Triphoridae (Gastropoda) from Martinique sampled by the MADIBENTHOS expedition, with notes on shallow-water species from Guadeloupe

Directeur De la publication / Publication director : Gilles Bloch Président du Muséum national d’Histoire naturelle réDactrice en chef / editor-in-chief : Laure Desutter-Grandcolas assistante De réDaction / assistant editor : Anne Mabille ([email protected]) Mise en page / Page layout : Anne Mabille coMité scientifique / scientific board : Nesrine Akkari (Naturhistorisches Museum, Vienne, Autriche) Maria Marta Cigliano (Museo de La Plata, La Plata, Argentine) Serge Gofas (Universidad de Málaga, Málaga, Espagne) Sylvain Hugel (CNRS, Université de Strasbourg, France) Marco Isaia (Università degli Studi di Torino, Turin, Italie) Rafael Marquez (CSIC, Madrid, Espagne) Jose Christopher E. Mendoza (Lee Kong Chian Natural History Museum, Singapour) Annemarie Ohler (MNHN, Paris, France) Jean-Yves Rasplus (INRA, Montferrier-sur-Lez, France) Wanda M. Weiner (Polish Academy of Sciences, Cracovie, Pologne) couverture / cover : Made with the figures of the article. 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Diffusion – Publications scientifiques Muséum national d’Histoire naturelle CP 41 – 57 rue Cuvier F-75231 Paris cedex 05 (France) Tél. : 33 (0)1 40 79 48 05 / Fax : 33 (0)1 40 79 38 40 [email protected] / https://sciencepress.mnhn.fr © Publications scientifiques du Muséum national d’Histoire naturelle, Paris, 2024 ISSN (imprimé / print) : 1280-9551/ ISSN (électronique / electronic) : 1638-9387 457 ZOOSYSTEMA • 2024 • 46 (18) © Publications scientifiques du Muséum national d’Histoire naturelle, Paris. www.zoosystema.com urn:lsid:zoobank.org:pub:650979F1-53CD-4B0A-B9A2-E6B1A49E9C2B Fernandes M. R. 2024. — Triphoridae (Gastropoda) from Martinique sampled by the MADIBENTHOS expedition, with notes on shallow-water species from Guadeloupe. Zoosystema 46 (18): 457-503. https://doi.org/10.5252/zoosystema2024v46a18. http://zoosystema.com/46/18 ABSTRACT This study aims to evaluate the taxonomic composition of triphorids from Martinique obtained by the MADIBENTHOS expedition, which comprised more than 500marine sampling events. In addition, some identifications of shallow-water triphorids from Guadeloupe (based on recent works) are corrected. After the analysis of 1615 specimens, 33 triphorid species were found in the shallow waters of Martinique, of which Inella spinosa n.sp., Isotriphora pardus n.sp. and Metaxia discus n.sp. are described as new species. New records are given for the Lesser Antilles, such as Cheirodonta dupliniana (Olsson, 1916), Strobiligera inaudita (Rolán & Lee, 2008) and Triphora martii Rolán & Fernández-Garcés, 1995. The taxonomy of some species is discussed (mainly on Nanaphora Laseron, 1958), and new generic allocations are proposed: Marshallora abacoensis (Rolán & Redfern, 2008) n. comb., Marshallora armandoi (Espinosa & Ortea, 2020) n. comb., Marshallora grenadensis (Rolán & Lee, 2008) n. comb., Marshallora monteiroi (Rolán & Fernández-Garcés, 2015) n. comb. The individual rarefaction curve of triphorids from MADIBENTHOS is nearly saturated but did not reach the asymptote. Seven species (all with planktotrophic development) were considered abundant or much abundant, whereas eight species (with planktotrophic or non-planktotrophic development) were represented by singletons or doubletons. Brushing and suction were efficient methods to obtain triphorids, and some dredging events recovered up to 13species (mainly deeper than 50m). More species were sampled in the Atlantic side of Martinique, but hyper-diverse stations were widespread along the island. The abundance of triphorids and species composition change in Martinique along the depth gradient; Iniforis turristhomae (Holten, 1802) is the most abundant triphorid between 0-30m, but Cosmotriphora melanura (C.B.Adams, 1850) predominates between 31-85m. Maurício Romulo FERNANDES Departamento de Zoologia, Instituto de Biociências, Universidade Federal do Estado do Rio de Janeiro (UNIRIO), Avenida Pasteur, 458, Urca, Rio de Janeiro (Brazil) [email protected] Submitted on 26 July 2023 | Accepted on 10 January 2024 | Published on 17 July 2024 Triphoridae (Gastropoda) from Martinique sampled by the MADIBENTHOS expedition, with notes on shallow-water species from Guadeloupe KEY WORDS Biodiversity, Lesser Antilles, larval development, depth zones, new combinations, new species. 458 ZOOSYSTEMA • 2024 • 46 (18) Fernandes M.R. RÉSUMÉ Les Triphoridae (Gastropoda) de Martinique échantillonnés lors de l’expédition MADIBENTHOS, avec des remarques sur les espèces des eaux peu profondes de Guadeloupe. Cette étude vise à évaluer la composition taxonomique des triphoridés de la Martinique collectés pendant l’expédition MADIBENTHOS, qui comprenait plus de 500 événements de collectes. De plus, certaines identifications des triphoridés peu profonds de Guadeloupe (basées sur des travaux récents) sont corrigées. Après l’analyse de 1 615 spécimens, 33 espèces de triphoridés ont été dénombrées dans les eaux peu profondes de la Martinique, dont Inella spinosa n.sp., Isotriphora pardus n.sp. et Metaxia discus n.sp. décrits comme nouvelles espèces. De nouveaux signalements sont proposés pour les Petites Antilles, comme Cheirodonta dupliniana (Olsson, 1916), Strobiligera inaudita (Rolán & H.G.Lee, 2008) et Triphora martii Rolán & Fernández-Garcés, 1995. La taxonomie de certaines espèces est discutée (principalement pour Nanaphora Laseron, 1958), et de nouvelles allocations génériques sont proposées: Marshallora abacoensis (Rolán & Redfern, 2008) n. comb., Marshallora armandoi (Espinosa & Ortea, 2020) n. comb., Marshallora grenadensis (Rolán & Lee, 2008) n.comb., Marshallora monteiroi (Rolán & Fernández-Garcés, 2015) n. comb. La courbe de raréfaction individuelle des triphoridés de MADIBENTHOS est presque saturée mais n’a pas atteint l’asymptote. Sept espèces (toutes à développement planctotrophe) sont considérées comme abondantes ou très abondantes, tandis que huit espèces (à développement planctotrophe ou non-planctotrophe) sont représentées par des “singletons” ou des “doubletons”. Le brossage et l’aspiration se sont révélés efficaces pour récolter des triphoridés, et certains dragages ont permis de récolter jusqu’à 13espèces (principalement à des profondeurs inférieures à 50m). Plus d’espèces ont été échantillonnées sur la façade atlantique de la Martinique, mais des stations hyper-diversifiées étaient répandues le long de l’île. L’abondance des triphoridés et la composition des espèces changent en Martinique le long du gradient de profondeur ; Iniforis turristhomae (Holten, 1802) est le triphoridé le plus abondant entre 0 et 30m, alors que Cosmotriphora melanura (C.B. Adams, 1850) prédomine entre 31 et 85m. INTRODUCTION Triphoridae Gray, 1847 is regarded as one of the “Big Five” families of marine gastropods in terms of species richness (Bouchet etal. 2002; Albano etal. 2011), which may be a consequence of their parasitic/micro-predatory feeding on sponges, i.e., the specialization on particular sponge hosts possibly triggered speciation and high diversity (Marshall 1983; Fernandes& Pimenta 2020; Nützel 2021). In fact, many taxonomic problems are derived from this great diversity, even more when only empty (and often worn) shells are available (Bouchet& Strong 2010). The discovery of Caribbean triphorids began in the early 19th century (Holten 1802), but reached a higher ground after the study of species from Jamaica (C. B. Adams 1850). Since then, several authors increased the knowledge about Caribbean triphorids after general catalogues of molluscs from particular localities (e.g. Dall& Simpson 1901; Warmke& Abbott 1962; Nowell-Usticke 1969; Vokes& Vokes 1983; De Jong& Coomans 1988; Merlano& Hegedus 1994; Hewitt& van Leeuwen 2017) or studies focused on the description of new species (e.g. Moolenbeek& Faber 1989; Faber& Moolenbeek 1991; Rolán& Luque 1999). Recent taxonomic revisions (with good illustrations) of Triphoridae from Caribbean sites include material from Cuba (e.g. Rolán& Fernández-Garcés 2007, 2008) and Bahamas (Redfern 2013). Based on Rosenberg (2009), the triphorids of Guadeloupe and Martinique were untouched until the description of Triphora guadaloupensis Rolán& Fernández-Garcés, 2008, endemic from Guadeloupe. Further, Rolán& FernándezGarcés (2015) studied the shallow-water triphorids from Guadeloupe, sampled by the KARUBENTHOS expedition (hereafter, KARUBENTHOS 1). Most of their identifications were followed by Lamy& Pointier (2018), which also included new material from Guadeloupe and Martinique; to the latter site, only Inella triserialis (Dall, 1881), Iniforis turristhomae (Holten, 1802), Marshallora nigrocincta (C. B.Adams, 1839), Monophorus olivaceus (Dall, 1889) and Nototriphora decorata (C. B. Adams, 1850) were recorded. These works greatly improved the knowledge about triphorids from the French Antilles, but there were mistakes on some of these taxonomic identifications, requiring corrections. The MADIBENTHOS expedition was organized by the MNHN (Muséum national d’Histoire naturelle, France) team and conducted in Martinique between September-October 2016, comprising 503 sampling events in the shore and sea, such as dredging, trap lines, brushing of hard substrata and suction sampling (Bouchet etal. 2019). Sampling stations were performed down to 100m, covering all portions of the island (Bouchet etal. 2019). Several new species of gastropods were recently described from material obtained by the expedition (e.g. Espinosa& Ortea 2017; Ortea& Buske 2018; Garrigues& Lamy 2019; Cecalupo& Perugia 2020), and others are still to come. The objective of this study is to evaluate the taxonomic composition of triphorids obtained by the MADIBENTHOS. In addition, some identifications of shallow-water triphorids from Guadeloupe (Rolán& Fernández-Garcés 2015; Lamy& Pointier 2018) are corrected. The deep-sea species from Guadeloupe, obtained by the KARUBENTHOS 2 expedition, will be treated in another study. MOTS CLÉS Biodiversité, Petites Antilles, développement larvaire, zones de profondeur, combinaisons nouvelles, espèces nouvelles. 459 Triphoridae from Martinique ZOOSYSTEMA • 2024 • 46 (18) MATERIAL AND METHODS Sampling and taxonomic procedureS Stations in which triphorids were obtained are listed under each species, following the five main geographic zones within Martinique proposed by Bouchet etal. (2019); their coordinates, depths and sampling methods are listed in Appendix1. All shallow-water species from Guadeloupe examined by the author are also listed, obtained by KARUBENTHOS 1 (Rolán& Fernández-Garcés 2015) and KARUBENTHOS 2 expeditions. Afew shells from Guadeloupe were illustrated; in such cases asterisks (*) are given in the plates. Live triphorids from Guadeloupe were stored in ethanol during MNHN expeditions, and these will be studied in an ongoing molecular phylogeny of Triphoroidea (unpublished data); those from Martinique were mainly stored in dry capsules, with a few exceptions stored in ethanol. During expeditions, live specimens were photographed under a Canon EOS 50D (macro lens 65mm or 90mm) camera by the MNHN team. Live triphorids from Guadeloupe are also marked with asterisks (*) in the plates. At the laboratory, shells were photographed under a Nikon D5000 camera coupled to a stereomicroscope Leica MZ16. Shells were not coated prior to SEM (Scanning Electron Microscope) images in order to preserve their color; the SEM equipment used was a JEOL 6490-LV. In legend captions, shell size refers to length. Descriptions are solely based on the material examined, following procedures and terminology on Fernandes& Pimenta (2015, 2019a). Empty shells and live specimens are respectively indicated in the material examined by ‘sh’ and ‘spm’. Most species are treated in the taxonomic section briefly, with reduced data on synonymic lists and no descriptions or lists of geographic distribution; for those species, see previous studies (e.g. Fernandes& Pimenta 2020) for complete data. abbreviationS