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European Journal of Taxonomy 851: 1–141 ISSN 2118-9773
https://doi.org/10.5852/ejt.2022.851.1999 www.europeanjournaloftaxonomy.eu
2022 · Sabroux R. et al.
This work is licensed under a Creative Commons Attribution License (CC BY 4.0).
Research article
urn:lsid:zoobank.org:pub:7317EA8C-7C05-4E24-A38C-30F860013694
Sea spiders (Arthropoda: Pycnogonida) collected during the
Madibenthos Expedition from Martinique shallow waters
Romain SABROUX 1,*, Alexandre HASSANIN 2 & Laure CORBARI 3
1,2,3 Institut de Systématique, Évolution, Biodiversité (ISYEB), Muséum national d’Histoire naturelle
CNRS, Sorbonne Université, EPHE, UA. 57 rue Cuvier, CP 51, 75005 Paris, France.
* Corresponding author: pycnogonides@gmail.com
2 Email: alexandre.hassanin@mnhn.fr
3 Email: corbari@mnhn.fr
1 urn:lsid:zoobank.org:author:F48B4ABE-06BD-41B1-B856-A12BE97F9653
2 urn:lsid:zoobank.org:author:6CAA213C-384A-4111-8F70-1E8E07D3347D
3 urn:lsid:zoobank.org:author:9E5EBA7B-C2F2-4F30-9FD5-1A0E49924F13
Abstract. This study presents the inventory of sea spiders (Pycnogonida) sampled during the Madibenthos
Expedition in Martinique (West Indies). Species were discriminated leaning on morphological and
Thirteen new species are described: Ammothella dirbergi sp. nov., A. krappi sp. nov., Tanystylum boucheti
sp. nov., T. ingrallis sp. nov., Ascorhynchus iguanarum sp. nov., Eurycyde kaiouti sp. nov., Nymphon
dorlis sp. nov., N. ludovici sp. nov., N. martinicum sp. nov., N. timons sp. nov., Anoplodactylus
madibenthos sp. nov., Pycnogonum cesairei sp. nov. and Rhynchothorax sidereus sp. nov. We describe
a neotype for Anoplodactylus micros Bourdillon, 1955 from the type locality. Martinique now includes
79 species of sea spiders, mostly endemic to the Tropical Northwestern Atlantic, cosmopolitan or
shared with the South America Atlantic coast. Some species are potentially introduced. However, our
knowledge of the distribution of species found in Martinique is probably biased by the scarcity of
diagnostic morphological characters. Also, nine potentially cryptic species (discriminated on genetic
Northwestern Atlantic. Therefore, we call for a more widespread use of barcoding in sea spiders.
Keywords. Pantopoda, Caribbean, West Indies, biodiversity, integrative taxonomy.
Sabroux R., Hassanin A. & Corbari L. 2022. Sea spiders (Arthropoda: Pycnogonida) collected during the
Madibenthos Expedition from Martinique shallow waters. European Journal of Taxonomy 851: 1–141.
https://doi.org/10.5852/ejt.2022.851.1999
Introduction
Sea spiders (Arthropoda: Pycnogonida) of the Tropical Northwestern Atlantic (TNWA; Spalding et al.
2007) have been the focus of about 45 publications (listed in Sabroux et al. 2019b), with 132 species
known from the region. Despite that this diversity already represents about 9% of the total of known
species of Pycnogonida, we demonstrated in a previous account of Martinique fauna based on the
European Journal of Taxonomy 851: 1–141 (2022)
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Madibenthos Expedition samples (Sabroux et al. 2019b) how lacunar our knowledge of TNWA sea
spiders is regarding all together a high rate of undescribed species and the poor knowledge of species
repartition. Species delimitation analyses performed with DNA barcoding data also give a hint on how
abyssal the unexplored pit of cryptic species is.
While our previous work focused on the macroecology of Martinique’s sea spiders and the knowledge
bias regarding the Caribbean fauna, this study will present the taxonomic diversity of sea spiders
collected during the Madibenthos Expedition in detail. This study presents several improvements
from the former listing (recounting, updated taxonomic status), provides a detailed list of specimens
and introduces the description of 13 new species. A total of 72 species is listed from the Madibenthos
samples. This shallow water material from Martinique adds to the contributions of Bourdillon (1955)
and Müller (1990a), which already included 20 species (of which 13 are recovered in the Madibenthos
material; Table 1), including one undetermined species of Ascorhynchus and the so-far endemic species
Nymphon macabou Müller, 1990, which was not recovered in the Madibenthos samples. It is also worth
noting that two deep-water species were sampled by the research vessel ‘Blake’ in 1879 and described
Ephyrogymna circularis Hedgpeth, 1943 (1052 m) and
Hedgpeth, 1943 (871 m).
Material and methods
Studied material
The material was collected in Martinique’s shallow waters (mostly 0–100 m) during the Madibenthos
expedition in September–October 2016. Sampling was performed by scuba-diving methods (catching,
vacuum sampling, brushing sampling), sampling along the intertidal zone and dredging. Sampling
stations with sea spider occurrences are presented in Fig. 1.
in Sabroux et al. (2019b) for most specimens examined herein in order to resolve taxonomic issues.
species, with corresponding names in Sabroux et al. (2019b), is provided in Table 2.
Types and examined material are deposited in the Muséum national d’Histoire naturelle, Paris
(collection numbers MNHN-IU-xxx-xxx). MK numbers correspond to GenBank accession numbers
published in Sabroux et al. (2019b). Measurements of described holotypes were taken on drawings
which were produced using a camera lucida; the trunk was measured from the chelifore insertion to the
measurements were performed measuring the distance from the proximal insertion to the tip.
Acronyms for the collections in which previously described material is hosted are as follow:
EMB-IOC = Estação de Biologia Marinha do Instituto Oswaldo Cruz, Rio de Janeiro, Brazil
MCZ = Museum of Comparative Zoology, Harvard University, Boston, MA, USA (Invertebrate
Zoology Collection)
MNHN = Muséum national d’Histoire naturelle, Paris, France (Invertébrés marins, Collection
Crustacés)
MOM = Musée Océanographique de Monaco, Monte-Carlo, Monaco
MP = Museu Paranaense, Curitiba, Paraná, Brazil
NHMD = Natural History Museum of Denmark, Copenhagen (Crustacea collection)
NHMUK = Natural History Museum, London, UK (LS Chelicerates)
NL = Naturalis Biodiversity Center, Leiden, the Netherlands (Collection Chelicerata and
Myriapoda)
SABROUX R. et al., Pycnogonida from the Madibenthos Expedition to Martinique
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Table 1. List of the 20 species previously recorded from Martinique. Precise localities and references are indicated.
