. Rostrum width half of interorbital length bifurcated, base elongated and fused, abruptly diverging forming a Y-shape, ending in acute tips. Interorbital re- gion slightly depressed medially. Hepatic region broad. One metagastric spine. Four long, strong, conical lateral spines (two in each branchial region), in line with the car- diac spine. One short intestinal spine. Orbital region very prominent, eyes completely protected in orbit when re- tracted, ocular peduncle visible when not retracted. Pre-or- bital spine acute, directed upwards, tip curved, longer than post-orbital spine, ventral margin of pre-orbital spine with a small crenulation; post-orbital spine curved upwards. Antennular fossae wider than long, margins smooth. Interantennular septum elongated, compressed laterally, forming distinct, ventrally-directed lobe. First and second antennal articles fused to epistome, suture between an- 211 tenna and epistome visible, antennal gland opening near suture line. Basal article of antenna with three spines, one tubercle, not visible in dorsal view. Antennal flagella lon- ger than rostral spines, behind rostrum in dorsal view. Epistome anterior margin narrower than antennular fossae, smooth. Buccal field sub-quadrate, narrower at posterior edge with acute spine at anterolateral angle in line with antennal spines. Third maxillipeds completely covering buccal field. Exopod long, nearly reaching dis- tal margin of merus. Pterygostomial region subtriangu- lar, slightly inflated, separated from subhepatic region by marked groove, one long, strong spine slightly curved upwards on medial margin, visible in dorsal view. Chelipeds equal, longer than pereopods in adults, more robust in adult males; females chelipeds shorter than males, slender. In males, ischium, merus, carpus and propodus segments granulate. Ischium with one prominent tubercle laterodistally. Merus with one spine on proximal half, one on distal margin. Carpus with four tubercles sparsely distributed. Dactylus arched in adult males, a small gap between fingers, distinctly shorter than palm. Cutting edges with sub-equal teeth in distal half, one distinct proximal tooth in larger males; distal half with dark brown colour in fixed specimens. Female ischium, merus, carpus and propodus with smaller tu- bercles than males, dactylus slightly arched, without gap between fingers. Pereopods short, slender, cylindrical. P2 longest, P3— PS progressively decreasing in length. P2 merus with dis- tinct spine in distolateral margin. Dactylus slightly curved, shorter than propodus, smooth ventrally, with corneous tip. Male thoracic sternites I-IV fused, broadly triangular, smooth, anterior half declivous in ventral view, forming a carina along the sterno-pleonal cavity margin. Thoracic sternal somite I'V with lateral margins straight. Telson ful- ly fitted to cavity, anterior margin smooth. Sternite VHI Figure 3. A. Habitus, dorsal view and B. ventral view of Pohleus septemspinosus (Stimpson, 1871) gen. nov. et comb. nov., female (R2 unnumbered). Colour in life with debris and algae for camouflage. Scale bars: 10 mm zse.pensoft.net 212 concealed by pleon. Episternites IV and V forming a con- tinuous line with the sternite, slightly downward directed. Pleonal somites I-VI, telson free in males and females, slightly raised medially forming low longitudinal ridge. Male telson sub-triangular, apex rounded. Female pleon markedly arched, with row of setae marginally. Female telson transversally ovate. G1 longer than thoracic sternal suture IV—V, stout, straight, parallel, with torsion in distal half, apex bilobed; mesial lobe short, with tip curved upwards; lateral lobe long, slightly arched, ending in an acute tip; lateral mar- gin smooth. G2 slender, straight, very short (one fifth of G1 length), with disto-medial process. Colour in life. Carapace light brown; cephalothorax ven- tral, pleon and pereopods pinkish to purple (Fig. 3). Neotype locality. USA, Florida, west of Sarasota, 27°44'N, 83°45'W. Geographic distribution. Western Atlantic: USA (from North Carolina); Gulf of Mexico; Bahamas; Venezuela and Brazil (from Ceara to Bahia) (Fig. 4). Remarks. The type material of Pericera septemspino- sa was probably lost in the Great Chicago Fire in 1871 (see Evans 1967; Deiss and Manning 1981; Manning 1993; Vasile et al. 2005; Manning and Reed 2006). The male (USNM 1256361) is here designated as neotype of Pericera septemspinosa Stimpson, 1871 due to the close morphological similarity to the original description. The neotype is from a region close to the type locality. The specimen chosen here is from west of Sarasota, 27°44'N, 83°45'W (cf. ICZN art.75 and 76). Pohleus septemspinosus gen. nov. et comb. nov. can be distinguished from its Pacific congener, Pohleus hep- tacanthus gen. nov. et comb. nov., by the following char- acters: (1) rostrum width half of the