ANSP The Academy of Natural Sciences of Drexel University, Philadelphia; BMSM Bailey-Matthews National Shell Museum, Sanibel; FLMNH Florida Museum of Natural History, Gainesville; MNCN Museo Nacional de Ciencias Naturales, Madrid; MNHN Muséum national d’Histoire naturelle, Paris; MNRJ Museu Nacional, Universidade Federal do Rio de Janeiro, Rio de Janeiro; NHMUK Natural History Museum, London; USNM/NMNH National Museum of Natural History, Washington D.C.; ZMA.MOLL Mollusca collection, Naturalis Biodiversity Center, Leiden. ecological analySeS The individual rarefaction curve and the abundance distribution of triphorids from MADIBENTHOS were obtained in the software PAST 4.02, regarding the total number of shells/ specimens, as well as indices of dominance, Shannon’s diversity, Pielou’s equitability and Buzas& Gibson’s evenness (all with 9999 bootstrap replications). The estimative of species richness (i.e., to evaluate the number of missing species) was calculated with a Chao1 estimator (100randomization runs; bias-corrected) in the software EstimateS 9.1. The software Excel was used for simple graphs, such as the number and proportion of species sampled by each sampling method (AB,brushing; AD, dredging; AM, visual at intertidal; AR, visual at diving; AS, suction), following the codes in Bouchet etal. (2019); stations AN (trap lines) were excluded because they did not contain triphorids. Shells identified as Nanaphora cf. verbernei (Moolenbeek& Faber, 1989) were included within N. verbernei in all ecological analyses. At the comparison between species found in Martinique vs other sites in the West Atlantic, only species from the continental shelf were considered, i.e., with at least one record shallower than 150m; the records of Inella triserialis and “Inella” aff. harryleei Rolán& Fernández-Garcés, 2008 respectively from Martinique and Guadeloupe (Lamy& Pointier 2018) were regarded as from the deep-sea, even though the minimum depth was indicated as 55m (see Discussion). In addition, some morphs without formal names were considered (e.g. three undescribed species of Marshallora Bouchet, 1985 in Brazil – Fernandes etal. 2021), but nomina dubia such as Metaxia vicina (C. B. Adams, 1850), recorded for the Aruba-Bonaire-Curaçao Is. (hereafter ABC Is.) (De Jong& Coomans 1988), were ignored. To evaluate whether triphorids from Martinique are endemic or have a wider range, a species is here considered to belong to the entire West Atlantic (and not restricted to the Lesser Antilles or the NW Atlantic) if crossing the Amazon filter. For example, Isotriphora tricingulata Rolán& Fernández-Garcés, 2015 occurs in part of the Caribbean and in Amapá state (North Brazil) (Fernandes& Pimenta 2020), but it is absent below the Amazon mouth, thus it is regarded as a NW Atlantic species. For bathymetric comparisons, only shells not much worn were considered, to avoid bias of post-mortem dislodgement. Shells were assigned to each of five depth zones (0-10m; 1120m; 21-30m; 31-60m; 61-85m). When wide depth ranges are given in a station, the predominant range was considered (e.g. in a depth of 8-16m, thus range 11-20m was selected) or only the minimum depth, if distributed in equal parts (e.g. in a depth of 8-12m, thus range 0-10m was selected). Two ordinations of nMDS (non-metric multidimensional scaling) were calculated in PAST 4.02 for the depth and geographic zones within Martinique (as proposed by Bouchet etal. 2019: 217), one regarding the relative abundance (percentage) of each species per zone (using a Bray-Curtis index after an ‘arcsin square root’ transformation), the other regarding the presence/absence of each species per zone (using a Jaccard index); depth/geographic zones with less than 20 triphorid shells were excluded in order to reduce sampling bias. RESULTS After the analysis of 1615shells/specimens, 33 triphorid species were sampled in the shallow waters of Martinique by the MADIBENTHOS expedition, of which three are described as new species. The taxonomic findings are treated individually in the next section (Figs1-17), also comprising shallow-water 460 ZOOSYSTEMA • 2024 • 46 (18) Fernandes M.R. records from Guadeloupe (mostly KARUBENTHOS 1, with a few records from KARUBENTHOS 2), and live specimens are illustrated further (Figs18-20). The ecological analyses (Figs21-26) are treated in the Discussion. Family triphoridae Gray, 1847 Genus Cheirodonta Marshall, 1983 Cheirodonta dupliniana (Olsson, 1916) (Fig.1A) Triphora dupliniana Olsson, 1916: 138, pl.3 fig.8. material examined.— Guadeloupe. KARUBENTHOS 1 • 1sh [identified by Rolán& Fernández-Garcés (2015): fig.4L as Marshallora modesta]; sta. GS22; MNHN • 1sh [previously identified as Sagenotriphora osclausum]; sta. GB25; MNHN • 1sh [idem]; sta. GB34; MNHN. Martinique. MADIBENTHOS • 1sh; ‘Nord Atlantique’; sta. AD244; MNHN • 1sh; ‘Nord Caraibe’; sta. AM040; MNHN • 2sh; ‘Nord Caraibe’; sta.AS570; MNHN • 1sh; sta. AS576; MNHN. bathymetric diStribution.— Recorded depth in Guadeloupe: 10-45m. Recorded depth in Martinique: 0-59m. Previous recorded depth in the West Atlantic: 6-183m (Fernandes& Pimenta 2020). Genus Coriophora Laseron, 1958 Coriophora novem (Nowell-Usticke, 1969) (Figs1B; 18A, B) Triphora novem Nowell-Usticke, 1969: 12, pl.2 fig.403. material examined.— Guadeloupe. KARUBENTHOS 1 • 1sh; sta. GB06; MNHN • 1sh; sta. GS07; MNHN • 2sh; sta. GB08; MNHN • 1 spm; sta. GM08; MNHN • 1sh; sta. GB09; MNHN • 1sh; sta. GD10; MNHN • 2sh; sta. GR13; MNHN • 5sh; sta. GS13; MNHN • 1sh; sta. GB20; MNHN • 1sh; sta. GS24; MNHN • 1sh; sta. GD25; MNHN • 1sh [juvenile]; sta. GD31; MNHN • 2sh; sta. GD58; MNHN • 1sh; sta. GD59; MNHN.— KARUBENTHOS 2 • 1sh [worn]; sta. DW4545; MNHN. Martinique. MADIBENTHOS. ‘Nord Atlantique’ • 1sh; sta. AB197; MNHN • 1spm; sta. AD254; MNHN • 1sh; sta. AB260; MNHN • 2sh; sta. AD260; MNHN • 3sh; sta. AD261; MNHN • 4sh; sta. AD263; MNHN • 3sh; sta. AD275; MNHN • 3sh; sta. AB350; MNHN • 2sh; sta. AB562; MNHN • 3sh; sta. AS565; MNHN. — ‘Sud Atlantique’ • 2sh; sta. AS081; MNHN • 2sh; sta. AB181; MNHN • 1sh [juvenile]; sta. AB183; MNHN • 1sh; sta. AS255; MNHN • 1sh; sta. AB400; MNHN • 2sh; sta. AB401; MNHN • 1sh; sta. AB405; MNHN • 1sh [worn]; sta. AB419; MNHN.—‘Nord Caraibe’ • 1sh; sta. AB161; MNHN • 2sh; sta. AD280; MNHN • 4sh; sta. AD290; MNHN • 1sh; sta. AB362; MNHN • 1sh; sta. AB460; MNHN • 2sh; sta. AS574; MNHN • 2sh; sta. AS576; MNHN. ‘Sud Caraibe’ • 3sh; sta. AB058; MNHN • 1sh; sta. AB060; MNHN • 4sh; sta. AS071; MNHN • 1sh; sta. AB123; MNHN • 1sh [juvenile]; sta. AB155; MNHN • 1sh; sta. AB169; MNHN • 1sh [juvenile]; sta. AD218; MNHN • 1sh [juvenile]; sta. AB358; MNHN • 2sh; sta. AB369; MNHN • 1sh; sta. AD616; MNHN.— ‘Baie de Fort-de-France’ • 1sh; sta. AB398; MNHN. bathymetric diStribution.— Recorded depth in Guadeloupe: 1-160m (live specimen at 1m). Recorded depth in Martinique: 4-80m (live specimen at 62m). Previous recorded depth in the West Atlantic: 2-110m (Fernandes& Pimenta 2020). Genus Cosmotriphora Olsson& Harbison, 1953 Cosmotriphora arnoldoi Faber& Moolenbeek, 1991 (Fig.1C) Cosmotriphora arnoldoi Faber& Moolenbeek, 1991: 81, figs1-2. material examined.— Martinique. MADIBENTHOS. ‘Nord Atlantique’ • 1sh; sta. AD260; MNHN. bathymetric diStribution.— Recorded depth in Martinique: 78m. Previous recorded depth in the West Atlantic: 20-155m (Fernandes& Pimenta 2020). Cosmotriphora melanura (C. B. Adams, 1850) (Figs1D; 18C-F) Cerithium melanura C. B. Adams, 1850: 117. material examined.— Guadeloupe. KARUBENTHOS 1 • 1sh; sta. GB01; MNHN • 3sh, 1 spm; sta. GB02; MNHN • 1sh, 1spm; sta. GB03; MNHN • 3sh; sta. GS04; MNHN • 2sh; sta. GS05; MNHN • 2sh, 2 spm; sta. GB06; MNHN • 4sh; sta. GS06; MNHN • 2sh; sta. GS07; MNHN • 2sh; sta. GB08; MNHN • 2sh [juveniles]; sta. GM08; MNHN • 11sh; sta. GS08; MNHN • 8sh; sta. GB09; MNHN • 3sh; sta. GS09; MNHN • 3sh; sta. GD10; MNHN • 3sh; sta. GR10; MNHN • 14sh; sta. GB11; MNHN • 1sh; sta. GS11; MNHN • 1sh; sta. GB12; MNHN • 1sh; sta. GS12; MNHN • 6; sta. GB13; MNHN • 25sh; sta. GS13; MNHN • 1sh; sta. GB14; MNHN • 1sh; sta. GR14; MNHN • 2sh; sta. GS14; MNHN • 1sh; sta. GB15; MNHN • 1sh; sta. GS15; MNHN • 13sh; sta. GB16; MNHN • 1sh; sta. GS16; MNHN • 3 spm; sta. GR17; MNHN • 3sh, 1 spm; sta. GS17; MNHN • 2sh; sta. GB18; MNHN • 3sh; sta. GS18; MNHN • 1sh; sta. GB19; MNHN • 3sh; sta. GR19; MNHN • 6sh, 1 spm; sta. GB20; MNHN • 1sh; sta. GD20; MNHN • 5sh; sta. GS20; MNHN • 1sh; sta. GD21; MNHN • 1sh; sta. GR21; MNHN • 4sh; sta. GB22; MNHN • 2sh; sta. GS22; MNHN • 1sh; sta. GB23; MNHN • 2sh; sta. GS23; MNHN • 1sh; sta. GB24; MNHN • 1sh; sta. GD24; MNHN • 2sh; sta. GS24; MNHN • 2sh; sta. GS25; MNHN • 1sh; sta. GB26; MNHN • 5sh; sta. GS29; MNHN • 2sh; sta. GB30; MNHN • 10sh; sta. GB31; MNHN • 1sh; sta. GD31; MNHN • 1sh; sta. GS32; MNHN • 1sh; sta. GD33; MNHN • 3sh; sta. GB34; MNHN • 4sh; sta. GS34; MNHN • 1sh; sta. GB36; MNHN • 3sh; sta. GR36; MNHN • 4sh; sta. GS36; MNHN • 1sh; sta. GS37; MNHN • 1sh; sta. GD52; MNHN • 1sh; sta. GD55; MNHN • 2sh; sta. GD58; MNHN • 1sh; sta. GD59; MNHN • 1sh; sta. GD69; MNHN. — KARUBENTHOS 2 • 9sh; sta. DW4545; MNHN • 1sh; sta. DW4546; MNHN • 1sh; sta. DW4550; MNHN • 1sh; sta. DW4555; MNHN • 1sh; sta. DW4586; MNHN • 1sh [juvenile]; sta. DW4587; MNHN • 1sh [juvenile]; sta. DW4599; MNHN. Martinique. MADIBENTHOS. ‘Nord Atlantique’ • 4sh; sta. AB197; MNHN • 2sh; sta. AD244; MNHN • 2sh; sta. AD248; MNHN • 1sh [juvenile]; sta. AD250; MNHN • 2sh; sta. AD254; MNHN • 2sh; sta. AD257; MNHN • 1sh; sta. AB260; MNHN • 7sh; sta. AD260; MNHN • 7sh; sta. AD261; MNHN • 10sh; sta. AD263; MNHN • 7sh; sta. AD275; MNHN • 1sh; sta. AB350; MNHN • 1sh; sta. AS557; MNHN • 3sh; sta. AB562; MNHN • 3sh; sta. AS563; MNHN • 13sh; sta. AS565; MNHN. — ‘Sud Atlantique’ • 4sh; sta. AS078; MNHN • 1sh; sta. AS081; MNHN • 6sh; sta. AB181; MNHN • 2sh; sta. AB189; MNHN • 2sh; sta. AB193; MNHN • 1sh; sta. AD222; MNHN • 2sh; sta. AD224; MNHN • 1sh; sta. AD232; MNHN. — ‘Nord Caraibe’. 2sh; sta. AS054; MNHN • 1sh; sta. AB062; MNHN • 2sh; sta. AB108; MNHN • 1sh; sta. AD109; MNHN • 6sh; sta. AB159; MNHN 461 Triphoridae from Martinique ZOOSYSTEMA • 2024 • 46 (18) • 3sh; sta. AB183; MNHN • 2sh; sta. AD276; MNHN • 3sh; sta. AD280; MNHN • 2sh; sta. AD289; MNHN • 12sh; sta. AD290; MNHN • 3sh; sta. AD298; MNHN • 1sh; sta. AD299; MNHN • 2sh; sta. AR308; MNHN • 3sh; sta. AS363; MNHN • 2sh; sta. AB372; MNHN • 1sh; sta. AS373; MNHN • 7sh; sta. AS375; MNHN • 1sh; sta. AB377; MNHN • 1sh; sta. AS378; MNHN • 3sh; sta. AB386; MNHN • 1sh; sta. AB460; MNHN • 5sh; sta. AB463; MNHN • 2sh; sta. AB567; MNHN • 3sh; sta. AS570; MNHN • 2sh; sta. AS572; MNHN • 3sh; sta. AS574; MNHN • 2sh; sta. AS576; MNHN • 3sh; sta. AD612; MNHN. ‘Sud Caraibe’ ABCD E FGHI J Fig. 1. — Triphoridae Gray, 1847 from Martinique: A, Cheirodonta dupliniana (Olsson, 1916), MNHN, sta. AS576, 5.4 mm; B, Coriophora novem (Nowell-Usticke, 1969), MNHN, sta. AD280, 5.0 mm; C, Cosmotriphora arnoldoi Faber & Moolenbeek, 1991, MNHN, sta. AD260, 2.6 mm; D, Cosmotriphora melanura (C. B. Adams, 1850), MNHN, sta. AB123, 4.1 mm; E, Eutriphora bermudensis (Bartsch, 1911), MNHN, sta. AS257, 4.2 mm; F, Iniforis gudeliae Rolán & Fernández-Garcés, 2009, MNHN, sta. AS403, 4.7 mm; G, Iniforis pseudothomae Rolán & Fernández-Garcés, 1993, MNHN, sta. AB062, 7.0 mm; H, Iniforis turristhomae (Holten, 1802), MNHN, sta. AS075, 5.7 mm; I, Isotriphora tricingulata Rolán & Fernández-Garcés, 2015, MNHN, sta. AD263, 4.9 mm; J, Isotriphora sp., MNHN, sta. AS403, 2.6 mm. Scale bars: 1 mm. 462 ZOOSYSTEMA • 2024 • 46 (18) Fernandes M.R. • 2sh; sta. AS057; MNHN • 1sh; sta. AB058; MNHN • 1sh; sta. AD067; MNHN • 3sh; sta. AS068; MNHN • 4sh; sta. AS071; MNHN • 2sh; sta. AS075; MNHN • 3sh; sta. AS092; MNHN • 1spm; sta. AS096; MNHN • 1sh, 1spm; sta. AB123; MNHN • 6sh; sta. AB126; MNHN • 2sh; sta. AB150; MNHN • 5sh; sta. AB152; MNHN • 1sh; sta. AS154; MNHN • 1sh; sta. AB163; MNHN • 4sh; sta. AB169; MNHN • 1sh; sta. AB173; MNHN • 1sh; sta. AB177; MNHN • 1sh, 1spm; sta. AD214; MNHN • 3sh; sta. AD216; MNHN • 5sh; sta. AD218; MNHN • 1sh; sta. AD220; MNHN • 1sh; sta. AD271; MNHN • 3sh; sta. AB354; MNHN • 1sh [juvenile]; sta. AB358; MNHN • 1sh; sta. AR359; MNHN • 1sh; sta. AB360; MNHN • 2sh; sta. AB369; MNHN • 1sh; sta. AS370; MNHN • 1sh; sta. AB578; MNHN • 2sh; sta. AS579; MNHN • 2sh; sta. AD615; MNHN • 1sh; sta. AD616; MNHN. — ‘Baie de Fort-de-France’ • 1sh; sta. AB392; MNHN • 1sh; sta. AB398; MNHN. bathymetric diStribution.