Family Genus Species Localities Reference Recovered in
present study
Ammotheidae
Dohrn, 1881
Achelia
Hodge, 1864
Achelia gracilis Verrill, 1900 Fort-de-France port Bourdillon 1955 X
Achelia sawayai Marcus, 1940 Cap Chevalier Müller 1990e X
Ammothella
Verrill, 1900
Ammothella appendiculata (Dohrn, 1881) Petite Anse du Diamant Bourdillon 1955 X
Ammothella exornata Stock, 1975 Ilet Cabrits, Petite Anse Macabou, Baie de
Tartane
Müller 1990e X
Ammothella rugulosa Verrill, 1900 Fort-de-France port Bourdillon 1955
Ammothella marcusi Hedgpeth, 1948 Baie de Tartane Müller 1990e X
Ammothella spinifera Cole, 1904 Vauclin, Petite Anse Macabou, Baie de
Tartane
Müller 1990e X
Tanystylum
Miers, 1879
Tanystylum acuminatum Stock, 1954 Petite Anse Macabou Müller 1990e X
Tanystylum birkelandi Child, 1979 Petite Anse Macabou Müller 1990e X
Tanystylum geminum Stock, 1954 Ilet Cabrit, Petite Anse Macabou, Baie de
Tartane
Bourdillon 1955,
Müller 1990e
Tanystylum isthmiacum Stock, 1955 Petite Anse Macabou Müller 1990e
Tanystylum tayronae Müller & Krapp, 2009 Petite Anse Macabou Müller 1990e,
Müller & Krapp 2009
X
Ascorhynchidae
Hoek, 1881
Ascorhynchus
Sars, 1877
Ascorhynchus sp. Fort-de-France port Bourdillon 1955
Callipallenidae
Hilton, 1942
Callipallene
Flynn, 1929
Callipallene emaciata (Dohrn, 1881) Petite Anse Macabou Müller 1990e
Nymphonidae
Wilson, 1878
Nymphon
Fabricius, 1794
Nymphon macabou Müller, 1990 Petite Anse Macabou Müller 1990e
Phoxichilidiidae
Sars, 1891
Anoplodactylus
Wilson, 1878
Anoplodactylus batangensis (Helfer, 1938) Petite Anse du Diamant, Ilet Cabrits, Cap
Chevalier, Petite Anse Macabou, Baie de
Tartane
Bourdillon 1955,
Müller 1990e
X
Anoplodactylus californicus Hall, 1912 Saint-Pierre, Petite Anse Macabou Bourdillon 1955,
Müller 1990e
(cf.)
Anoplodactylus digitatus (Böhm, 1879) Baie de Fort-de-France Bourdillon 1955 X
Anoplodactylus micros Bourdillon, 1955 Petite Anse Macabou Bourdillon 1955 X
Anoplodactylus monotrema Stock, 1979 Petite Anse Macabou Müller 1990e X
European Journal of Taxonomy 851: 1–141 (2022)
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SMF = Senckenberg-Museum Frankfurt am Main, Germany (Marine Zoology, Crustacean
Collections)
UFPB = Universidade Federal da Paraíba, Brazil (Pycnogonida collection)
USNM = Smithsonian National Museum of Natural History, Washington D.C., USA (Invertebrate
Zoology Collections)
YPM = Yale Peabody Museum of Natural History, New Haven, CT, USA (Invertebrate Zoology
Dept)
ZMH = Zoological Museum Hamburg, Germany (Zoological collections, Arachnology)
ZMB = Museum für Naturkunde, Berlin, Germany (Arachnida, Myriapoda and stem-group
Arthropoda)
Fig. 1. Madibenthos sampling stations with occurrences of sea spiders in Martinique. Bathymetric range
is shown by grey isolines each 20 m (0–100 m depth) and each 500 m (deeper than 500 m). The 100 m
isobath (above which most samplings were performed) is marked by a black isoline. Sampling stations
are indicated as black circles. See Sabroux et al. (2019b) for alternative representation.
SABROUX R. et al., Pycnogonida from the Madibenthos Expedition to Martinique
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Table 2 (continued on next 2 pages). List of species recorded from the Madibenthos Expedition. Corresponding names in Sabroux et al. (2019b) are
indicated, as well as the number of suspected cryptic species. In column with number of potential cryptic species, the COI inter-cluster uncorrected
p-distance is given in parentheses.
Family Genus Species Name in Sabroux et al. 2019b Potential cryptic
species within
Ammotheidae
Dohrn, 1881
Achelia
Hodge, 1864
Achelia gracilis Verrill, 1900 2 (0.105–0.112)
Achelia sawayai Marcus, 1940 3 (0.133–0.191)
Ammothella
Verrill, 1900
Ammothella appendiculata (Dohrn, 1881)
Ammothella dirbergi sp. nov. Ammothella sp. 1 [pro parte]
Ammothella exornata Stock, 1975
Ammothella krappi sp. nov. Ammothella sp. 1 [pro parte]
Ammothella marcusi Hedgpeth, 1948
Ammothella . krappi sp. nov. Ammothella sp. 1 [pro parte]
Ammothella spinifera Cole, 1904
Nymphopsis
Haswell, 1884
Nymphopsis duodorsospinosa Hilton, 1942
Tanystylum
Miers, 1879
Tanystylum acuminatum Stock, 1954
Tanystylum boucheti sp. nov. Tanystylum hummelincki Stock, 1954 [pro parte]
Tanystylum birkelandi Child, 1979
Tanystylum duospinum Hilton, 1939
Tanystylum hummelincki Stock, 1954 Tanystylum hummelincki Stock, 1954 [pro parte]
Tanystylum ingrallis sp. nov. Tanystylum sp. 3
Tanystylum orbiculare Wilson, 1878 Tanystylum conirostre (Dohrn, 1881)
Tanystylum tayronae Müller & Krapp, 2009
Ascorhynchidae
Hoek, 1881
Ascorhynchus
Sars, 1877
Ascorhynchus castellioides Stock, 1957
Ascorhynchus horologium Child, 1992
Ascorhynchus iguanarum sp. nov. Ascorhynchus sp. 4
Ascorhynchus latipes (Cole, 1906)
Ascorhynchus sp. 5
Eurycyde
Schiödte, 1857
Eurycyde clitellaria Stock, 1955
Eurycyde kaiouti sp. nov. Eurycyde raphiaster Loman, 1912 [pro parte]
Eurycyde raphiaster Loman, 1912 Eurycyde raphiaster Loman, 1912 [pro parte]
European Journal of Taxonomy 851: 1–141 (2022)
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Table 2 (continued). List of species recorded from the Madibenthos Expedition. Corresponding names in Sabroux et al. (2019b) are indicated, as well
as the number of suspected cryptic species. In column with number of potential cryptic species, the COI inter-cluster uncorrected p-distance is given
in parentheses.