interorbital length in P. septemspinosus gen. nov. et comb. nov. (vs. rostrum with one-third or less of the interorbital length in P. hep- tacanthus gen. nov. et comb. nov.; Fig. 5C, D); (11) ple- onal somite II smooth in P. septemspinosus gen. nov. et comb. nov. (vs. pleonal somite IT with one spine or tuber- cle in P. heptacanthus gen. nov. et comb. nov.; Fig. 5C, D); (111) merus of the second pereopod armed with a spine in P. septemspinosus gen. nov. et comb. nov. (vs. merus of second pereopod unarmed in P. heptacanthus gen. nov. et comb. nov.; Fig. 5C, D). Unfortunately, no male spec- imens of Pohleus heptacanthus gen. nov. et comb. nov. were available for study, thus, the gonopodal differences cannot be ascertained. The southeast record of this species as Espirito Santo State, in Brazil (Serejo et al. 2006: fig. 8C) is not valid, since the specimen (MNRJ 17062) was re-identified as Macrocoeloma concavum. Therefore, the distribution of to Pohleus septemspinosus gen. nov. et comb. nov. is cor- rected to the Brazilian coast, from Ceara to Bahia. zse.pensoft.net Colavite, J et al.: A new genus for Pericera septemspinosa and Pericera heptacantha Pohleus heptacanthus (Bell, 1836) gen. nov. et comb. nov. Fig. 5C, D Pericera heptacantha Bell, 1836: 173 [type locality: Puerto Potrero, Central America, 23.7 m depth; type material: syntypes, 1 male (non-extant), 1 female (OUM 13764)]-— Bell 1836b: 61, pl. 12, fig. 6 and 6r-u; White 1847: 10; A. Milne-Edwards 1873: 55. Macrocoeloma heptacantha — Miers 1886: 79, 81. Macrocoeloma heptacanthum — Rathbun 1898a: 576; 1925: 473, pl. 173, fig. 1; pl. 269, fig. 8-11, text-figs 133, 134; Garth 1958: 415; Di Mauro 1982: 170; Ng et al. 2008: 119. Lectotype (Here designated). Costa Rica * 1 female, cl: 35 mm, cw: 43 mm; Central America, Puerto Potrero, 23.7 m depth; H Cumming leg. (OUM 13764) (Fig. 5C, D). Material examined. Mexico « | juv. female; Off Cape San Lucas, R/V Albatross, stn 2829, 22°52'00"N, 109°55'00"W, RKY leg., 56.6 m depth, 74.1 °F; 01 May 1888, M J Rathbun det. (USNM 21933 illustrated). PANA- MA * | juv. female; Panama Bay, R/V Albatross, stn 2798, 8°10'30"N, 78°50'30"W, 114.6 m depth; 05 Mar1888, M J Rathbun det. (USNM 21932). Diagnosis. Rostrum width less than one-third of interor- bital length, bifurcated, base elongated, fused, diverging abruptly forming a Y-shape, ending in acute tip. Pleon- al somite II with one spine or tubercle. Merus of second pereopod smooth. Description based on female lectotype (male char- acters modified from Bell 1836b). Cephalothorax and appendages slightly covered with short, velvet-like pu- bescence. Carapace subglobose wider than long, con- vex, with long lines of hooked and simple setae in all regions. Rostrum short, less than one-third of interor- bital length, bifurcated, base elongated, fused, diverg- ing abruptly forming a Y-shape, ending in acute tips. Interorbital region slightly depressed medially. Hepatic region broad. One metagastric spine. Four long, strong, conical lateral spines (two in each branchial region), in line with cardiac spine. One short intestinal spine. Orbital region very prominent, eyes completely pro- tected when retracted, ocular peduncle visible when not retracted. Pre-orbital spine directed upwards, slight- ly curved on tip, longer than post-orbital spine, ven- tral margin of pre-orbital spine with small crenulation; post-orbital spine curved upwards. Antennular fossae wider than long, margins smooth. Interantennular septum elongate, laterally compressed, forming distinct ventrally-directed lobe. First and second antennal articles fused to epistome, with suture between antenna and epistome visible, antennal gland opening near suture line. Basal article of antenna with two spines, not visible in dorsal view: proximal spine smallest. An- Zoosyst. Evol. 96 (1) 2020, 205-216 North America Eastern Pacific 2250 km 213 Western Atlantic Figure 4. Geographic distribution Pohleus heptacanthus (Bell, 1836) gen. nov. et comb. nov.; orange circles = distribution based on examined material; green star = type locality and Pohleus septemspinosus (Stimpson, 1871) gen. nov. et comb. nov.; red circles = distribution based on examined material; yellow star = neotype locality. tennal flagella longer than rostral spines, behind rostrum in dorsal view. Epistome anterior margin narrower than antennular fos- sae, smooth, posterior margin slightly depressed. Buccal field sub-rectangular, narrower at posterior edge with one acute spine in anterolateral angle aligned with antennal spines. Third maxillipeds covering buccal frame when closed, leav- ing a small gap between ischia. Exopod long, nearly reach- ing distal margin of merus. Pterygostomial region subtrian- gular, slightly inflated, separated from subhepatic region by marked groove, with one long, strong spine slightly curved upwards on medial margin, visible in