— Recorded depth in Guadeloupe: 0-150m (live specimens at 3-23m); some intact adult shells obtained by KARUBENTHOS 2 between 204-482m, but they must be regarded with caution. Recorded depth in Martinique: 0-80m (live specimens at 4-70m). Previous recorded depth in the West Atlantic: 0-160m, with discrepant records down to 480m (Fernandes& Pimenta 2020). Genus Eutriphora Cotton& Godfrey, 1931 Eutriphora bermudensis (Bartsch, 1911) (Fig.1E) Triphoris bermudensis Bartsch, 1911: 305, pl.28, figs2, 4. material examined.— Guadeloupe. KARUBENTHOS 1 • 1sh [previously identified as Iniforis turristhomae]; sta. GS05; MNHN • 1sh [idem]; sta. GS06; MNHN • 1sh [idem]; sta. GB08; MNHN • 1sh [idem]; sta. GS18; MNHN • 1sh [idem]; sta. GB26; MNHN. Martinique. MADIBENTHOS. ‘Sud Atlantique’ • 1sh; sta. AS253; MNHN • 1sh; sta. AS257; MNHN. — ‘Nord Caraibe’ • 1sh; sta. AB161; MNHN. ‘Sud Caraibe’: • 1sh; sta. AB169; MNHN. — ‘Baie de Fort-de-France’ • 1sh; sta. AB392; MNHN. bathymetric diStribution.— Recorded depth in Guadeloupe: 17-49m. Recorded depth in Martinique: 13-50m. Previous recorded depth in the West Atlantic: 4-56m (Rolán& Fernández-Garcés 1995). remark The single shell identified by Rolán& Fernández-Garcés (2015: fig.4T) as E. bermudensis from Guadeloupe, and followed by Lamy& Pointier (2018: pl.91, fig.6), actually refers to Triphora martii Rolán& Fernández-Garcés, 1995. Genus Inella Bayle, 1879 Inella spinosa n.sp. (Fig.4) urn:lsid:zoobank.org:act:42468F50-0E06-472E-8F1D-5738EF4F29B5 t ype material . — Holotype. Martinique. MADIBENTHOS • sh; Macouba, region ‘Nord Atlantique’; sta. AD275, 14°55’00”N, 61°08’54”W; depth 80m; MNHN-IM-2000-38595. Paratypes. Martinique. MADIBENTHOS. ‘Nord Atlantique’ • 2sh; sta. AD263; MNHN-IM-2000-38596. type locality.— Martinique: Macouba, region ‘Nord Atlantique’; MADIBENTHOS expedition sta. AD275, 14°55’00”N, 61°08’54”W; 80m. other material examined.— Guadeloupe. KARUBENTHOS 2 • 1sh; sta. DW4545; MNHN. Martinique. MADIBENTHOS. ‘Nord Atlantique’ • 1sh [juvenile, worn]; sta. AD260; MNHN. etymology.— From Latin, spinosus. The species is named due to the spiny appearance of the spiral cords of teleoconch. diagnoSiS.— Spirally-elongated, pointed nodules in median and abapical cords, giving a spiny appearance to the teleoconch profile; adapical spiral cord initially very narrow, never reaching the same size of other cords; shell with a cream background. geographic diStribution.— Guadeloupe and Martinique. bathymetric diStribution.— Recorded depth in Guadeloupe: 60-82m. Recorded depth in Martinique: 78-80m. deScription Shell sinistral, conical/fusiform, rectilinear profile, up to 4.4mm long, 1.1mm wide, length/width ratio 3.5-3.9; adult shells reach at least 2.7mm in length. Protoconch paucispiral with truncated apex, without clear differentiation from the teleoconch, of 2.75-3.0 whorls, 0.57-0.61mm long, 0.50-0.52mm wide; the small nucleus rises in an adapical direction and further goes abapical in an oblique descent; two smooth, keeled spiral cords of equal size, situated at 26-34% and 66-71% of last whorl height, in addition to a narrow and smooth subsutural cord. Teleoconch with up to 8.5 whorls; two main spiral cords (median and abapical) at the beginning, continuous to those of protoconch, assuming a keeled-shape along the teleoconch, with median cord always more prominent, except on the body whorl where both cords have nearly the same size; adapical spiral cord initially very narrow, gradually strengthening along the teleoconch but never reaching the same size of other cords or forming distinct nodules; a smooth sutural cord is also evident, gradually strengthening and reaching the same size of adapical cord in larger shells; spirallyelongated, pointed nodules in median and abapical cords, giving a spiny appearance to the teleoconch profile; nine to ten orthocline axial ribs on seventh teleoconch whorl; suture distinct; smooth subperipheral cord, one narrow, smooth basal cord; supranumerical cords absent; elliptical aperture, 0.51-0.67mm long, 0.37-0.44mm wide, length/width ratio 1.4-1.5; open, very short anterior canal, 0.05-0.12mm long, 0.18-0.27mm wide, length/width ratio 0.3-0.5; posterior canal absent. Shell with a cream background, white nodules on teleoconch, internodular spaces light brown. remarkS The spiny appearance of the spiral cords of the teleoconch, combined with the color pattern and small shell size (reaching only 4.4mm long), makes I. spinosa n.sp. unique among Inella and other West Atlantic triphorids. Because of its 463 Triphoridae from Martinique ZOOSYSTEMA • 2024 • 46 (18) probable lecithotrophic development and bathymetric range restricted to the upper 100m (absent from deeper waters of Guadeloupe, based on samples from KARUBENTHOS 2), I.spinosa n.sp. probably has a narrow geographic range in the Lesser Antilles. The single shell obtained from Guadeloupe (Fig.4D) has the axial sculpture much weaker than shells from Martinique (Fig.4A-C), attenuating the spiny appearance of the spiral cords. ABCDE FGHI J Fig. 2 . — Triphoridae Gray, 1847 from Martinique - cont.: A, Latitriphora albida (A. Adams, 1854), MNHN, sta. AB149, 4.0 mm; B, Marshallora cf. modesta (C. B. Adams, 1850), MNHN, sta. AB150, 4.3 mm; C, Metaxia excelsa Faber & Moolenbeek, 1991, MNHN, sta. AB123, 3.6 mm; D, Metaxia rugulosa (C. B. Adams, 1850), MNHN, sta. AD265, 6.3 mm; E, Metaxia taeniolata (Dall, 1889), MNHN, sta. AD261, 5.5 mm; F, Monophorus olivaceus (Dall, 1889), MNHN, sta. AS576, 9.5 mm; G, Nototriphora decorata (C. B. Adams, 1850), MNHN, sta. AB390, 6.8 mm; H, Sagenotriphora osclausum (Rolán & Fernández-Garcés, 1995), MNHN, sta. AS565, 3.4 mm; I, Similiphora intermedia (C. B. Adams, 1850), MNHN, sta. AD067, 5.6 mm; J, Similiphora sp., MNHN, sta. AS552, 5.1 mm. Scale bars: 1 mm. 470 ZOOSYSTEMA • 2024 • 46 (18) Fernandes M.R. remarkS Lamy& Pointier (2018: pl.92, fig.16) illustrated the holotype of I. tricingulata under the name Triphorasp. 5, providing a depth record of 50m, contrary to the true depth record of the type locality (i.e., 85m in station GD31 of KARUBENTHOS 1, not 81m, as recorded in the original description). I disregard the depth record of 50m provided by Lamy& Pointier (2018), due to the uncertainty of which shells were studied. Isotriphorasp. (Fig.1J) m aterial examined .— Martinique. MADIBENTHOS. ‘Sud Atlantique’ • 1sh; sta. AS403; MNHN. bathymetric diStribution.— Only known from 11m. Genus Latitriphora Marshall, 1983 Latitriphora albida (A. Adams, 1854) (Fig.2A) Triphoris albidus A. Adams, 1854: 278. material examined.— Guadeloupe. KARUBENTHOS 1 • 1sh; sta. GS17; MNHN • 1sh; sta. GB36; MNHN. Martinique. MADIBENTHOS. ‘Nord Atlantique’ • 2sh; sta. AB562; MNHN. — ‘Sud Atlantique’ • 1sh; sta. AB117; MNHN • 2sh; sta. AB149; MNHN. — ‘Nord Caraibe’ • 1sh, 1spm; sta. AB386; MNHN. ‘Sud Caraibe’ • 1sh [worn]; sta. AM033; MNHN • 1sh; sta. AS075; MNHN • 1sh [juvenile]; sta. AB126; MNHN • 1sh; sta. AD216; MNHN. ‘Baie de Fort-de-France’ • 1sh; sta. AB390; MNHN. bathymetric diStribution.— Recorded depth in Guadeloupe: 3-16m. Recorded depth in Martinique: 2-26m (live specimen at 17m). Previous recorded depth in the West Atlantic: 0-107m (Fernandes& Pimenta 2020). remark Rolán& Fernández-Garcés (2015: fig.4S) illustrated a shell of Nototriphora decorata (C. B. Adams, 1850) under the name L. albida, which was followed by Lamy& Pointier (2018: pl.91, fig.15), and there were other 14 MNHN lots of N. decorata from Guadeloupe previously identified as L. albida; only two shells of L. albida were correctly identified. Genus Marshallora Bouchet, 1985 Marshallora cf. modesta (C. B. Adams, 1850) (Fig.2B) Cerithium modestum C. B. Adams, 1850: 117. material examined.— Guadeloupe. KARUBENTHOS 1 • 2spm; sta. GM01; MNHN • 1sh, 3 spm; sta. GM02; MNHN • 2spm; sta. GM03; MNHN • 2sh; sta. GM11; MNHN • 1sh; sta. GM29; MNHN • 1sh; sta. GS30; MNHN • 3sh; sta. GD39; MNHN • 1sh; sta. GD49; MNHN • 1sh; sta. GD51; MNHN. — Martinique. MADIBENTHOS. ‘Sud Atlantique’ • 1sh; sta. AM042; MNHN • 2sh [one with hermit crab]; sta. AM043; MNHN • 1sh; sta. AM325; MNHN. ‘Sud Caraibe’ • 1sh; sta. AB150; MNHN. ‘Baie de Fort-de-France’ • 1sh; sta. AM034; MNHN • 8sh; sta. AB390; MNHN • 1sh; sta. AB392; MNHN • 3sh; sta. AB394; MNHN • 1sh; sta. AD603; MNHN • 1sh; sta. AD607; MNHN • 1sh; sta. AD623; MNHN. bathymetric diStribution.— Recorded depth in Guadeloupe: 0-15m (live specimens at 1m). Recorded depth in Martinique: 0-13m. Previous recorded depth in the West Atlantic: 0-20m (Rosenberg 2009). remarkS As abovementioned, the illustration of M. modesta from Gua - deloupe by Rolán& Fernández-Garcés (2015: fig.4L) actually refers to C. dupliniana, whereas their record of Marshallora nigrocincta (C. B. Adams, 1839) must be refused, because this species is known with certainty only from Canada and USA, based on DNA evidences and some shell features (Fernandes etal. 2021). Accordingly, the record of M. nigrocincta from Guadeloupe and Martinique by Lamy& Pointier (2018) is rejected. The material from the French Antilles will be studied under an integrative taxonomic basis, together with new samples from the West Atlantic (unpublished data). Meanwhile, shells are herein identified as M. cf. modesta, regarding the existence of at least two Caribbean species in this complex (Fernandes etal. 2021). Marshallora monteiroi (Rolán& Fernández-Garcés, 2015) n. comb. (Figs7; 8; 20I-J) “Triphora” monteiroi Rolán& Fernández-Garcés, 2015: 47, pl.3. Triphora portoricensis non Rolán& Redfern, 2008 – Lamy& Pointier 2018: 289, pl.92, fig.11. type material. — Holotype. Guadeloupe • sh; Basse-Terre, Baie de Baille-Argent; KARUBENTHOS 1 sta. GD21, 16°15’33”N, 61°48’48”W; depth 40m; MNHN-IM-2000-30473. type locality.— Guadeloupe: Basse-Terre, Baie de Baille-Argent; KARUBENTHOS 1 sta. GD21, 16°15’33”N, 61°48’48”W; 40m. material examined.— Guadeloupe. KARUBENTHOS 1 • 1sh [previously identified as Marshallora modesta – this shell is from the type locality]; sta. GD21; MNHN. Martinique. MADIBENTHOS. ‘Nord Atlantique’ • 1sh; sta. AD263; MNHN • 1sh; sta. AD275; MNHN • 1sh; sta. AB350; MNHN. — ‘Sud Atlantique’ • 1sh; sta. AS252; MNHN. — ‘Nord Caraibe’ • 10sh, 1spm; sta. AD283; MNHN • 1sh; sta. AD290; MNHN • 1sh; sta. AS482; MNHN. bathymetric diStribution.— Recorded depth in Guadeloupe: 40m (including a live specimen). Recorded depth in Martinique: 10-80m (live specimen at 37-40m). geographic diStribution.