Callipallenidae
Hilton, 1942
Callipallene
Flynn, 1929
Callipallene brevirostris (Johnston, 1837)
Callipallene cinto Müller & Krapp, 2009
Callipallene longicoxa Stock, 1955
Pallenoides
Stock, 1951
Pallenoides cf. amazonicus Stock, 1974 Pallenoides sp. 1
Pallenoides spinulosum Stock, 1955
Parapallene
Carpenter, 1892
Parapallene bermudensis Lebour, 1949
Endeidae
Norman, 1908
Endeis Philippi,
1843
Calman, 1923
Endeis mollis (Carpenter, 1904) Endeis cf. mollis (Carpenter, 1904)
Endeis meridionalis (Böhm, 1879) Endeis meridionalis (Böhm, 1879)
Endeis sp. 3
Nymphonidae
Wilson, 1878
Nymphon
Fabricius, 1794
Nymphon aemulum Stock, 1975
Nymphon dorlis sp. nov. Nymphon sp. 1
Nymphon ludovici sp. nov. Nymphon sp. 3 2 (0.104–0.111)
Nymphon martinicum sp. nov. Nymphon sp. 4
Nymphon timons sp. nov. Nymphon sp. 2
Pallenopsidae
Fry, 1978
Pallenopsis
Wilson, 1881
Pallenopsis candidoi Mello-Leitao, 1949
Pallenopsis schmitti Hedgpeth, 1943
Phoxichilidiidae
Sars, 1891
Anoplodactylus
Wilson, 1878
Anoplodactylus cf. arcuatus Child, 1977
Anoplodactylus batangensis (Helfer, 1938)
Anoplodactylus bahamensis Child, 1977 Anoplodactylus micros Bourdillon, 1955 [pro parte]
Anoplodactylus cf. californicus Hall, 1912 Anoplodactylus sp. 2
Anoplodactylus digitatus (Böhm, 1879)
Anoplodactylus evelinae Marcus, 1940
Anoplodactylus ganchiformis Anoplodactylus stictus Marcus, 1940
Anoplodactylus glandulifer Stock, 1954
Anoplodactylus imswe Child, 1982
Family Genus Species Name in Sabroux et al. 2019b Potential cryptic
species within
SABROUX R. et al., Pycnogonida from the Madibenthos Expedition to Martinique
7
Table 2 (continued). List of species recorded from the Madibenthos Expedition. Corresponding names in Sabroux et al. (2019b) are indicated, as well
as the number of suspected cryptic species. In column with number of potential cryptic species, the COI inter-cluster uncorrected p-distance is given
in parentheses.
Phoxichilidiidae
Sars, 1891 (cont.)
Anoplodactylus
Wilson, 1878
(cont.)
Anoplodactylus insignis (Hoek, 1881)
Anoplodactylus justi Müller, 1992 ? Anoplodactylus cf. justi Müller, 1992
Anoplodactylus madibenthos sp. nov. Anoplodactylus sp. 1
Anoplodactylus maritimus Hodgson, 1914
Anoplodactylus massiliformis Stock, 1975
Anoplodactylus micros Bourdillon, 1955 Anoplodactylus micros Bourdillon, 1955 [pro parte] 2 (0.077–0.1)
Anoplodactylus monotrema Stock, 1979
Anoplodactylus pectinus Hedgpeth, 1948
Anoplodactylus petiolatus (Krøyer, 1844)
Anoplodactylus quadratispinosus Hedgpeth, 1943
Anoplodactylus robustus (Dohrn, 1881)
Anoplodactylus sp. 4
Anoplodactylus sp. 7
Anoplodactylus sp. 8
Pycnogonidae
Wilson, 1878
Pentapycnon
Bouvier, 1910
Pentapycnon geayi Bouvier, 1911
Pycnogonum
Brünnich, 1764
Pycnogonum cesairei sp. nov. Pycnogonum sp. 1
Pycnogonum cf. ornans Stock, 1992
Pycnogonum cf. pusillum Dohrn, 1881
Rhynchothoracidae
D’Arcy Thompson,
1909
Rhynchothorax
Costa, 1861
Rhynchothorax crenatus Child, 1982
Rhynchothorax sidereus sp. nov. Rhynchothorax sp. 1
Family Genus Species Name in Sabroux et al. 2019b Potential cryptic
species within
European Journal of Taxonomy 851: 1–141 (2022)
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Molecular distances
Molecular distances were calculated as uncorrected p-distances among CO1 sequences made available
by Sabroux et al. (2019b). These calculations were performed with MEGA7 (Kumar et al. 2016), using
pairwise deletion for missing data.
Integrative taxonomy
An integrative taxonomic approach, based on comparisons between morphological and molecular data,
for mitochondrial marker of the cytochrome oxidase subunit 1 (CO1) gene. Species delimitation was
then performed using the ABGD method (Puillandre et al. 2012). When the barcoding method suggested
detail. Following the recommandations of Dietz et al. (2015a), only groups which were discriminated
suggested by molecular data alone and we could not identify discriminating morphological characters
Results (description authority: Sabroux)
et al. (2019b) are
due to a very few specimens that were recounted, damaged or newly found in unsorted samples, etc.
The 9 families, 16 genera and 72 species recorded from the Madibenthos Expedition are listed in Table 2.
Unless otherwise mentioned, all material was collected during the Madibenthos Expedition, Martinique
(West Indies). Station codes indicate the sampling method (AB for brushing, AD for dredging, AM for
intertidal catches, AR for sight catches, AS for vaccum) (see Bouchet et al. 2019).
or development could not be determined.