dorsal view. Male chelipeds equal, longer than pereopods; covered with sparse granulation, unarmed. Dactylus arched in adult males, leaving small gap between fingers, distinctly shorter than palm. Cutting edges with sub-equal teeth in distal half, one distinct proximal tooth in larger males; distal half with light brown colour in fixed specimens. Pereopods short, slender, cylindrical. P2 longest; P3-P5 progressively decreasing in length. Dactylus slightly curved, covered with short setae. Female chelipeds equal, longer than pereopods, slen- der and smooth. Dactylus arched in adult, shorter than palm, sub-equal teeth in distal half. Pereopods, slender, cylindrical. P2 longest, P3-P5 progressively decreasing in length. Dactylus slightly curved, shorter than propodus, smooth ventrally, with corneous tips. Male thoracic sternites I-IV fused, broadly triangular, smooth, anterior half declivous in ventral view. Telson fully fitted to cavity, anterior margin smooth. Female pleonal somites I-VI markedly arched, telson free, transversally oval, with a row of setae on margin and one small spine in first somite. Male pleon rather prom- inent, pleonal somites I-VI, telson free, somite II with a mesial tubercle. Somite HI with slight elevations. Somite VI longest, with a mesial tubercle and a small projection each side. Colour in life. Light brown, covered with darker hair, first pair of pereopods reddish (Bell 1936b). zse.pensoft.net 214 Colavite, J et al.: A new genus for Pericera septemspinosa and Pericera heptacantha Figure 5. Pohleus septemspinosus (Stimpson, 1871) gen. nov. et comb. nov. A, B. female (MNHN IU 2013-32682). Pohleus hep- tacanthus (Bell, 1836) gen. nov. et comb. nov.; C, D. female lectotype (OQUM 13764). A, C. habitus; B, D. ventral view. Note the Spine on merus of the second pereopod (white arrow). Scale bars: 10 mm. Type locality. Costa Rica, Central America, Puerto Po- trero, in sand at a depth of 23.7 m. Geographic distribution. Eastern Pacific: Mexico (Cape San Lucas), Costa Rica (Guanacaste) and Panama (Pana- ma Bay) (Fig. 4). Remarks. Bell (1836) described Pericera heptacan- tha, based on two specimens as syntypes, one male and one female. The male syntype is considered lost and the female is deposited in the dry crustacean collection of the Oxford University Museum (OUM 13764) (Di Mauro 1982). Thus, the female syntype (OUM 13764) is here designated as the lectotype of Pericera hep- tacantha since it 1s the only specimen from the type series remaining (Fig. SC, D). The locality Puerto Po- trero in Central America probably refers to the Puerto Potrero, Guanacaste, in Costa Rica. Rathbun (1937: 136) examining Lithadia cumingii Bell, 1855, a spe- cies described from the same locality by Bell (1855), referred to the type locality as from Potrero, Costa Rica, the same case as for Pohleus heptacanthus gen. nov. et comb. nov. zse.pensoft.net Acknowledgements We are grateful to Rafael Lemaitre and Karen Reed (USNM) for facilitating access to materials and informa- tion from the USNM collections and to Marcos Tavares (MZUSP) and Alexandre Almeida for access to the com- parative material and all visited institutions for access to materials and information. Thanks to the Laboratory of Analytical Biology at Smithsonian Institute and Labora- tory of Molecular Biology at MZUSP for allowing the use of their facilities during the analyses. Thanks to Patricia Souza and Andressa Cunha for the photos of the fresh specimen arranged in this paper. This work was supported by the Funda¢ao de Amparo a Pesquisa do Estado de Sao Paulo (FAPESP) [2013/01201-0 and 2014/15549-0]. JC thanks FAPESP [2016/02775-8] for supporting her doc- torate fellowship at UNESP. We also thank Darryl Felder at the University of Louisiana at Lafayette for use of DNA sequences generated by AMW under the USA Na- tional Science Foundation grant NSF/AToL EF—0531603 to DF. WS thanks Gerhard W. Pohle for his friendship and support over the years, looking forward to our further fruitful cooperation. This work greatly benefited from Zoosyst. Evol. 96 (1) 2020, 205-216 the comments of Peter Ng (National University of Singa- pore), Ngan Kee Ng (National University of Singapore) and Paul Clark (Natural History Museum, UK). References Abele LG, Kim W (1986) An illustrated guide to the marine decapod crustaceans of Florida (Vol. 8). State of Florida Department of Envi- ronmental Regulation Technical Series, 760 pp. Almeida AO, Guerrazzi MC, Coelho PA (2007) Stomatopod and deca- pod crustaceans from Camamu Bay, state of Bahia, Brazil. 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Windsor, William Santana Data type: Species data Explanation note: Taxa included in the molecular phy- logenetic analyses to place the newly described taxa within the context of the family Pisidae. 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.96.50360.suppl1