— Guadeloupe and Martinique. emended deScription Shell sinistral, conical to slightly pupoid, rectilinear to slightly convex profile, up to 4.6mm long, 1.2mm wide, length/width 471 Triphoridae from Martinique ZOOSYSTEMA • 2024 • 46 (18) ratio 2.4-3.9; adult shells reach at least 2.0mm in length. Protoconch sub-columnar, slightly inflated, of 2.75-3.0 whorls, 0.44-0.51mm long, 0.41-0.48mm wide, weak distinction between protoconch and teleoconch; first smooth, globose, but on subsequent whorls axial ribs soon appearing, with their adapical part emerging on the abapical part of the smooth whorl above, assuming an irregular shape, often considerably sigmoid on second whorl, but usually slightly sigmoid or nearly orthocline (sometimes opisthocline) towards the end of protoconch, where it numbers 20-22 ribs; one main spiral cord emerging soon after the axial ribs, keel-shaped and often smooth (but wrinkled in some shells), situated at 47-60% of last whorl height. Teleoconch with up to nine whorls; two spiral cords (adapical and abapical) at the beginning, the abapical one continuous to that of protoconch; median spiral cord emerging at the beginning of second to the end of third whorl, reaching the same size than other cords after one to two whorls; at the body whorl, the three cords are equidistant and have nearly the same size; 22-24 nearly orthocline axial ribs on the sixth whorl; rounded nodules of medium size; distinct suture, with a sutural cord; slightly nodulose to smooth subperipheral cord, two smooth, well-developed basal cords; no supranumerical cords; aperture ovate (with a minute notch in the posterior end), 0.50-0.81mm long, 0.40-0.61mm wide, length/width ratio 1.3-1.5; anterior canal short, open, 0.13-0.25mm long, 0.14-0.28mm wide, length/width ratio 0.8-0.9. Protoconch brownish, cream, golden or white; teleoconch dark to light brown, rarely beige; head-foot white, translucent. remarkS This species was previously known only from the holotype, which has a light brown teleoconch and white protoconch (Rolán& Fernández-Garcés 2015). An additional shell from the type locality in Guadeloupe is illustrated (Fig.7J), with a darker teleoconch, which is the common pattern in shells of Marshallora monteiroi n. comb. from Martinique (Fig.7). An exception is a shell with a gradual (not abrupt) color transition between the lighter protoconch and the cream/beige teleoconch (Fig.7K), which is more similar to the planktotrophic species Marshallora ostenta Rolán& Fernández-Garcés, 2008, and it perhaps does not belong to M. monteiroi n. comb. The color of the protoconch is variable among shells of M. monteiroi n. comb., from white to brownish (Fig.7). There are also slight variations in the protoconch sculpture, with a spiral cord usually smooth and situated at the mid portion of the whorl, but sometimes wrinkled or situated at a lower portion, in addition to the variable axial ribs (Fig.8). Remarkably, the adult shell length ranges from 2.0 to 4.6mm. AB CD EF GH Fig. 8. — Marshallora monteiroi (Rolán & Fernández-Garcés, 2015) n. comb.: A, MNHN, sta. AD283 (Fig. 7A); B, MNHN, sta. AS482 (Fig. 7B); C, MNHN, sta. B350 (Fig. 7C); D, MNHN, sta. AD275 (Fig. 7D); E, MNHN, sta. AD283 (Fig. 7E); F, MNHN, sta. AD283 (Fig. 7F); H, MNHN, sta. AS252 (Fig. 7I); H, MNHN, sta. AD263 (Fig. 7K). Scale bars: 100 μm. 472 ZOOSYSTEMA • 2024 • 46 (18) Fernandes M.R. This species is placed under the Atlantic genus Marshallora owing to the brown teleoconch, smooth basal cords (Fig.7L-M), absence of supranumerical cords (Fig.7N) and white head-foot (Fig.20I-J), among other features. There are other Caribbean species with paucispiral protoconchs currently allocated under the catch-all taxon “Triphora” that, otherwise, fit into the concept of Marshallora, following the above-mentioned features [including the white head-foot of one species – Redfern (2013: fig.356B)]. These species are herein transferred to Marshallora, pending future anatomical and genetic studies to test these allocations: Marshallora abacoensis (Rolán& Lee, 2008) n. comb., Marshallora armandoi (Espinosa& Ortea, 2020) n. comb., Marshallora calva (Faber& Moolenbeek, 1991) [already cited in this genus by Fernandes& Pimenta 2020] and Marshallora grenadensis (Rolán& Lee, 2008) n.comb. The latter, only known from the holotype, is the most similar species to M. monteiroi n.comb., regarding slight differences in the color and sculpture of the protoconch (Rolán& Fernández-Garcés 2015), and possibly having a less inflated protoconch and slightly later emergence of the median spiral cord (emerging narrowly at the end of fourth teleoconch whorl – Rolán& Fernández-Garcés 2008: fig.30D), requiring additional material for further comparisons. Genus Metaxia Monterosato, 1884 Metaxia discus n.sp. (Fig.9A-F) urn:lsid:zoobank.org:act:B613849C-897F-4E0A-BE99-EBD074B89222 type material. — Holotype. Martinique. • sh; North of Presqu’Île de la Caravelle, region ‘Nord Atlantique’; MADIBENTHOS expedition; sta. AD245; 14°53’06”N, 60°3’24”W; depth 60m; MNHN-IM-2000-38601. Paratype. Martinique. MADIBENTHOS • 1sh; type locality; MNHN-IM-2000-38602. type locality.— Martinique: North of Presqu’Île de la Caravelle, region ‘Nord Atlantique’; MADIBENTHOS expedition sta. AD245, 14°53’06”N, 60°3’24”W; 60m. etymology.— From Latin, discus. The species is named due to the discoid shape of the protoconch. diagnoSiS.— Protoconch with 2.5much convex whorls, first whorl discoid, broad, with same width than second one, and with two spiral cords of varying expression; shell white. geographic diStribution.— Martinique (this study). bathymetric diStribution.— Only known from 60m. deScription Shell dextral, conical/fusiform, rectilinear profile, up to 3.2mm long, 0.8mm wide, length/width ratio 3.5-4.1. Protoconch paucispiral, of 2.5much convex whorls, 0.41-0.44mm long, 0.34-0.37mm wide; first whorl discoid, broad, with same width than second one, with two spiral cords on the adapical half of the whorl, having varying expression, the abapical cord more promi - nent and initially undulating (zigzag) to further nearly straight, and the adapical cord undulating (zigzag); on the second whorl, the two spiral cords having nearly the same width and being no longer restricted to the adapical portion of the whorl, situated respectively at c.39% and c.71% of whorl height, with small nodules, with incomplete, orthocline to slightly prosocline axial riblets located above adapical cord. Teleoconch with up to seven whorls; four equidistant spiral cords, with the first (adapical) cord being much smaller and the third cord the most prominent, in addition to a much reduced, smooth sutural cord; elliptical nodules of small size; c.15 weak, orthocline axial ribs on the sixth teleoconch whorl; smooth and narrow subperipheral cord, no distinct basal cord; aperture rounded (but partially broken), with small anterior notch. Shell white. remarkS Metaxia discus n.sp. superficially resembles the morph named Metaxiasp. 1 (Fig.9G-H), from Barbados (Rolán& Fernández-Garcés 2008). The teleoconch of Metaxiasp. 1has three main spiral cords, similarly to Metaxia excelsa (Faber& Molenbeek, 1991) or Metaxia taeniolata (Dall, 1889), whereas M. discus n.sp. has four main spirals in late whorls (i.e., the upper cord is stronger than that of Metaxiasp. 1, which seems a weak subsutural cord), similarly to Metaxia rugulosa (C. B. Adams, 1850). The protoconch of Metaxiasp. 1 is illustrated here (Fig.9H), and, despite slightly worn, is more similar to M. rugulosa than to M. discus n.sp. (see comparisons below). Despite having a nearly identical teleoconch, M. discus n.sp. is differentiated from the widespread West Atlantic species M. rugulosa by the discoid and truncated shape of the protoconch, often with undulating spiral cords (Fig.9D), whereas the protoconch of M. rugulosa is always convex, with a dome-shaped first whorl and straight spiral cords in the second whorl (Rolán& Redfern 1996; Fernandes& Pimenta 2011). Metaxia discus n.sp. has a protoconch similar to Metaxia quadrata Faber, 2010, which is known only by the holotype, from Aruba. However, the protoconch of M. quadrata seems to have three spiral cords on the first whorl (vs two cords in M. discus n.sp.), with a more accentuated zigzag pattern than in M. discus n.sp., although the second whorl of protoconch has a nearly rectilinear adapical spiral cord (but more undulating in M. discus n.sp.). In addition, the shell of M. quadrata is yellowish brown and it has a weaker axial sculpture (Faber 2010) vs entirely white in M. discus n.sp. Metaxia excelsa Faber& Moolenbeek, 1991 (Figs2C; 18J, L-O) Metaxia excelsa Faber & Moolenbeek, 1991: 83. material examined.— Guadeloupe. KARUBENTHOS 1 • 2sh; sta. GB02; MNHN • 1spm; sta. GB03; MNHN • 3sh; sta. GB06; MNHN • 1sh, 1spm; sta. GS06; MNHN • 2sh, 1spm; sta. GB08; MNHN • 1sh; sta. GB11; MNHN • 1sh; sta. GS11; MNHN • 1spm; sta. GB12; MNHN • 1sh; sta. GB13; MNHN • 15sh, 3spm; sta. GS13; MNHN • 1sh; sta. GB14; MNHN • 1spm; sta. GS15; MNHN • 4sh; sta. GB16; MNHN • 1sh; sta. GS17; MNHN • 4sh; sta. GS18; MNHN • 1sh; sta. GS19; MNHN • 1sh; sta. GS22; MNHN • 1sh; sta. GB23; MNHN • 473 Triphoridae from Martinique ZOOSYSTEMA • 2024 • 46 (18) 1sh; sta. GS23; MNHN • 1sh; sta. GB25; MNHN • 1sh; sta. GB36; MNHN • 1sh; sta. GD68; MNHN. — KARUBENTHOS 2 • 1sh; sta. DW4545; MNHN. Martinique. MADIBENTHOS. ‘Nord Atlantique’ • 1spm; sta. AB197; MNHN • 1sh [worn]; sta. AD255; MNHN • 2sh; sta. AD260; MNHN • 2sh; sta. AD263; MNHN • 3sh; sta. AD267; MNHN • 5sh; sta. AD275; MNHN • 2sh; sta. AS565; MNHN. — ‘Sud Atlantique’ • 1sh; sta. AD222; MNHN • 1sh; sta. AB401; MNHN. — ‘Nord Caraibe’ • 1sh; sta. AB159; MNHN • 3sh, 1spm; sta. AB161; MNHN • 2sh; sta. AD280; MNHN • 1sh; sta. AD289; MNHN • 3sh; sta. AD290; MNHN • 1spm; sta. AB362; MNHN • 1sh; sta. AS363; MNHN • 3sh; sta. AB372; MNHN • 1sh; sta. AS375; MNHN • 1sh; sta. AB382; MNHN • 4sh; sta. AB386; MNHN • 1sh; sta. AB463; MNHN • 1sh; sta. AB567; MNHN • 1sh; sta. AS572; MNHN • 3sh; sta. AS576; MNHN. ‘Sud Caraibe’ • 1sh; sta. AB058; MNHN • 1spm; sta. AB123; MNHN AB CDE F GH Fig. 9 . — A-F, Metaxia discus n. sp. A, B, D, F, MNHN-IM-2000-38601, holotype, 3.2 mm; C, E, MNHN-IM-2000-38602, paratype, sta. AD245, 2.7 mm; G, H, Metaxia sp. 1 (morph named by Rolán & Fernández-Garcés [2008]), USNM 87301, 4.2 mm (vs 5.0 mm, according to Rolán & Fernández-Garcés [2008]); credits: Yolanda Villacampa (NMNH). Scale bars: A-C, G, 1 mm; D, E, 100 μm; F, H, 200 μm. 474 ZOOSYSTEMA • 2024 • 46 (18) Fernandes M.R. • 2sh; sta. AB150; MNHN • 2sh; sta. AB152; MNHN • 2sh; sta. AB169; MNHN • 1sh; sta. AB173; MNHN • 2sh; sta. AB177; MNHN • 1sh; sta. AD203; MNHN • 2sh; sta. AD216; MNHN • 1sh [juvenile]; sta. AD271; MNHN • 1sh; sta. AB354; MNHN • 1sh; sta. AS370; MNHN • 1sh; sta. AD617; MNHN. ‘Baie de Fortde-France’ • 1sh; sta. AB392; MNHN • 1sh; sta. AB394; MNHN. bathymetric diStribution.— Recorded depth in Guadeloupe: 3-60m (live specimens at 14-50m). Recorded depth in Martinique: 2-80m (live specimens at 4-25m). Previous recorded depth in the West Atlantic: 1-147m (Fernandes& Pimenta 2020). Metaxia rugulosa (C. B. Adams, 1850) (Fig.2D) Cerithium rugulosum C. B. Adams, 1850: 121-122. material examined.— Guadeloupe. KARUBENTHOS 1 • 1sh; sta. GS13; MNHN • 1sh; sta. GB14; MNHN • 1sh; sta. GB30; MNHN. — KARUBENTHOS 2 • 1sh; sta. DW4586; MNHN. Martinique. MADIBENTHOS. ‘Nord Atlantique’ • 1sh; sta. AD244; MNHN • 1sh; sta. AD260; MNHN • 2sh; sta. AD261; MNHN • 2sh; sta. AD263; MNHN • 1sh; sta. AD265; MNHN • 4sh; sta. AD275; MNHN. — ‘Sud Atlantique’ • 1sh; sta. AB120; MNHN • 1sh; sta. AB185; MNHN. — ‘Nord Caraibe’ • 1sh; sta. AD277; MNHN • 1sh; sta. AD280; MNHN • 1sh; sta. AS363; MNHN • 1sh; sta. AS576; MNHN. ‘Sud Caraibe’ • 1sh; sta. AD067; MNHN • 1sh; sta. AB150; MNHN • 1sh; sta. AD218; MNHN. ‘Baie de Fort-de-France’ • 1sh; sta. AB398; MNHN. bathymetric diStribution.— Recorded depth in Guadeloupe: 16-50m; one intact adult shell obtained by KARUBENTHOS 2 between 204-251m, but it must be regarded with caution. Recorded depth in Martinique: 2-80m. Previous recorded depth in the West Atlantic: 0-198m (Fernandes& Pimenta 2020). remarkS The shell from Guadeloupe illustrated by Rolán& FernándezGarcés (2015: fig.4G) and Lamy& Pointier (2018: pl.92, fig.3) as M. rugulosa actually refers to M. taeniolata (Dall, 1889). Metaxia taeniolata (Dall, 1889) (Figs2E; 19A, B) Cerithiopsis metaxae var. taeniolata Dall, 1889: 256. material examined.— Guadeloupe. KARUBENTHOS 1 • 1sh [previously identified as M. rugulosa]; sta. GB08; MNHN • 1sh [idem]; sta. GD10; MNHN • 1sh [idem]; sta. GD31; MNHN. Martinique. MADIBENTHOS. ‘Nord Atlantique’ • 1spm; sta. AD261; MNHN • 1sh; sta. AD275; MNHN • 1sh; sta. AS552; MNHN. — ‘Nord Caraibe’ • 1sh; sta. AS572; MNHN. bathymetric diStribution.— Recorded depth in Guadeloupe: 17-85m. Recorded depth in Martinique: 18-80m (live specimen at 80m). Previous recorded depth in the West Atlantic: 4-101m (Fernandes& Pimenta 2020). Metaxiasp. (Fig.10) m aterial examined .— Martinique. MADIBENTHOS. ‘Sud Atlantique’ • 1sh; sta. AB181; MNHN. geographic diStribution.— Only known from Martinique. bathymetric diStribution.