Subphylum Chelicerata Heymons, 1901
Class Pycnogonida Latreille, 1810
Order Pantopoda Gerstaecker, 1863
Family Ammotheidae Dohrn, 1881
Subfamily Achelinae Wilson, 1881
Genus Achelia Hodge, 1864
Type species
Achelia echinata Hodge, 1864, by subsequent designation (Child 1998a).
Achelia gracilis Verrill, 1900
Achelia (?) gracilis
Ammothea gracilis –
Ammothea (Achelia) gracilis – Giltay 1934b: 5.
SABROUX R. et al., Pycnogonida from the Madibenthos Expedition to Martinique
9
Achelia gracilis –
Varela 2012Lucena et al. 2019: 3 (list),
20––Sabroux et al. 2019 León-Espinosa et al. 2021:
Ramírez-Tello et al. 2022: tab. 1.
Achelia (Pigrolavatus) gracilis –
Type material
Holotype: YPM IZ 003366.CR (not examined). Type locality: Flatts Inlet, Bermuda.
Material examined
depth 20 m; 20 Sep. 2016; st. AB260; MNHN-IU-2016-1062/MK411040 • 1 juv.; Les Anses-d’Arlet;
1180/
depth 10 m; 19 Sep. 2016; st. AB195; MNHN-
IU-2016-
1 juv.; same collection data as for preceding; MNHN-IU-2016-1384.
Remarks
This species was distinguished from Achelia sawayai Marcus, 1940 based on the key in Müller & Krapp
(2009), relying on the number of palp articles (7 in A. gracilis, 8 in A. sawayai) and the number of
A. gracilis, 3 in A. sawayai).
Achelia gracilis was previously recorded from Martinique by Bourdillon (1955) on the hulls of scows
in Fort-de-France port. Madibenthos sampling reveals that this species is present on both the Atlantic
and Caribbean coasts of Martinique. Achelia gracilis is a common species in the Caribbean, almost as
common as its close congener, Achelia sawayai (Child 1996a).
DNA barcoding data (Sabroux et al. 2019b) suggest that this species name hides two potential
cryptic species (intracluster p-distance = 0–0.010; intercluster p-distance = 0.105–0.112), here named
group 1 (MNHN-IU-2016-849, -851, -857, 881, -888, -1114, -1180, -1276, -1379 and -1380) and group 2
(MNHN-IU-2016-1062 and -1330) (Table 2 and Appendix).
Distribution
Tropical West Atlantic (Bermuda, Caribbean and Brazil).
Depth range
0–18 m. One record at 157 m is regarded as a likely contamination by shallow fouling on the hull of a
ship. We extend this depth-range to 28 m.
European Journal of Taxonomy 851: 1–141 (2022)
10
Achelia sawayai Marcus, 1940
Achelia sawayai
Achelia sawayai –
: 497; 1979: 7–8; 1982a: 356–357; 1992b: 7 (key), 11–12,
Birkeland et al.
Kraeuter 1976: 342–1990a: 277; 1990b: 186–188; 1990e:
105–106;–
et al. 2009: 10,
Müller & Krapp 2009: 10–12, 14 (key), 17–21, 132–
4–et al.
Sabroux et al. 2017: appendix 1–2019Lucena &
Lucena et al. 2019: 3 (list), 4–5, 20– Prata et al. 2020:
–cLeón-Espinosa et al. 2021Ramírez-Tello et al. 2022:
152, 162, tab. 1.
Achelia Sawayai – Fage 1949: 28.
Achelia (Pigroglavatus) sawayai –
155 (distribution).
Achelia sawayai f. typica – Stock 1986: tab. 1.
nec Tanystylum calcirostre Schimkewitsch, 1890 – Kraeuter 1973: 496.
Type material
Brazil.
Material examined
MARTINIQUE • depth 65 m; 11 Sep. 2016; st. AD223;
depth 3 m; 15 Sep.
same collection data as for preceding;
MNHN-IU-2016-
MK411103 depth 2 m; 24 Sep. 2016; st. AB452; MNHN-
same collection data as for preceding; MNHN-IU-2016-885/MK411194 •
data as for preceding; MNHN-IU-2016-1378 depth
65 m; 11 Sep. 2016; st. AD223; MNHN-IU-2016-886/MK411195
SABROUX R. et al., Pycnogonida from the Madibenthos Expedition to Martinique
11
MNHN-IU-2016-1383/MK411109.
Remarks
DNA barcoding (Sabroux et al. 2019) suggests that A. sawayai is an umbrella-name including three
potential cryptic species (intracluster p-distance = 0–0.013; intercluster p-distance = 0.133–0.191) here
designated as group 1 (MNHN-IU-2016-859 and -1329), group 2 (MNHN-IU-2016-850, -886, -1108,
-1136, -1331 and -1332) and group 3 (MNHN-IU-2016-884, -885, -1073 and -1383) (Table 2, Appendix).
Achelia sawayai was previously recorded from Martinique by Müller (1990a) at Cap Chevalier (South-
East Martinique), and the present sampling reveals that this species is found on both the Atlantic and
Caribbean coasts. The three possible cryptic species here detected in Martinique suggest that A. sawayai
may represent a species complex including many more species. Indeed, it is a common species of the
Tropical Western Atlantic, and its distribution extends to the Eastern Atlantic, the Mediterranean, the
Distribution
Apparently a circumtropical species, mostly found in the West Atlantic (US Georgia, Caribbean, Gulf of
Mexico and Brazil), occasionally recorded in the East Atlantic (Gulf of Guinea, Mauritania, Cap Verde),
Ocean (French Polynesia, Fiji).
Depth range
0–115 m. One record at 157 m is regarded as a likely contamination by shallow fouling on the hull of a
ship.
Genus Ammothella Verrill, 1900
Type species
Ammothea (Ammothella) rugulosa Verrill, 1900, by original designation.
Ammothella appendiculata (Dohrn, 1881)
Ammothea appendiculata Dohrn, 1881: 7, 15, 18, 22, 24, 49, 51, 135 (text, key), 152–155, 159, 165, pl.
Ammothella appendiculata
–b;
2004: 149 (key); et al.
Arnaud 1987: 39––
Bamber 1997: 144– –622; 2004: 2–
et al. 2006: 88–89;
et al.
et al. 2008:
Arabi et al.
European Journal of Taxonomy 851: 1–141 (2022)
12
–
Lehmann et al.
Soler-Membrives 2014: 95 (key), 96–
Dietz et al.
et al.et al. 2019b: 1531, tab. 1,
et al. 2020: tab. 1. Ramírez-Tello et al. 2022: 165, tab. 3.
Ammothea appendiculata –
Ammothea (Ammothella) appendiculata
Type material
Syntypes (?) (see Stock 1955; Dunlop et al. 2007): ZMB_Pyc_46 (not examined). Type locality: Santa
Lucia, Naples Bay, Italy.
Material examined
MARTINIQUE • depth 0–2 m; 12 Sep. 2016; st.
AM012; MNHN-IU-2016-1278/MK411096 • 1 juv.; same collection data as for preceding; MNHN-
IU-2016- same collection data as for preceding; MNHN-IU-2016-1573 •
depth 0–1 m; 9 Oct. 2016; st. AM325; MNHN-
depth 10–17 m; 3 Oct. 2016; st. AR383;
MNHN-IU-2017-220.
Remarks
This species was already recorded in Martinique by Bourdillon (1955) from the Petite Anse du Diamant,
and was sampled during Madibenthos from both the Atlantic and Caribbean coasts. It is commonly
sampled in the Atlantic, and available illustrations (e.g., Dohrn 1881; Stock 1955; Child 1982a; Müller &
and a stout form co-exist, which he suspected to correspond to two adult stages separated by a moult,
though this theory was proven invalid by observations of Child (1982a) on Belize material. Madibenthos
material corresponds to the stout form and strongly resembles the drawing of Child (1992b); this stout
form is the one represented by the type material according to the drawings in Stock (1955) (although
the type status of this material is not totally clear). The slender form mentioned by Stock (1955) may
correspond to one of the hereafter newly described species, Ammothella dirbergi sp. nov. or A. krappi
sp. nov., or to A. krappi
CO1 data support the distinction of these three species from A. appendiculata (Sabroux et al. 2019b).
The p-distances between A. appendiculata and these species are very high and vary between 0.172 and
0.197 (see Appendix).
Another harsh debate regarding this species is the possible synonymy of Ammothella appendiculata and
A. rugulosa. Lucena et al. (2019)
their appendages.
Distribution
Rather cosmopolitan, with records in the West and East Mediterranean, West Atlantic (Florida, Gulf of
Caledonia).
Depth range
0–76 m.
SABROUX R. et al., Pycnogonida from the Madibenthos Expedition to Martinique
13
Ammothella dirbergi sp. nov.
urn:lsid:zoobank.org:act:7CCAA126-D880-4F80-B6D7-A7CB0C532E6C
Fig. 2
Amothella sp. 1 – Sabroux et al.
Material examined
Holotype
MNHN-IU-2016-833/MK411148.
Paratypes
same
collection data as for preceding; MNHN-IU-2016-
1 Oct. 2016; st. AD283; MNHN-IU-2016-1054 • 1 juv.; same collection data as for preceding; MNHN-
IU-2016-1224
AB388; MNHN-IU-2016-1058 • 2 juvs; same collection data as for preceding; MNHN-IU-2016-1308 •
MNHN-IU-2016- depth 24 m;
depth 9 m;
depth 16 m; 18 Sep. 2016; st. AS255; MNHN-IU-2016-1176
preceding; MNHN-IU-2016-1315 depth 44–47 m;
depth
depth
depth same
collection data as for preceding; MNHN-IU-2016-1239
AR383; MNHN-IU-2016-1306.
Etymology
Latinized name, 2nd decl. (masc.), genitive singular. This species is named after Guillaume Dirberg
(MNHN), who collected many specimens of this species by sight, which is remarkable when considering
the inconspicuousity of these animals.
European Journal of Taxonomy 851: 1–141 (2022)
14
Description
Trunk completely segmented, cuticle smooth. No dorsomedian ornamentation. Ocular tubercle
long, about 5.5 times as long as base width, distal part broad and ovoid carrying pigmented eyes, pointy
tip and 2 small pointy lateral sense organs (see Lehmann et al. 2017; Brenneis 2022). Lateral processes
about twice as long as wide, well separated by about half of their own diameter, unornamented.
Pyriform, movable, longer than chelifore or trunk, with blunt tip.
club-shaped setae and simple setae medially and distally. Abdomen insertion articled on trunk.
3-articled. Scape 2-articled, 1st scape article about half as long as 2nd. 2nd article long,
trumpet-shaped, carrying simple and club-shaped setae. Chela reduced as rounded buds, with distal seta
on outer side.
9-articled, carrying many setae mostly on last 5 articles. 1st article shortest, 2nd slightly shorter than
4th. 3rd article about as long as wide. 4th article longest, about as long as 2nd, about 6 times as long as wide,
with scarce setae. 5th article about 2.5 times as long as wide. 6th article shorter than 5th and twice as long
as 8th, about 4.5 times as long as wide. 7th to 9th articles shorter than 6th, 9th slightly longer.
10-articled, with scarce setae. 1st article as long as wide. 2nd about 5 times as long as wide, 3rd
about 3 times as long as wide. 4th nd. 5th article
th article. 7th article longest within striglis, 10th shortest. Strigilis
Slender, carrying several short, simple and club-shaped setae, some of them in bouquet. Coxa
1 about 1.5 times as long as wide, with 2 dorsodistal club-shaped setae and often 2 tiny normal setae
laterally. Coxa 2 subequal to coxae 1 and 3 together. Coxa 3 about twice as long as wide. Femur 6 times
as long as wide; cement gland opening standing on distal margin on dorsal surface, as long tube about
spines and setae on distal half of sole; main claw about half as long as propodus; auxiliary claws present,
thinner, about same size as main claw.
(mm). Trunk 0.54; abdomen 0.62; proboscis 0.70; chelifore 0.47; coxa 1 0.14; coxa 2
0.40; coxa 3 0.30; femur 0.84; tibia 1 1.03; tibia 2 0.92; tarsus 0.05; propodus 0.34; main claw 0.17.
Sexual dimorphism
Individual variability
Number of ovigeral spines variable, 1 to 4 on 1st strigilis article, 1 or 2 on 2nd strigilis article, 1 or 2 on
3rd strigilis article. Lateral processes variable in length. Club-shaped setae sometimes slightly cleft, as
on coxa 1 of left 2nd leg of holotype.