— Only known from 22m. AB CD Fig. 10. — Metaxia sp. MNHN, sta. AB181, 2.6 mm. Scale bars: A, B, 1 mm; C, 200 μm; D, 100 μm. 475 Triphoridae from Martinique ZOOSYSTEMA • 2024 • 46 (18) A B CD E FG HI Fig. 11 . — A-E, Nanaphora decollata (Rolán & Fernández-Garcés, 1994), MNHN, sta. GB09 (KARUBENTHOS 1), 2.9 mm; a weak micro-sculpture on the teleoconch is shown in B; F-I, Nanaphora cf. verbernei (Moolenbeek & Faber, 1989); F, G, MNHN, sta. AB130, both 3.0 mm; H, I, MNHN, sta. AB149, 2.8 mm, 3.1 mm. Scale bars: A, F-I, 1 mm; B, E, 100 μm; C, D, 200 μm. 476 ZOOSYSTEMA • 2024 • 46 (18) Fernandes M.R. remarkS The single available shell of Metaxiasp. has a faint axial sculpture and a protoconch with 2.5 whorls (measuring 0.32mm long, 0.31mm wide), and its embryonic shell has a distinct, abapical zigzag spiral cord and a smaller, adapical spiral cord bordering the adapical portion of the whorl (Fig.10D). It could be an atypical shell of M. rugulosa, regarding the wide variation in the embryonic shell of this species (Rolán& Redfern 1996), although Caribbean shells of M.rugulosa are usually white and have a coarse axial sculpture (Rolán& Fernández-Garcés 2007, 2008; Lee 2009). Most Brazilian shells of M.rugulosa are brown (Fernandes& Pimenta 2011, 2020), similarly to Metaxiasp. (Fig.10A, B), which demands further studies – desirably with genetic evidence. Metaxia propinqua Rolán& Fernández-Garcés, 2008, from southern USA, has shells entirely white or with a white protoconch and light brown teleoconch. The white paratype of M. propinqua illustrated by Rolán& Fernández-Garcés (2008: fig.1E-G) seems a typical shell of M.rugulosa, contrary to the bicolor holotype. Another remarkably similar species to Metaxiasp. is Metaxia quadrata Faber, 2010, described from Aruba, based on a single (juvenile) shell. This species was described as yellowish brown, and the axial sculpture of the teleoconch is much faint, as observed in Metaxiasp. (Faber 2010). The apparent single difference between them is that M. quadrata has three spiral cords (vs two in Metaxiasp.) on the first protoconch whorl. Because Metaxia species are mainly differentiated by subtle differences in protoconch morphology, more material from the Lesser Antilles is required to properly evaluate the taxonomy of this morph from Martinique. Genus Monophorus Grillo, 1877 Monophorus olivaceus (Dall, 1889) (Figs2F; 19C-F) Triforis decorata var. olivacea Dall, 1889: 244. material examined.— Guadeloupe. KARUBENTHOS 1 • 1sh, 1spm; sta. GB03; MNHN • 1sh; sta. GS06; MNHN • 1sh; sta. GS07; MNHN • 3sh; sta. GS08; MNHN • 3sh; sta. GD10; MNHN • 2sh; sta. GB11; MNHN • 1sh; sta. GB12; MNHN • 15sh, 1spm; sta. GS13; MNHN • 5sh; sta. GB14; MNHN • 2sh; sta. GD15; MNHN • 1sh; sta. GS16; MNHN • 2sh; sta. GS18; MNHN • 2sh [juveniles]; sta. GB19; MNHN • 1spm; sta. GD21; MNHN • 1sh; sta. GS21; MNHN • 3sh; sta. GB22; MNHN • 1sh; sta. GS22; MNHN • 1sh; sta. GB24; MNHN • 1sh; sta. GB25; MNHN • 2sh; sta. GS26; MNHN • 1sh; sta. GS29; MNHN • 2sh [juveniles]; sta. GR34; MNHN • 1sh; sta. GB36; MNHN • 1sh; sta. GD61; MNHN. Martinique. MADIBENTHOS. ‘Nord Atlantique’ • 1sh; sta. AR143; MNHN • 1sh; sta. AB197; MNHN • 1sh; sta. AB199; MNHN • 1sh; sta. AD254; MNHN • 1sh [juvenile]; sta. AD260; MNHN • 1sh; sta. AD263; MNHN • 6sh; sta. AB556; MNHN • 8sh; sta. AS557; MNHN • 3sh; sta. AB559; MNHN • 1sh; sta. AB562; MNHN • 7sh; sta. AS565; MNHN. — ‘Sud Atlantique’ • 1sh; sta. AD084; MNHN • 1spm; sta. AB185; MNHN • 1sh; sta. AS255; MNHN • 3sh [juveniles]; sta. AB405; MNHN. — ‘Nord Caraibe’ • 5sh; sta. AB108; MNHN • 8sh; sta. AB159; MNHN • 2sh; sta. AB161; MNHN • 1sh, 1spm; sta. AD283; MNHN • 1sh; sta. AD294; MNHN • 4sh; sta. AR308; MNHN • 1sh; sta. AB362; MNHN • 1sh; sta. AS365; MNHN • 1sh; sta. AB372; MNHN • 2sh; sta. AS373; MNHN • 10sh; sta. AS375; MNHN • 7sh; sta. AB377; MNHN • 2sh; sta. AS378; MNHN • 2sh; sta. AB386; MNHN • 1sh; sta. AB388; MNHN • 1spm; sta. AR461; MNHN • 1sh [juvenile]; sta. AB510; MNHN • 5sh; sta. AS570; MNHN • 2sh; sta. AS572; MNHN • 1sh; sta. AS574; MNHN • 5sh; sta. AS576; MNHN. ‘Sud Caraibe’ • 1sh; sta. AS066; MNHN • 1sh; sta. AS071; MNHN • 1sh; AR100; MNHN • 1sh; sta. AR122; MNHN • 1sh; sta. AB126; MNHN • 6sh; sta. AB150; MNHN • 1spm; sta. AB152; MNHN • 1sh; sta. AB155; MNHN • 3sh; sta. AB177; MNHN • 1sh; sta. AD271; MNHN. ‘Baie de Fort-de-France’ • 1sh; sta. AB394; MNHN • 2sh; sta. AS581; MNHN. bathymetric diStribution.— Recorded depth in Guadeloupe: 9-80m (live specimens at 22-50m). Recorded depth in Martinique: 2-80m (live specimens at 4-40m). Previous recorded depth in the West Atlantic: 0-120m, with discrepant records down to 260m (Fernandes& Pimenta 2020). Genus Nanaphora Laseron, 1958 Nanaphora decollata (Rolán& Fernández-Garcés, 1994) (Fig.11A-E) Cheirodonta decollata Rolán& Fernández-Garcés, 1994: 20, figs1921, 23-24, 30CD; 2007: 20, pl.1, figs6-11.— Zhang 2011: 99, fig.289.— Redfern 2013: 127, fig.361.— Lamy& Pointier 2018: 284, pl.91, fig.2. Nanaphora decollata – Fernandes& Pimenta 2015: 502.— Rolán& Fernández-Garcés 2015: 53, fig.4V. type material.— Holotype. Cuba • sh; MNCN 15.05/11142. Paratypes. See the original description. type locality.— Cuba: Marianao beach, La Habana. material examined.— Guadeloupe. KARUBENTHOS 1 • 1sh; sta. GB09; MNHN. geographic diStribution.— Bahamas (Redfern 2013); Cuba (Rolán& Fernández-Garcés 1994); Cayman (this study); Antigua (Zhang 2011); Guadeloupe (Rolán& Fernández-Garcés 2015; Lamy& Pointier 2018; this study). bathymetric diStribution.— Recorded depth in Guadeloupe: 6m. Previous recorded depth in the West Atlantic: 2m (Rolán& Fernández-Garcés 1994) to 14m (Redfern 2013). remarkS Nanaphora decollata was originally described in Cheirodonta Marshall, 1983, a genus with only two confirmed species, i.e., the type species Cheirodonta pallescens (Jeffreys, 1867) from the East Atlantic and Cheirodonta dupliniana (Olsson, 1916) from the West Atlantic, plus a third species [Cheirodonta labiata (A. Adams, 1854), from Australia] with a tentative allocation (Marshall 1983; Fernandes& Pimenta 2015). The Atlantic species of Cheirodonta have radula with multicuspid teeth, of which the marginal are much elongated (Bouchet& Guillemot 1978; Bouchet 1985; Fernandes& Pimenta 2019b). In fact, there are significant 477 Triphoridae from Martinique ZOOSYSTEMA • 2024 • 46 (18) similarities between the radula of N. decollata (Rolán& Fernández-Garcés 1994) and those of Cheirodonta, mainly in the morphology of the lateral teeth, with the central and marginal teeth of N. decollata being also similar to those of C. dupliniana, but slightly different from those of C. pallescens, which shows central tooth with a diastema and marginal teeth with cusps restricted to the distal part of the basal plate. The protoconch of N. decollata is also slightly similar to that of Cheirodonta, bearing granules on the embryonic shell and two spiral cords on the larval shell (Fig.11E), although the former protoconch is shorter and its whorls are more convex. The most discrepant differences between N. decollata and Cheirodonta rely on teleoconch features, with the former showing a small and ovoid shell shape, nodules much larger (including the subperipheral and basal cords) and a much reduced suture (Fig.11A). These conditions are also observed in other West Atlantic species, which, perhaps prematurely, led Fernandes& Pimenta (2015) to include them in Nanaphora, a probably non-monophyletic genus which shows species with different patterns of embryonic and larval shell sculptures, and whose type species has a paucispiral protoconch (hindering comparisons) and unknown radula (Marshall 1983; Fernandes& Pimenta 2015). Citing Fernandes& Pimenta (2015: 502), “the affinity among the genera Nanaphora, Opimaphora Laseron, 1958 and Cheirodonta makes necessary a taxonomic revision of them to achieve a precise delimitation of each genus”, including a further molecular phylogeny. Two unusual teleoconch features of N. decollata were indicated by Rolán& Fernández-Garcés (1994) and Redfern (2013), and are confirmed here. The presence of micro-sculpture (Fig.11B) is not widespread in Triphoridae (Marshall 1983; Fernandes& Pimenta 2020), but some species currently allocated in Nanaphora may show it (Marshall 1983; see also the next species). Another remarkable feature of N. decollata is the emergence of the median spiral cord not between the adapical and abapical cords, as observed in most triphorids, but derived from a split of the adapical cord on the body whorl (Fig.11C), after the adapical cord had become axially elongated – it does not seem a supranumerical cord (Bouchet 1985; Fernandes& Pimenta 2015). The splitting of one spiral cord into two cords is observed in some species of Cerithiopsidae H. Adams& A. Adams, 1853, and it was reported in a Triphoridae species from Ascension Island (Bakker& Swinnen 2021), although with a different formation and major divergences in the shell. Rosenberg (2009) cited two new localities for the Caribbean range of Nanaphora verbernei (Moolenbeek& Faber, 1989), i.e., Cayman and Grenada, based on material from the ANSP collection. After checking images of the single lots for such localities (ANSP 200078, Grand Cayman Is., 2m - 2.8mm long, with protoconch; ANSP 296542, Prickly Bay, Grenada, 0-1m - 3.5mm long, without protoconch), the material from Cayman actually refers to N. decollata, whereas that from Grenada is worn and precludes further identification (although it seems N. decollata). The main difference between both species is the spiral sculpture of the protoconch, since N. decollata has always two spiral cords (Fig.11E). Shells without apex may be much similar owing to the ovoid shape and bifurcating adapical spiral cord of teleoconch (for N. verbernei, see Fig.13B and Moolenbeek& Faber 1989: fig.8). Rolán& Fernández-Garcés (1994) argued that the white band is only seen in the adapical cord of N. decollata, but it can be also present in the abapical cord in the penultimate whorl of N. verbernei, which is seen in most shells from Martinique without apex (Fig.11F-I); even though, it is preferred to name these decollate shells as N. cf. verbernei, and so far N. decollata is absent from Martinique. AB CD Fig. 12 . — Types of Nanaphora verbernei (Moolenbeek & Faber, 1989); credits to MSc. Hannco Bakker; A, ZMA.MOLL.136613, holotype; B-D, ZMA.MOLL.136655, paratypes. Scale bars: 1 mm. 478 ZOOSYSTEMA • 2024 • 46 (18) Fernandes M.R. Nanaphora verbernei (Moolenbeek& Faber, 1989) (Figs12; 13) Triphora verbernei Moolenbeek& Faber, 1989: 77, figs6-8. Triphora exiguum non C. B. Adams, 1850 – De Jong& Coomans 1988: 49. Cheirodonta verbernei – Rolán& Fernández-Garcés 1994: 20, figs17-18, 22, 30 CV. Cosmotriphora verbernei – Rolán& Fernández-Garcés 2007: 20, pl.1, figs17-18. Nanaphora verbernei – Fernandes& Pimenta 2015: 500 (new generic allocation, but the species was misidentified – see below). type material.— Holotype. Curaçao • sh; ZMA.MOLL.136613. Paratypes. Curaçao • 19sh; type locality; ZMA.MOLL.136655. type locality.— Curaçao: Boca Labadera, Santa Catarina, beach. material examined.— Guadeloupe. KARUBENTHOS 1 • 1sh [worn – previously identified as Nanaphora decollata]; sta. GM06; MNHN • 1sh [idem]; sta. GM11; MNHN • 1sh [juvenile – previously identified as Coriophora novem]; sta. GS13; MNHN. — KARUBENTHOS 2 • 1sh; sta. DW4545; MNHN • 1sh [juvenile]; sta. DW4551; MNHN. Martinique. MADIBENTHOS. ‘Nord Atlantique’ • 1sh; sta. AD261; MNHN • 1sh; sta. AD275; MNHN. — ‘Nord Caraibe’ • 1sh; sta. AD280; MNHN. ‘Sud Caraibe’ • 1sh; sta. AD115; MNHN. m aterial examined of N aNaphora cf . verberNei (Figs1F-I;1I).— Martinique. ‘Sud Atlantique’ • 4sh; sta. AB130; MNHN • 2sh, 1spm; sta. AB149; MNHN. geographic diStribution.— Cuba (Rolán& Fernández-Garcés 1994); Puerto Rico (Moolenbeek& Faber 1989); Guadeloupe and Martinique (this study); Bonaire and Curaçao (Moolenbeek& Faber 1989). bathymetric diStribution.