Remarks
Ammothella indica Stock, 1954, which is a common species in
group of Ammothella with club-shaped spines, and the lengths of the ocular tubercle, of leg and chelifore
articles, and of the cement gland tube in males, are about the same. The relative lengths of palp articles
SABROUX R. et al., Pycnogonida from the Madibenthos Expedition to Martinique
15
Fig. 2. Ammothella dirbergi sp. nov. A–F, I–L G–H. Paratype,
A. Body, dorsal view. B. Body, lateral view (same scale as A). C. Ocular
tubercle, frontal view. D. Chelifore. E. Chela bud. F. Palp. G. Oviger. H. Oviger strigilis. I. Oviger.
J. Oviger strigilis. K. Third leg. L. Tarsus and propodus of third leg.
European Journal of Taxonomy 851: 1–141 (2022)
16
A. indica and A. dirbergi sp. nov. are the following: (i) lateral sense organs are inconspicuous in
A. indica; (ii) the abdomen of A. dirbergi does not present a diadem of setae as in A. indica; and (iii) the
abdomen of A. dirbergi is not as curved as in A. indica.
Ammothella based on the long
ocular tubercle and the club-shaped spines on the legs and abdomen. Species sharing these features are
Lee & Arango, 2003, A. gertrudae Müller & Krapp, 2009, A. panamensis Child,
2004, A. prolixa Child, 1990 and A. setacea (Helfer, 1938). Ammothella dirbergi
in lacking tubular spines on the lateral processes and cephalon, and by its longer ocular
A. gertrudae in lacking long tubercles on the lateral processes;
A. panamensis in lacking club-shaped setae on the lateral processes, has longer tibiae
A. prolixa in having a shorter distance between the
A. setacea in
lacking long tubercles on the lateral processes and in having a shorter ocular tubercle and a completely
segmented trunk (Arnaud & Child 1988; Child 1990, 2004; Lee & Arango 2003; Müller & Krapp 2009).
Finally, Ammothella krappi sp. nov. is a close relative of A. dirbergi sp. nov. Ammothella dirbergi
A. dibergi, more than 7
times in A. krappiA. dirbergi, half as long
in A. krappi). The two species can be distinguished based on CO1 data (Sabroux et al. 2019): while
in A. dirbergi
A. dirbergi and A. krappi are between 0.123 and 0.134, and those between A. dirbergi and A. krappi
are between 0.089 and 0.097.
Distribution
Only known from Martinique. Ammothella dirbergi was sampled on the Atlantic and Caribbean coasts.
Depth range
0.5–47 m.
Ammothella exornata Stock, 1975
Ammothella exornata
Ammothella exornata – Stock19792004
et al.
et al. 2019b:
Type material
Holotype: NL ZMA.PYC.P.1062 (not examined). Type locality: southeast of Oyster Pond, St Martin.
Material examined
MARTINIQUE • depth 7 m; 9 Sep. 2016; st.
depth
0–2 m; 12 Sep. 2016; st. AM012; MNHN-IU-2016-
SABROUX R. et al., Pycnogonida from the Madibenthos Expedition to Martinique
17
• 1 juv.; same collection data
as for preceding; MNHN-IU-2016-1285
1098 • 1 juv.; Banc du
depth
depth
depth 12 m; 26 Sep. 2016; st. AB360; MNHN-
depth 0–1 m; 27 Sep. 2016; st.
AM033; MNHN-IU-2016-1464.
Remarks
Ammothella exornata was recorded from Martinique by Müller (1990a) from Ilet Cabrits, Petite Anse
Macabou and Baie de Tartane. We found it on both the Atlantic and Caribbean coasts.
Distribution
Exclusively in the Caribbean area.
Depth range
0–10 m. We extend this depth range to 18 m.
Ammothella krappi sp. nov.
urn:lsid:zoobank.org:act:016C5DC5-7B37-4A2F-B4CC-3176173A3C65
Fig. 3
Ammothella sp. 1 – Sabroux et al. 2019b
Material examined
Holotype
MARTINIQUE • depth 63 m, 21 Sep. 2016; st. AD257,
MNHN-IU-2016-1132/MK411059.
Paratypes
MARTINIQUE • depth 20–25 m; 8 Oct. 2016;
depth 66 m; 7
same collection data as for preceding;
depth 17 m; 9 Oct. 2016; st.
depth 80 m; 27 Sep.
depth 17–19 m;
depth
depth 44–47 m; 8 Oct. 2016; st. AS572; MNHN-IU-2019-3395.
Etymology
Latinized name, 2nd decl. (masc.), genitive singular. This species is named after Franz Krapp, who
passed away in early 2022. Through this dedication, we wish to express not only our recognition of his
author.
European Journal of Taxonomy 851: 1–141 (2022)
18
Description
Trunk completely segmented; cuticle smooth. No dorsomedian ornamentation. Ocular tubercle
long, about 8.5 times as long as base width, distal part broad and round, carrying pigmented eyes, blunt
tip and 2 small, rounded lateral sense organs. Lateral processes about 3 times as long as base width, well
separated by more than their own diameter, sometimes carrying setae.
Pyriform with slender elongated base, movable, longer than chelifore or trunk, with blunt
tip.
anus. Dorsally with club-shaped setae and with simple setae medially and distally.
3-articled. Scape 2-articled, 1st scape article about half as long as 2nd. 2nd article long,
distal seta on outer side.
9-articled, carrying many setae mostly on last 5 articles. 1st article shortest, 2nd article slightly
shorter than 4th. 3rd article about as long as wide. 4th article longest, about as long as 2nd, about 6 times as
long as wide, with scarce setae. 5th article about 2.5 times as long as wide. 6th article shorter than 5th and
twice as long as 8th, about 4.5 times as long as wide. 7th to 9th articles shorter than 6th, 9th longest among
them.
10-articled, with scarce setae. 1st article about as long as wide, 2nd about 7 times as long as wide,
3rd about 3 times. 4th article as long as 2nd but slenderer, slightly curved. 5th
4th article. 7th article longest within striglis, about as long as 6th, 10th shortest. Strigilis spines compound,
Slender, carrying several short, simple and club-shaped setae, some of them in bouquet. Coxa
laterally. Coxa 2 longer than coxae 1 and 3 together. Coxa 3 about 3 times as long as wide. Femur about
8 times as long as wide, cement gland opening standing on distal margin on dorsal surface, as very long
tube about half as long as femur. Tibia 1 longest. Tibia 2 slenderer, slightly shorter than tibia 1 but longer
than femur. Tarsus short, trapezoid. Propodus about 0.4 times as long as tibia 2, with 3 large basal spines,
and smaller spines and setae on distal half of sole; main claw about half as long as propodus; auxiliary
(mm). Trunk 0.52; abdomen 0.78; proboscis 0.92; chelifore 0.56; coxa 1 0.17; coxa 2
0.53; coxa 3 0.26; femur 0.91; tibia 1 1.13; tibia 2 0.96; tarsus 0.06; propodus 0.37; main claw 0.19.
Sexual dimorphism
Individual variability
Ocular tubercle size from 7 to 9 times as long as base width. Number of ovigeral spines variable, 1 to 4
on 1st strigilis article, and 1 or 2 on 2nd–3rd strigilis articles.
Remarks
This species is distinguished from most other species of Ammothella using the same criteria as for
A. dirbergi sp. nov., of which this species seems to be a close relative. Distinction from A. dirbergi is
supported by CO1 data (Sabroux et al. 2019b): while for A. krappi
SABROUX R. et al., Pycnogonida from the Madibenthos Expedition to Martinique
19
Fig. 3. Ammothella krappi sp. nov. A–G, J–K H–I. Paratype,
A. Body, dorsal view. B. Body, lateral view (same scale as A). C. Ocular
tubercle, frontal view. D. Chelifore. E. Third leg. F. Tarsus and propodus of third leg. G. Palp.
H. Oviger. I. Oviger strigilis. J. Oviger. K. Oviger strigilis.
European Journal of Taxonomy 851: 1–141 (2022)
20
between 0.011 and 0.036, the p-distances in A. dirbergi are between 0.123 and 0.134. Morphologically,
A. krappi has a longer ocular tubercle than A. dirbergi (more than seven times as long as wide in
A. krappi, less than six times as long as wide in A. dirbergi), the tip of the ocular tubercle is blunt rather
than acute, and the lateral sense organs are smaller than in A. dirbergi. The cement gland tube is also
longer in A. krappi, as well as the leg articles.
The species was sampled on the Caribbean and northern coasts of Martinique.
Distribution
Only known from Martinique.
Depth range
17–80 m.
Ammothellakrappi sp. nov.
Fig. 4
Ammothella sp. 1 – Sabroux et al. 2019b
Material examined
MARTINIQUE • depth 28 m; 7 Sep. 2016; st.
AB157; MNHN-IU-2016-832 • 1 juv.; same collection data as for preceding; MNHN-IU-2016-553 •
preceding; MNHN-IU-2016-1316 depth 0–15 m; 11 Sep.
2016; st. AS066; MNHN-IU-2016-
MNHN-IU-2016-1042 • same collection data as for preceding; MNHN-IU-2019-3400
8 Oct. 2016; st. AS572; MNHN-IU-depth
37–40 m; 1 Oct. 2016; st. AD283; MNHN-IU-
depth 1–8 m; 25 Sep. 2016; st. AR455; MNHN-IU-2019-3399.
Remarks
A. krappi sp. nov., even sharing
with this species a taller ocular tubercle than in A. dirbergi
slightly from the holotype of A. krappi by having a more acute ocular tubercle tip and more conspicuous
lateral sense organs. This distinction from A. krappi is supported by a high molecular barcode divergence
(p distance = 0.129–0.131). Several adults that could not be barcoded (listed above) present a similar
shape of the ocular tubercle, and they are likely to belong to the same species as the juvenile, MNHN-
IU-2016-834. Their ocular tubercle is taller, but this is coherent with the fact that the barcoded specimen
SABROUX R. et al., Pycnogonida from the Madibenthos Expedition to Martinique
21
Fig. 4. Ammothella krappi sp. nov. A–B, D–I, L–M C. Juvenile,
MNHN-IU-2016-834. J–K A. Body, dorsal view. B. Body, lateral view
(same scale as A). C. Ocular tubercle, frontal view. D. Ocular tubercle, frontal view. E. Fourth leg.
F. Tarsus and propodus of fourth leg. G. Palp. H. Oviger. I. Oviger strigilis. J. Oviger. K. Oviger
strigilis. L. Chelifore. M. Close-up of chela bud.
European Journal of Taxonomy 851: 1–141 (2022)
22
is a juvenile. However, all these specimens are extremely similar to the holotype of A. krappi, except
regarding the shape of the distal region of the ocular tubercle. While we regard this group as likely to
represent a new species, the similarities between A. krappi and the adults listed here as A. krappi
These specimens were sampled on both the Atlantic and Caribbean coasts.
Distribution
Only recorded in Martinique at the moment.
Depth range
2–72 m.
Ammothella marcusi Hedgpeth, 1948
Ammothella marcusi Hedgpeth, 1948: 247.
Ammothella marcusi – Hedgpeth1979:
Sabroux et al. 2019León-Espinosa et al. 2021: 160, tab. 1.
Ramírez-Tello et al. 2022: tab. 1.
Type material
Holotype: USNM 81099 (not examined). Type locality: Logger-head Key, Tortugas, southwest Florida.
Material examined
AB350; MNHN-IU-2016-1302.
Remarks
Only one specimen was collected, from which we did not succeed in amplifying the CO1 barcode
sequence due to the poor quality of the DNA extract (probably due to the advanced state of decay of
the specimen, which hardly consists of more than an empty cuticle, indicating that it was probably dead
upon sampling). This species is generally rare in samplings, though Child (1979) examined a rather
rich collection at the Smithsonian Tropical Reseach Institute in Panama. The species was recorded in
Martinique by Müller (1990a) in Baie de Tartane, so both records of this species in Martinique are from
Distribution
Depth range
0–5 m. We extend this depth range to 15 m.
Ammothella spinifera Cole, 1904
Ammothella spinifera
SABROUX R. et al., Pycnogonida from the Madibenthos Expedition to Martinique
23
Ammothella spinifera
1979: 9; 1992a: 2, 9–11; 2004: 147
––et al.
10–Müller & Krapp 2009: 21 (key), 33–38, 132–
et al. 2019: 3 (list), 6–Sabroux et al. 2019b:
León-Espinosa et al. 2021–Ramírez-Tello
et al. 2022: 162, tab. 1.
Ammothea spinifera – Hall 1913: 132 (key).
Type material
Holotype: USNM 231856 (not examined). Type locality: San Diego, California.
Material examined
MARTINIQUE • depth 3 m; 15 Sep. 2016; st. AB126;
depth 2 m; 12 Sep.