— Recorded depth in Guadeloupe: 0-82m. Recorded depth in Martinique: 11-80m. Recorded depth of N. cf. verbernei in Martinique: 1-2m (live specimen at 2m). Previous recorded depth in the West Atlantic: 0-90m (Moolenbeek& Faber 1989). emended deScription Shell sinistral, cyrtoconoid, convex profile, up to 3.9mm long, 1.4mm wide, length/width ratio 2.8-3.0. Protoconch multispiral, conical/columnar, of 5.0-5.5 slightly convex whorls, 0.54-0.57mm long, 0.42-0.44mm wide; embryonic shell dome-shaped, covered by rounded granules; larval shell initially with two spiral threads but adapical one soon disappearing, with one spiral cord (abapical) remaining until the emergence of adapical cord in the penultimate whorl, the two cords situated at 32-36% and 61-68% of last whorl height; c.27 nearly rectilinear to slightly sigmoid axial ribs. Teleoconch with up to nine whorls; two spiral cords (adapical and abapical) at the beginning, abapical one continuous to that of protoconch; median spiral cord emerging narrowly at sixth or seventh whorl, bordering close the adapical cord, reaching the same size of abapical cord (adapical one is slightly more prominent on body whorl) after 1.5-2.0 whorls; rounded nodules of large size; 17-19 opisthocline axial ribs on seventh whorl; shallow suture, with small sutural cord; spiral micro-sculpture present in all teleoconch; subperipheral cord nodulose, adapical basal cord slightly nodulose to wavy, nearly smooth abapical basal cord; supranumerical cords not discernible (but peristome never intact); nearly circular aperture, 0.55-0.57mm long, 0.50-0.58mm wide, length/width ratio 0.9-1.1; anterior canal almost closed, crossed in its base by projection of outer lip, slightly curved backward, 0.34-0.50mm long, 0.22-0.26mm wide, length/width ratio 1.3-2.2; posterior canal as a small hole, detached from aperture. Light brown to orange protoconch; teleoconch light brown to nearly orange, with adapical spiral cord often whitish on body whorl, and some shells also show abapical cord lighter than adapical cord on mid whorls. remarkS The author initially allocated this species in Coriophora due to the spiral sculpture of the protoconch (Fig.13H, K), combined with a micro-sculpture on teleoconch (Fig.13G, J), shell shape, opisthocline axial ribs, shallow suture and late emergence of median spiral cord of teleoconch (Fig.13A-E) – regarding there are substantial divergences between the single species from the West Atlantic (C. novem) and the type species of Coriophora (Fernandes& Pimenta 2020). However, after reviewing details on the protoconch sculpture of Nanaphora verbernei in the literature from the Caribbean (Moolenbeek& Faber 1989; Rolán& Fernández-Garcés 1994), an error was detected. Fernandes& Pimenta (2015) interpreted the embryonic shell of the Caribbean species as reticulated (following shells from Brazil), owing to the inadequate SEM images previously available, despite being described with hemispheric tubercles in the material from Cuba (Rolán& Fernández-Garcés 1994). Fernandes& Pimenta (2015) also considered that N. verbernei had always two spiral cords in the larval shell, following shells from Brazil and the dubious image of a shell from Cuba (Rolán& Fernández-Garcés 1994: fig.22, which is bright in part of the protoconch, hampering a proper visualization), but they equivocally contested the description of the larval shell as having initially one but further two spiral cords (Moolenbeek& Faber 1989: fig.7). In fact, the Caribbean N.verbernei has rounded granules on the embryonic shell and the pattern 2-1-2-(1) spiral cords on the larval shell (Fig.13H, K). The Brazilian morph, despite showing a much similar teleoconch morphology, is not N.verbernei, as indicated by discrepant features on the radula and on the color of living specimens (Fernandes& Pimenta 2019b), and it will be discussed under the following species. Accordingly, the record of N.verbernei from Antigua (Zhang 2011) is related to the next species, and the records from Cayman and Grenada (Rosenberg 2009) are invalid (see remarks of N. decollata). The identification of N.verbernei from Guadeloupe by Rolán& Fernández-Garcés (2015), and followed by Lamy& Pointier (2018), is also incorrect (see next species); the real N.verbernei was hidden under the names of two other species (see material examined). Because E. Rolán was one reviewer of the manuscript that resulted in Fernandes& Pimenta (2015), nearly at the same time of the publication of Rolán& Fernández-Garcés (2015), it may have influenced the wrong identification of shells from Guadeloupe. Comparing the material from Gua- 479 Triphoridae from Martinique ZOOSYSTEMA • 2024 • 46 (18) ABCDE FG H IJ K Fig. 13. — Nanaphora verbernei (Moolenbeek & Faber, 1989): A, MNHN, sta. GM11 (KARUBENTHOS 1), 3.8 mm; B, MNHN, sta. AD115, 3.9 mm; C, MNHN, sta. AD261, 3.7 mm; D, F-I, MNHN, sta. AD280, 3.9 mm; E, J, K, MNHN, sta. DW4545 (KARUBENTHOS 2 expedition), 3.8 mm. A weak micro-sculpture on the teleoconch is shown in G, J. Scale bars: A-E, 1 mm; F, 500 μm; G-H, J-K, 100 μm; I, 200 μm. 486 ZOOSYSTEMA • 2024 • 46 (18) Fernandes M.R. material examined.— Guadeloupe. KARUBENTHOS 1 • 1sh; sta. GD35; MNHN. Martinique. MADIBENTHOS. ‘Sud Caraibe’ • 1sh; sta. AD218; MNHN. bathymetric diStribution.— Recorded depth in Guadeloupe: 66m. Recorded depth in Martinique: 60m. Previous recorded depth in the West Atlantic: 58-163m (Fernandes& Pimenta 2020). remark This species was identified as “Triphora”sp. by Rolán& Fernández-Garcés (2015: fig.4U) and as Isotriphorasp. 1 by Lamy& Pointier (2018: pl.91, fig.14). Genus “Triphora” Blainville, 1828 s.l. Triphora ellyae De Jong& Coomans, 1988 (Figs3B; 20G-H) Triphora ellyae De Jong& Coomans, 1988: 50, pl.34, fig.242. material examined.— Guadeloupe. KARUBENTHOS 1 • 2sh [previously identified as Similiphora intermedia]; sta. GB05; MNHN • 1sh, 1spm [idem]; sta. GB06; MNHN • 1sh [idem]; sta. GB11; MNHN • 1sh [identified as Monophorus ateralbus by Rolán& Fernández-Garcés (2015: fig.4X)]; sta. GD13; MNHN • 1sh [previously identified as Similiphora intermedia]; sta. GS13; MNHN. Martinique. MADIBENTHOS. ‘Nord Atlantique’ • 1sh; sta. AD260; MNHN • 1sh; sta. AD263; MNHN • 1sh; sta. AB350; MNHN • 1sh; sta. AB556; MNHN • 2sh; sta. AS557; MNHN. — ‘Nord Caraibe’ • 2sh; sta. AB159; MNHN • 1sh; sta. AD283; MNHN • 1sh; sta. AD290; MNHN • 1sh; sta. AD299; MNHN • 1spm; sta. AB308; MNHN • 1sh; sta. AS375; MNHN • 3sh; sta. AB463; MNHN • 1sh; sta. AS570; MNHN • 2sh; sta. AS572; MNHN. ‘Sud Caraibe’ • 1sh; sta. AB062; MNHN • 1sh; sta. AB169; MNHN. bathymetric diStribution.— Recorded depth in Guadeloupe: 5-50m (live specimen at 23m). Recorded depth in Martinique: 5-80m. Previous recorded depth in the West Atlantic: 3-110m (Fernandes& Pimenta 2020). Triphora martii Rolán& Fernández-Garcés, 1995 (Fig.3C) “Triphora” martii Rolán& Fernández-Garcés, 1995: 16, figs39-42. material examined.— Guadeloupe. KARUBENTHOS 1 • 1sh [identified as Eutriphora bermudensis by Rolán& Fernández-Garcés (2015: fig.4T)]; sta. GS18; MNHN • 1sh [previously identified as Iniforis turristhomae]; sta. GD59; MNHN. Martinique. MADIBENTHOS. ‘Sud Atlantique’ • 1sh; sta. AS252; MNHN. bathymetric diStribution.— Recorded depth in Guadeloupe: 49-88m. Recorded depth in Martinique: 10-12m. Previous recorded depth in the West Atlantic: 20-40m (Rolán& Fernández-Garcés 1995). Triphora portoricensis Rolán& Redfern, 2008 (Figs16; 17) “Triphora” portoricensis Rolán& Redfern in Rolán& Fernández-Garcés 2008: 158, fig.32A-E. Iniforissp. – Redfern 2001: 66, pl.33, fig.278A, B. “Triphora” portoricensis – Redfern 2013: 127, fig.358A, B.— Fernandes& Pimenta 2015: 505, fig.6.— Fernandes& Pimenta 2020: 168. type material.— Holotype. Puerto Rico • sh; FLMNH 363895. Paratype. Bahamas • 1sh; Abaco; 26°44’00”N, 77°09’00”W; 9.IX.1987; C. Redfern leg.; depth 52m; BMSM 55395. type locality.— Puerto Rico. material examined.— Martinique. MADIBENTHOS. ‘Nord Atlantique’ • 3sh; sta. AB350; MNHN. ‘Sud Atlantique’ • 1sh; sta. AB183; MNHN •1sh; sta. AB191; MNHN • 1sh; sta. AB193; MNHN • 1sh; sta. AS255; MNHN • 5sh; sta. AB405; MNHN. geographic diStribution.— Bahamas (Rolán& Fernández-Garcés 2008; Redfern 2013); Puerto Rico (Rolán& Fernández-Garcés 2008); Martinique (this study); Brazil. Rio Grande do Norte to Rio de Janeiro (Fernandes& Pimenta 2015). bathymetric diStribution.— Recorded depth in Martinique: 14-23m. Previous recorded depth in the West Atlantic: 23-100m (Fernandes& Pimenta 2015, 2020). emended deScription Shell sinistral, conical to slightly pupoid, rectilinear to slightly convex profile, reaching 3.6mm long, 1.3mm wide, length/ width ratio 2.6-3.1; adult shells reach at least 1.9mm in length. Protoconch sub-columnar, slightly inflated, of 2.753.25 whorls, 0.40-0.58mm long, 0.41-0.51mm wide, weak distinction between protoconch and teleoconch; initial 0.5 whorl smooth, but axial ribs soon appear, with their adapical part emerging on the abapical part of the smooth whorl above, assuming an irregular shape, often arrow-shaped initially but varying from sigmoid, nearly orthocline, strongly opisthocline or strongly prosocline on the last whorl, where it numbers 20-34 ribs; one main spiral cord often occupying the mid portion of the whorl (sometimes the abapical portion), situated at 44-61% of last whorl height, keel-shaped, in addition to a small sutural cord. Teleoconch with up to seven whorls; two spiral cords (adapical and abapical) at the beginning, the abapical one continuous to that of protoconch; median spiral cord emerging at the end of second whorl or at the third whorl, reaching the same size than other cords after c.1.5 whorl; at the body whorl, the three cords being equidistant and nearly the same size; 20-21 nearly orthocline axial ribs on the fifth whorl; rounded nodules of medium size, with square interspaces among nodules; distinct suture, with a sutural cord; slightly nodulose to wavy subperipheral cord, one to two smooth basal cords, often narrow, but sometimes well-developed; no distinct supranumerical cords, but a minute spiral thread sometimes appearing between abapical and subperipheral cords prior to the peristome; aperture rounded to slightly rhomboid (with a discrete notch in the posterior end), 0.47-0.83mm long, 0.43-0.59mm wide, length/width ratio 1.1-1.4; anterior canal very short, open, 0.07-0.15mm long, 0.13-0.20mm wide, length/width ratio 0.4-0.7. White to light cream shell, with adapical spiral cord and base slightly darker. 487 Triphoridae from Martinique ZOOSYSTEMA • 2024 • 46 (18) AB C D M EF GH I K L O N J Fig. 18 . — Live triphorids from Martinique and Guadeloupe (marked with asterisk): A, B, Coriophora novem (Nowell-Usticke, 1969), MNHN, sta. GM08 (A), AD254 (B); C-F, Cosmotriphora melanura (C.B.Adams, 1850), MNHN, sta. GB03 (C), AS096 (D), GS17 (E), AD214 (F); G-I, K, Iniforis turristhomae (Holten, 1802), MNHN, sta. AS075 (G-H), AM005 (I), GB07 (K); J, L-O, Metaxia excelsa Faber & Moolenbeek, 1991, MNHN, sta. AB123 (J), GB12 (L, O), AB362 (M), AB197 (N). Credits: MNHN team. 488 ZOOSYSTEMA • 2024 • 46 (18) Fernandes M.R. remarkS The shell from Guadeloupe identified by Lamy& Pointier (2018: pl.92, fig.11) as Triphora portoricensis is actually the holotype of Marshallora monteiroi n. comb. Some shells of T. portoricensis from Martinique (Fig.16) resemble one shell tentatively identified as M. monteiroi n. comb. (Fig.7K), but the former morph has a slightly darker adapical spiral cord of teleoconch (vs spiral cords with a more homogeneous color in M. monteiroi n. comb.) and basal cords (Fig.17A, E) often considerably thinner than M. monteiroi n. comb. (Fig.7L-M). Shells of T. portoricensis from Martinique vary in the adult shell length (1.9-3.6mm) and in the axial sculpture of the protoconch, but the largest shells are similar to the holotype from Puerto Rico (4.4mm to eight teleoconch whorls vs up to seven whorls in shells from Martinique). In contrast, shells from Bahamas and Brazil are entirely white (Redfern 2013; Fernandes& Pimenta 2015); those from Brazil have a different teleoconch sculpture in late whorls, with a more distant adapical spiral cord (not discernible in Bahamian shells, only known by juveniles), and reach up to 10mm to 13 teleoconch whorls (Fernandes& Pimenta 2015). This suggests that the current concept of T. portoricensis is actually a species complex masked by the similar protoconch morphology, which could be derived from convergence or parallelism. Triphora cf. scylla Fernandes& Pimenta, 2015 (Fig.3D) Triphora scylla Fernandes& Pimenta, 2015: 509, fig.8. A B C D E F GHI C Fig. 19. — Live triphorids from Martinique and Guadeloupe (marked with asterisk): A, B, Metaxia taeniolata (Dall, 1889), MNHN, sta. AD261; C-F, Monophorus olivaceus (Dall, 1889), MNHN, sta. GD21 (C), AB185 (D), AR461 (E, F); G, H, Nanaphora leei Fernandes & Pimenta, 2015, MNHN, sta. AB400 (G), AB419 (H); I, Nanaphora cf. verbernei (Moolenbeek & Faber, 1989), MNHN, sta. AB149. Credits: MNHN team. 489 Triphoridae from Martinique ZOOSYSTEMA • 2024 • 46 (18) material examined.— Martinique. MADIBENTHOS. ‘Nord Atlantique’ • 1sh [juvenile]; sta. AD275; MNHN • 1sh [juvenile]; sta. AB562; MNHN. bathymetric diStribution.— Recorded depth in Martinique: 14-80m. Previous recorded depth in the West Atlantic: 20-150m (Fernandes& Pimenta 2020). DISCUSSION After more than 500 collection events and 33 triphorid species sampled from the shallow waters of Martinique, the MADIBENTHOS expedition significantly improved our knowledge about the diversity of Triphoridae from the Caribbean. Besides the description of three new species A B C D EFG HI J Fig. 20. — Live triphorids from Martinique and Guadeloupe (marked with asterisk): A-D, Nototriphora decorata (C.B. Adams, 1850), MNHN, sta. GR10 (A), AS253 (B), AB382 (C), AB161 (D); E, F, Sagenotriphora osclausum (Rolán & Fernández-Garcés, 1995), MNHN, sta. AB191; G, H, Triphora ellyae De Jong & Coomans, 1988, MNHN, sta. AB308; I, J, Marshallora monteiroi (Rolán & Fernández-Garcés, 2015) n. comb., MNHN, sta. AD283 (I), GD21 (J). Credits: MNHN team. 490 ZOOSYSTEMA • 2024 • 46 (18) Fernandes M.R. (all non-planktotrophic), there were some new records of planktotrophic species to the Lesser Antilles. Cheirodonta dupliniana and Strobiligera inaudita were reported along the North and South portions of the West Atlantic, but neither were previously recorded from any Caribbean island (Fernandes& Pimenta 2020). Despite the abovementioned No. species No. shells No. stations No. species Planktotrophic Planktotrophic Non-planktotrophic Non-planktotrophic No. shells No. species 675 600 525 225 150 75 0 0200 400 600 800 1000 1200 1400 1600 10 20 25 30 35 40 15 5 0 0 1 1 to 2 1 to 2 3 to 5 3 to 5 6 to 20 6 to 2021 to 50 21 to 5051 to 100 >100 >50 2 3 4 0 1 2 3 4 5 6 7 5 10 15 20 25 30 450 375 300 Abundance Species A B CD Iniforis turristhomae Cosmotriphora melanura Fig. 21 . — A, Individual rarefaction curve of triphorids from MADIBENTHOS; blue curves indicate the 95% confidence interval; B, abundance distribution of triphorids from MADIBENTHOS; C, D, number of species vs number of shells (C) or number of stations (D), regarding planktotrophic (black bars) and nonplanktotrophic (grey bars) species. 491 Triphoridae from Martinique ZOOSYSTEMA • 2024 • 46 (18) issues of identification, Nanaphora leei is now confirmed to the Caribbean, and not restricted to Brazil; the same may apply for Triphora cf. scylla, pending additional material. Triphora martii was previously known from Cuba (Rolán& Fernández-Garcés 1995; Espinosa etal. 2007; García& Capote 2013). On the other hand, no shell of Inella triserialis was sampled by MADIBENTHOS, confirming this is a deep-sea species; the identification from Martinique is uncertain (Lamy& Pointier 2018), regarding the poor condition of types, from the deep sea of the Yucatán Strait (Rolán& Fernández-Garcés 2008; Fernandes& Pimenta 2019a). The individual rarefaction curve of triphorids from MADIBENTHOS (Fig.21A) is nearly saturated but did not reach the asymptote, suggesting more than 33 species exist in Martinique. Accordingly, Chao1 estimator suggested the occurrence of 38 species, i.e., five species from Martinique were possibly missed (not sampled) by MADIBENTHOS. This number is considerably inferior than the total triphorid species found in similar efforts by the MNHN team in the Indo-Pacific, such as New Caledonia (174 species, 42 stations – Bouchet etal. 2002) and Vanuatu (259 species, 566 stations – Albano etal. 2011). Similarly, Marshall (1983) indicated the existence of at least 80 triphorid species in a single sand sample No. species AB Brushing (AB): 68 stations 0 sp (5.9%) 1 sp (11.7%) 2 spp (17.6%) 3 spp (11.7%) 4 spp (20.6%) 5 spp (14.7%) 6 spp (10.3%) 7 spp (2.9%) 9 spp (4.4%) Dredging (AD): 126 stations Visual at intertidal (AM): 51 stations Visual at diving (AR): 204 stations Suction (AS): 42 stations 0 sp (53.9%) 1 sp (23.8%) 2 spp (6.3%) 3 spp (5.5%) 4 spp (2.4%) 5 spp (3.2%) 8 spp (0.8%) 9 spp (1.6%) 11 spp (0.8%) 12 spp (0.8%) 13 spp (0.8%) ) %) 0 sp (82.3%) 1 sp (13.7%) 2 spp (3.9%) 0 sp (96.1%) 1 sp (2.4%) 2 spp (1.0%) 5 spp (0.5%) 0 sp (9.5%) 1 sp (16.7%) 2 spp (21.4%) 3 spp (16.7%) 4 spp (7.0%) 5 spp (7.0%) 6 spp (14.3%) 7 spp (2.4%) 8 spp (4.8%) 56 30 25 20 15 5 0 AR AM AB AS AD 10 22 23 26 Fig. 22. — A, number of triphorid species obtained in MADIBENTHOS by each sampling method (AB, brushing; AD, dredging; AM, visual at intertidal; AR, visual at diving; AS, suction); B, number of species sampled vs proportion of stations according to each sampling method in MADIBENTHOS. 492 ZOOSYSTEMA • 2024 • 46 (18) Fernandes M.R. from NE Australia. These contrasting numbers reflect the known higher diversity of Triphoridae in the Indo-Pacific (Albano etal. 2011). The abundance distribution of triphorids (Fig.21B) shows two species with skewed distributions, i.e., Iniforis turristhomae and Cosmotriphora melanura. This is reflected in the low evenness (0.276; 0.258-0.294), although the dominance was also low (D: 0.209; 0.192-0.226), whereas Shannon’s diversity (H: 2.208; 2.141-2.274) and Pielou’s equitability (J: 0.631; 0.612-0.650) were moderately high. Seven species (all with planktotrophic development) were considered abundant (51100shells) or much abundant (>100shells) and were sampled in more than 20 stations, whereas eight species (with planktotrophic or non-planktotrophic modes) were much rarer (i.e., represented by singletons or doubletons) and sampled in one or two stations (Fig.21C, D). Not all planktotrophic species West Atlantic (18 spp.; 54.5%) NW Atlantic (8 spp.; 24.2%) Lesser Antilles (2 spp.; 6.1%) Martinique (5 spp.; 15.2%) No. species Total Shared species with Martinique 2922 Caribbean Atlantic W est A t l ant ic ( 18 spp.; 54.5% ) NW A t l a nt ic ( 8 s pp .; 24.2% ) Lesse r A nt il les ( 2 spp.; 6.1% ) Martiniqu e ( 5 s pp .; 15.2% ) N o. s p ec i es Tota l S hared species with Martiniqu e 2 9 22 C aribbean A tl a nti c No. species 26 30 25 15 5 30 30 0 Nord Atlantique Sud Atlantique Nord Caraïbe Sud Caraïbe Baie de Fort-deFrance 22 20 18 11 A C D EF B5960 50 40 30 20 10 0 Brazil CubaMartinique Guadeloupe Abaco ABCIs. 20 40 21 33 29 25 22 14 14 -0.5 -0.60 0.60-0.45 0.45-0.30 0.30-0.15 0.00 0.15 -0.4 -0.3 -0.2 -0.1 0.0 0.1 0.2 0.3 0.4 -0.5 -0.60 0.60-0.45 0.45-0.30 0.30-0.15 0.00 0.15 -0.4 -0.3 -0.2 -0.1 0.0 0.1 0.2 0.3 0.4 18 Fig. 23. — A, Geographic range of triphorids from Martinique, whether endemic vs present in other sites in Lesser Antilles or beyond in the Northwest or West Atlantic; B, total shallow-water triphorids recorded in each well-sampled locality, including the number of shared species with Martinique; references from Brazil are those of M. Fernandes and colleagues, from Cuba those of E. Rolán (or J. Espinosa) and colleagues, from Aruba, Bonaire and Curaçao (ABC Is.) are De Jong & Coomans (1988), Moolenbeek & Faber (1989), Faber & Moolenbeek (1991) and Faber (2010), and Redfern (2013) from Abaco, Bahamas; C, number of triphorid species per geographic zone; D, number of triphorid species per Caribbean or Atlantic side in Martinique; E, F, nMDS of geographic and bathymetric zones regarding triphorid species, based on Bray-Curtis (E) and Jaccard (F) indexes: dark green, Nord Atlantique; light green, Sud Atlantique; dark blue, Nord Caraibe; light blue, Sud Caraibe; red, Baie de Fort-de-France; dots, 0-10 m; squares, 11-20 m; triangles, 21-30 m; X, 31-60 m; + - 61-85 m. 493 Triphoridae from Martinique ZOOSYSTEMA • 2024 • 46 (18) were abundant or present in many stations, because there might be biotic and abiotic restrictions for larval settlement and adult survivorship (such as the presence of particular sponge hosts). Similar higher abundances of planktotrophic triphorids were also observed in a survey from Vanuatu, SW Pacific (Albano etal. 2011), suggesting that non-planktotrophic triphorids are demographically rarer owing to certain biological (e.g. short or even null larval dispersion) or ecological restrictions. Two thirds (22 of 33) of the triphorid species from MADIBENTHOS are planktotrophic, a proportion higher than the nearly 50% found for triphorids from Brazil (Fernandes& Pimenta 2020) and southern Australia (Marshall 1983). However, the study of deep-sea material from Martinique will certainly alter this scenario by adding more non-planktotrophic species. There was a bias in the sampling methods on MADIBENTHOS, with 42 suction (AS) events, 51 visual efforts at intertidal (AM), 68 of brushing (AB), 126 of dredging (AD) and 204visual efforts at diving (AR). No method could sample all 33triphorid species due to the rarity of many of them, but the visual methods (AM and AR) were clearly poor at the total number of sampled species and on the required attempts to obtain a single specimen (Fig.22), due to the small size of triphorids (Albano etal. 2011). On the other hand, brushing and suction were once again regarded as efficient methods to obtain triphorids (Albano etal. 2011), sampling similar numbers of species (22 vs 23) and proportions of species (up to nine or eight) per station (Fig.22). Despite that c.54% of dredging stations did not recover any triphorid, some Atlantic side N Caribbean side 10 km 14º21’N, 60º57’W 14º40’N, 60º42’W 6 sp. 7 sp. 8 sp. 9 sp. 11 sp. 12 sp. 13 sp. Triphora portoricensis Iniforis gudeliae Marshallora cf. modesta A B C DEF Triphora ellyae Iniforis pseudothomae Fig. 24. — A, hyper-diverse stations of Triphoridae from MADIBENTHOS, containing more than six species; B-F, maps of geographic distribution of selected species. All maps obtained from Google Earth. 494 ZOOSYSTEMA • 2024 • 46 (18) Fernandes M.R. hyper-diverse dredging stations recovered up to 13 species, mainly at depths below 50m, in which brushing and suction did not occur. In addition to the restriction of depth, there were more dredging stations than the sum of brushing and suction stations, and the area covered by a dredging is often larger than of a AB or AS station, which explain the higher number of sampled species (Fig.22A). However, dredging usually samples empty shells, contrary to brushing and suction, which in most cases obtain live triphorids (rarely shells with hermit crabs; pers. obs. for brushing). Unfortunately, the MADIBENTHOS expedition did not aim for the storage of live triphorids in ethanol, and the amount of photographed live specimens probably does not reflect the actual number, avoiding further comparisons. The five species currently endemic from Martinique (“Inella”sp., Isotriphora pardus n.sp., Isotriphorasp., Metaxia discus n.sp., Metaxiasp.) have non-planktotrophic development (Fig.23A), but their extreme rarity (sampled as singletons or doubletons, with the exception of I.pardus n.sp.) and the lack of intensive sampling in other parts of the Lesser Antilles avoid such certainty. Other species sampled from Martinique but absent from Guadeloupe are Cosmotriphora arnoldoi, Triphora cf. scylla and Triphora portoricensis, although the two former are singleton or doubleton. In contrast, 25out of 29species from Guadeloupe (86.2%) are shared with Martinique (Fig.23B); Nanaphora decollata, Isotriphora peetersae Moolenbeek& Faber, 1989 (KARUBENTHOS stations: GS05 [2], GB11 [1], GS29 [1]) and Triphora elvirae (KARUBENTHOS station: GR10 [1]) were absent from Martinique, whereas the non-planktotrophic Triphora guadaloupensis was absent from MADIBENTHOS and KARUBENTHOS 1. The triphorid fauna from ABC Is. has 14 of the 18 recorded species (77.8%) shared with Martinique, whereas Abaco (Bahamas) shares with Martinique 14 of 22 species (63.6%) (Fig.23B). The similarity drops when comparing shallow-water triphorids from Martinique with those from Cuba (21 of 40 species shared, i.e., 52.5%), which has more species than Martinique due to a much larger coastline and with researchers actively describing species since 1992; many triphorids from Cuba are non-planktotrophic, which influences their endemism. The (bio)geographic distance between Martinique and Brazil partly explains the low number of shared species (20 of the 59 species from Brazil, i.e., 33.9%); Brazil has more known triphorids owing to a much larger coastline, comprising different marine provinces (Spalding etal. 2007), in addition to several species known from the end of the continental shelf (Fernandes& Pimenta 2019a, 2020), a depth zone poorly explored in Martinique. An expected but intriguing absence from Martinique is the planktotrophic Triphora atlantica Smith, 1890, which is widely distributed in the southern USA and in Brazil, but with a single Caribbean record from Puerto Rico (Rolán& Fernández-Garcés 2008; Fernandes& Pimenta 2020). Another sampling bias in MADIBENTHOS was related to the geographic zones within Martinique, with the increasing order (excluding AN stations): Baie de Fort-de-France (49stations), Nord Atlantique (69), Nord Caraibe (95), Sud Atlantique (131), Sud Caraibe (147); i.e., the Atlantic side had less stations (200) than the Caribbean side (291). Even though, the zones with most triphorid species are, in a decreasing order: Nord Atlantique (26 species / Chao1 estimator: 26.1 species), Sud Atlantique (22 / 25.7), Nord Caraibe (20 lsotriphora tricingulata lnella spinosa Cosmotriphora arnoldoi “lnella” sp. Metaxia taeniolata lsotriphora pardus Eutriphora bermudensis Coriophora novem Sagenotriphora osclausu m Similiphora intermedia Triphora ellyae Triphora martii Triphora portoricensis Marshallora monteiroi lniforis pseudothomae Triphora cf. scylla Metaxia sp. lsotriphora sp. Similiphora sp. Metaxia discus lniforis turristhomae Marshallora cf. modesta lniforis gudeliae Nanaphora leei Latitriphora albida Cheirodonta dupliniana Nototriphora decorata Cosmotriphora melanura Nanaphora verbernei Monophorus olivaceus Metaxia excelsa Metaxia rugulosa Strobiligera inaudita Depth (m) 0102030405060708090 Fig. 25. — Depth range of triphorid species in Martinique, sampled by MADIBENTHOS. 495 Triphoridae from Martinique ZOOSYSTEMA • 2024 • 46 (18) / 22.0), Sud Caraibe (18 / 27.0) and Baie de Fort-de-France (11 / 12.0) (Fig.23C, D). In the nMDS groupings (Fig.23EF), the shallower (0-30m) stations from the Caribbean side (including Baie de Fort-de-France) mainly grouped together, just like shallower stations from the Atlantic side, despite the skewed category ‘Sud Atlantique 0-10m’ in the presence/ absence matrix (Fig.23F). Deeper (31-85m) stations also grouped close (Fig.23E-F), except the category ‘Sud Caraibe 61-85m’, which had a highly discrepant relative abundance of Iniforis turristhomae (72.7%). Some hyper-diverse stations at Nord Atlantique derived from deeper dredging (Fig.24A) lye close to the arbitrary limit with Nord Caraibe (Bouchet etal. 2019: 217), and influenced the higher number of triphorids in the Atlantic side of Martinique (Fig.23C, D) despite the reduced number of stations, regarding other portions of the island lacked dredging in deeper levels. The wider ‘shelf’ of the Atlantic side might have also contributed to the increased number of triphorids owing to a supposed higher availability of microhabitats (e.g. perhaps more sponge hosts). As expected Iniforis turristhomae (34.3%) Nototriphora decorata (12.0%) Monophorus olivaceus (11.1%) Cosmotriphora melanura (10.8%) Marshallora cf. modesta (5.6%) Sagenotriphora osclausum (4.4%) Iniforis gudeliae (3.5%) Nanaphora leei (3.2%) Similiphora intermedia (2.6%) Metaxia excelsa (2.6%) Triphora ellyae (2.3%) Latitriphora albida (2.0%) Nanaphora verbernei (2.0%) Others (3.2%) Iniforis turristhomae (51.0%) Cosmotriphora melanura (10.8%) Monophorus olivaceus (6.6%) Nototriphora decorata (6.4%) Iniforis gudeliae (4.9%) Sagenotriphora osclausum (4.5%) Coriophora novem (3.6%) Metaxia excelsa (3.1%) Iniforis pseudothomae (1.4%) ) Similiphora intermedia (1.2%) Nanaphora leei (1.1%) ) Triphora portoricensis (1.1%) Others (4.1%) Iniforis turristhomae (48.5%) Cosmotriphora melanura (12.4%) Monophorus olivaceus (9.4%) Iniforis pseudothomae (6.4%) Metaxia excelsa (4.5%) Sagenotriphora osclausum (4.5%) Coriophora novem (4.2%) Nototriphora decorata (3.0%) Triphora portoricensis (1.5%) Similiphora intermedia (1.2%) Others (3.6%) Iniforis turristhomae (12.0%) Cosmotriphora melanura (24.1%) Marshallora monteiroi (14.4%) Monophorus olivaceus (7.2%) Sagenotriphora osclausum (7.2%) Coriophora novem (6.0%) Metaxia excelsa (4.8%) Metaxia rugulosa (3.6%) Nototriphora decorata (3.6%) Triphora ellyae (3.6%) Others (13.2%) Cosmotriphora melanura (29.5%) Iniforis turristhomae (24.5%) Coriophora novem (8.0%) Metaxia excelsa (8.0%) Nototriphora decorata (5.3%) Metaxia rugulosa (4.5%) Iniforis pseudothomae (3.1%) Sagenotriphora osclausum (3.1%) Inella spinosa (1.8%) Monophorus olivaceus (1.8%) Triphora ellyae (1.8%) Others (8.5%) 61-25 m (n = 224) 31-60 m (n = 83) 11-20 m (n = 637) 0-10 m (n = 341) 21-30 m (n = 330) Fig. 26. — Proportion of triphorid species sampled by MADIBENTHOS, according to depth zones and total number of shells/specimens (n). 502 ZOOSYSTEMA • 2024 • 46 (18) Fernandes M.R. Station Locality Latitude (N) Longitude (W) Initial depth (m) Final depth (m) Sampling method GS04 Grand Cul-de-sac marin 16°21.97’ 61°37.98’ – 11 Suction GB05 Pointe à Lézard 16°08.43’ 61°46.92’ – 12 Brushing GS05 Grand Cul-de-sac marin 16°21.72’ 61°36.35’ – 22 Suction GB06 Tête à l’Anglais 16°22.9’ 61°45.94’ – 23 Brushing GM06 Îlet Fortune 16°09’ 61°33.67’ – 1 Visual at intertidal GS06 Grand Cul-de-sac marin 16°21.75’ 61°36.07’ – 23 Suction GB07 Pointe du Quesy 16°06.07’ 61°46.37’ – 6 Brushing GS07 Pointe à Lézard 16°08.43’ 61°46.92’ – 12 Suction GB08 Pointe à Lézard 16°08.43’ 61°46.92’ – 12 Brushing GM08 Pointe de l’Ermitage 16°07.57’ 61°46.45’ – 1 Visual at intertidal GS08 Tête à l’Anglais 16°22.9’ 61°45.94’ – 23 Suction GB09 Baie de Bouillante 16°08.07’ 61°46.71’ – 6 Brushing GS09 Pointe de l’Ermitage 16°07.61’ 61°46.53’ – 11 Suction GB10 Pointe de Malendure 16°05.95’ 61°47.5’ – 8 Brushing GD10 N Baie de Bouillante 16°08.48’ 61°47.03’ – 54 Dredging GR10 Pointe à Lézard 16°08.43’ 61°46.92’ – 29 Visual at diving GB11 Côte Ferry 16°17.69’ 61°48.23’ – 10 Brushing GM11 Cabrit 16°11.97’ 61°34.28’ – 1 Visual at intertidal GS11 S Rocroy 16°02.38’ 61°45.71’ – 17 Suction GB12 Port-Louis 16°25.61’ 61°32.57’ – 14 Brushing GR12 Tête à l’Anglais 16°22.9’ 61°45.94’ – 21 Visual at diving GS12 Baie de Bouillante 16°08.07’ 61°46.71’ – 6 Suction GB13 Port-Louis 16°23.26’ 61°31.79’ – 10 Brushing GD13 Petite Anse 16°05.72’ 61°46.26’ – 5 Dredging GR13 Pointe de l’Ermitage 16°07.61’ 61°46.53’ – 11 Visual at diving GS13 Îlet Pigeon 16°02.4’ 61°45.6’ – 50 Suction GB14 Near Port-Louis 16°23.74’ 61°32.07’ – 49 Brushing GR14 Vieux Habitan 16°03.25’ 61°46.17’ – 27 Visual at diving GS14 Pointe de Malendure 16°05.95’ 61°47.5’ – 8 Suction GB15 S Port-Louis, W Petit Canal 16°22.57’ 61°31.74’ – 8 Brushing GD15 Anse à la Barque 16°05.39’ 61°46.48’ – 50 Dredging GR15 Pointe du Quesy 16°06.07’ 61°46.37’ – 11 Visual at diving GS15 Sec Ferry 16°17.51’ 61°48.96’ – 27 Suction GB16 Port-Louis 16°27.34’ 61°32.07’ – 19 Brushing GD16 Anse Caraïbe 16°12.37’ 61°47.2’ – 10 Dredging GS16 Port-Louis 16°25.61’ 61°32.57’ – 25 Suction GB17 Pointe Grigri 16°23.26’ 61°31.79’ – 13 Brushing GR17 Baie de Bouillante 16°08.07’ 61°46.71’ – 13 Visual at diving GS17 Port-Louis 16°23.26’ 61°31.79’ – 3 Suction GB18 Port-Louis 16°25.99’ 61°32.92’ – 45 Brushing GS18 Near Port-Louis 16°23.74’ 61°32.07’ – 49 Suction GB19 Port-Louis 16°23.26’ 61°31.79’ – 11 Brushing GR19 Îlet Pigeon 16°02.4’ 61°45.6’ – 15 Visual at diving GS19 S Port-Louis 16°22.57’ 61°31.74’ – 8 Suction GB20 Port-Louis 16°26.78’ 61°32.41’ – 16 Brushing GD20 Near Baie de Baille-Argent 16°15.54’ 61°48.71’ – 35 Dredging GR20 Pointe de Malendure 16°05.95’ 61°47.5’ – 8 Visual at diving GS20 Port-Louis 16°27.34’ 61°32.07’ – 19 Suction GB21 Trou à l’orage 16°22.88’ 61°31.43’ – 8 Brushing GD21 Near Baie de Baille-Argent 16°15.55’ 61°48.8’ – 40 Dredging GR21 Sec Ferry 16°17.51’ 61°48.96’ – 27 Visual at diving GS21 Pointe Grigri 16°23.26’ 61°31.79’ – 14 Suction GB22 Pointe Montagnier 16°30.57’ 61°28.45’ – 12 Brushing GS22 Pointe d’Antigues 16°25.99’ 61°32.92’ – 45 Suction GB23 Grotte Amédier 16°30.04’ 61°28.79’ – 16 Brushing GS23 Port-Louis 16°23.26’ 61°31.79’ – 7 Suction GB24 Sec Pâté 15°54’ 61°39.3’ – 25 Brushing GD24 Port-Louis 16°25’ 61°33’ – 150 Dredging GS24 Port-Louis 16°26.78’ 61°32.41’ – 16 Suction GB25 Marina de Rivière Sens 15°59’ 61°43.08’ – 25 Brushing GD25 Port-Louis 16°25’ 61°33’ – 160 Dredging GS25 Pointe Montagnier 16°30.57’ 61°28.45’ – 12 Suction GB26 Near Caret 16°22.28’ 61°38.14’ – 29 Brushing GS26 Grotte Amédier 16°30.04’ 61°28.79’ – 16 Suction GB27 Îlot Caret 16°21.26’ 61°37.79’ – 2 Brushing GN27 Port-Louis 16°23’ 61°33’ – 120 Trap lines GR28 Port-Louis 16°27.34’ 61°32.07’ – 19 Visual at diving GM29 Grand Cul-de-sac marin 16°17.42’ 61°33.19’ – 1 Visual at intertidal GS29 Near Caret 16°22.28’ 61°38.14’ – 29 Suction GB30 Caye Plate 16°10.97’ 61°32.41’ – 16 Brushing Appendix 1. — Continuation. 503 Triphoridae from Martinique ZOOSYSTEMA • 2024 • 46 (18) Station Locality Latitude (N) Longitude (W) Initial depth (m) Final depth (m) Sampling method GS30 Îlot Caret 16°21.26’ 61°37.79’ – 2 Suction GB31 Petite Terre 16°09.71’ 61°07.73’ – 15 Brushing GD31 Port-Louis 16°24.97’ 61°32.8’ – 85 Dredging GD32 Port-Louis 16°23.88’ 61°32.47’ – 80 Dredging GS32 W Fajou 16°21.35’ 61°35.79’ – 2 Suction GD33 Port-Louis 16°24’ 61°33’ – 130 Dredging GM33 Anse Tarare 16°15.37’ 61°11.92’ – 6 Visual at diving GS33 Caye Plate 16°10.97’ 61°32.41’ – 14 Suction GB34 Petite Terre 16°10.45’ 61°08.16’ – 10 Brushing GR34 Pointe de la Fontaine 16°27.74’ 61°31.84’ – 9 Visual at diving GS34 Petite Terre 16°09.71’ 61°07.73’ – 15 Suction GD35 Port-Louis 16°22.77’ 61°34.19’ – 66 Dredging GB36 W Petite Terre 16°09.45’ 61°10.5’ – 16 Brushing GD36 Vieux-Bourg 16°22.87’ 61°33.05’ – 65 Dredging GR36 Pointe Montagnier 16°30.57’ 61°28.45’ – 12 Visual at diving GS36 Petite Terre 16°07.87’ 61°12.52’ – 50 Suction GR37 Grotte Amédier 16°30.04’ 61°28.79’ – 16 Visual at diving GS37 Petite Terre 16°10.45’ 61°08.16’ – 10 Suction GD39 Grand Cul-de-sac marin 16°17.34’ 61°33.32’ – 1 Dredging GS39 W Petite Terre 16°09.45’ 61°10.5’ – 16 Suction GD42 Grand Cul-de-sac marin 16°21.12’ 61°32.33’ – 3 Dredging GD49 Petit Cul-de-sac marin 16°13.62’ 61°32.38’ – 3 Dredging GD50 Îlet Gosier 16°11.31’ 61°29.59’ – 22 Dredging GR50 Lagon de Petite Terre 16°10.55’ 61°06.67’ – 4 Visual at diving GD51 Îlet Gosier 16°11.44’ 61°29.34’ – 15 Dredging GD52 Îlet Gosier 16°12.09’ 61°29.71’ – 6 Dredging GD54 Near Îlet Fajou 16°22.2’ 61°35.01’ – 60 Dredging GD55 Near Îlet Fajou 16°22.48’ 61°35.46’ – 85 Dredging GD58 Near Îlet Fajou 16°22.68’ 61°34.95’ – 95 Dredging GD59 Near Îlet Fajou 16°22.55’ 61°35.38’ – 88 Dredging GD61 E Petite Terre 16°11.97’ 61°03.96’ – 80 Dredging GD65 Banc des Vaisseaux 16°08.14’ 61°16.96’ – 20 Dredging GD68 Banc des Vaisseaux 16°09.03’ 61°17.31’ – 33 Dredging GD69 Anse Tarare 16°15.97’ 61°10.18’ – 60 Dredging KARUBENTHOS 2 DW4545 N Grande-Terre 16°29.7’ 61°31.4’ 60 82 Dredging DW4546 N Grande-Terre 16°31’ 61°32’ 268 306 Dredging DW4550 N Grande-Terre 16°37’ 61°31’ 432 482 Dredging DW4551 N Grande-Terre 16°31.3’ 61°24.5’ 59 74 Dredging DW4552 E La Désirade 16°23.9’ 60°56.2’ 48 60 Dredging DW4555 E La Désirade 16°24’ 60°51’ 100 258 Dredging DW4574 E La Désirade 16°22’ 60°54’ 140 340 Dredging DW4583 N Marie-Galante 16°09’ 61°19’ 233 396 Dredging DW4586 W Marie-Galante 16°00’ 61°23’ 204 251 Dredging DW4587 W Marie-Galante 16°01’ 61°24’ 281 406 Dredging DW4599 W Marie-Galante 15°53’ 61°25’ 262 266 Dredging Appendix 1. — Continuation.