2016; st. AD230; MNHN-IU-2016-827
IU-2016-1141 depth 15 m; 10 Sep. 2016; st. AB062;
MNHN-IU-2016- depth 2 m; 12 Sep. 2016; st.
depth 0–2 m; 17 Sep. 2016;
st. AM021; MNHN-IU-2016-581 • 1 juv.; same collection data as for preceding; MNHN-IU-2016-582 •
4–10 m; 14 Sep. 2016; st. AB123; MNHN-IU-2016-1295.
Remarks
at Vauclin, Petite Anse Macabou and Baie de Tartane, and was abundantly collected during Madibenthos
from both the Atlantic and Caribbean coasts.
Distribution
Depth range
0–28 m.
European Journal of Taxonomy 851: 1–141 (2022)
24
Genus Nymphopsis Haswell, 1884
Type species
Nymphopsis armatus Haswell, 1884, by original designation.
Nymphopsis duodorsospinosa Hilton, 1942
Nymphopsis duodorsospinosa Hilton, 1942a: 303–305, pl. 45.
Nymphopsis duodorsospinosa –
et al. 1976: 134, 157.
: 1, 21; 1992b: 8 (key), 30–2009:
––et al. 2010: 447,
Sabroux et al. 2017: 14,
2019León-Espinosa et al. 2021: tab. 1.
Nymphopsis duodorsospinosum – Child 1982a: 363.
Type material
Holotype: USNM 170494 (not examined). Type locality: San Francisquito Bay, South California.
Material examined
MNHN-IU-2016-812/MK411134.
Remarks
This species is regularly sampled in the Tropical West Atlantic, but is never abundantly collected. This
N. duodorsospinosa for Martinique, on the Caribbean coast.
Distribution
Sea, Gulf of Mexico).
Depth range
0–72 m.
Genus Tanystylum Miers, 1879
Type species
Nymphon styligerum Miers, 1875, by monotypy.
Tanystylum acuminatum Stock, 1954
Tanystylum acuminatum
Tanystylum acuminatum – Stock
Müller & Krapp 2009: 11, 13, 39 (key), 39–41, 128 (ecology), 130 , 132 (phenology), 137
SABROUX R. et al., Pycnogonida from the Madibenthos Expedition to Martinique
25
et al. 2019: 478 (list),
et al.
Type material
Holotype: NL ZMA.PYC.P.1486 (not examined). Type locality: Gustavia, Saint Barthelemy (East
Caribbean).
Material examined
AB189; MNHN-IU-2016-856/MK411169.
Remarks
Lucena et al. (2019) described a velvety region of the femur posteriorly to the cementary tube in this
species, a character also observed in the single collected specimen from Martinique. The species was
from the Caribbean coast of Martinique.
Distribution
Caribbean and Brazil.
Depth range
0–15 m. We extend this depth range to 16 m.
Tanystylum birkelandi Child, 1979
Tanystylum birkelandi
Tanystylum birkelandi – Child
et al.
Type material
Holotype: USNM 170494 (not examined). Type locality: Galeta Island reef, Caribbean Panama.
Material examined
MARTINIQUE • depth 10 m; 19 Sep. 2016; st. AB195; MNHN-
IU-2016-1407.
Remarks
This species has rarely been recorded, mostly from Panama or Belize. The species was previously
recorded from West Indies from Martinique by Müller (1990a) in Petite Anse Macabou. Therefore, the
few records of Tanystylum birkelandi are all from the Atlantic coast. This species shows characteristic
setae with microstetae on coxae 1 (Child 1979).
Distribution
Caribbean (Panama, Belize, Martinique).
Depth range
0–15 m.
European Journal of Taxonomy 851: 1–141 (2022)
26
Tanystylum boucheti sp. nov.
urn:lsid:zoobank.org:act:E1915AAA-F3A8-496C-AF45-3BD9355DEB5E
Fig. 5A–H
nec Tanystylum hummelincki Stock, 1954 – Sabroux et al. 2019b
Material examined
Holotype
MNHN-IU-2016-1074/MK411045.
Etymology
Latinized name, 2nd decl. (masc.), genitive singular. The species is named after Philippe Bouchet, head
of the Madibenthos Expedition.
Description
Trunk completely unsegmented, discoidal; cuticle granular. No dorsomedian ornamentation.
Ocular tubercle lower than abdomen, about as tall as base width, with two acute terminal tips, one large
anteriorly and one smaller posteriorly, and 2 small lateral sense organs laterally, and carrying four large
pigmented eyes. Cephalon with two low lateral tubercles at anterolateral margin. Low dorsal rising at
base of abdomen. Lateral processes about as long as wide, jointed; 1st lateral process with one submedian
tubercle, 2nd and 3rd lateral processes with two dorsolateral tubercles on distal margin, 4th lateral process
with one anterior tubercle on distal margin. Lateral process tubercles carrying one short seta.
Conical in dorsal view, rounded proximally and conical distally in lateral view, with blunt
tip, about as long as two anterior trunk segments.
Vertically oriented, higher than ocular tubercle, not reaching beyond lateral processes. Setae
present on distal part. No basal segmentation.
1-articled, composed of one elongated knob with two distal setae.
6-articled, with setae on all articles except 1st article. 1st article shortest, wider than long. 2nd article
about 2.5 times as long as wide. 3rd article shorter. 4th article longest, about 2.5 times as long as 2nd
article. 5th article about as long as wide, with ventral setae. 6th article about 4 times as long as wide.
10-articled with scarce setae. 1st article as long as wide. 2nd article about twice as long as wide.
3rd article about 1.5 times as long as wide. 4th article about twice as long as 3rd. 5th longest, curved.
6thth article about as long as wide, presenting one distal low spur carrying
long setae. 8th and 9th articles subequal. 10th article shortest. Strigilis spines simple on articles 7th to 9th,
compound on 10th article. Strigilis formula 2:1:1:2.
Stout and nodulous, with scarce small setae. Coxa 1 shorter than wide, carrying 3 distal tubercles:
tip. Coxa 2 longer than coxa 1 or 3. Coxa 3 as long as broad, with ventral setae. Femur stout, about twice
as long as wide, with one elevated, dorsdistal cement gland tube. Tibia 1 and tibia 2 with large rounded
tubercles and long setae on dorsal surface. Femur, tibia 1 and tibia 2 subequal. Tarsus trapezoid, short,
about as long as wide, carrying one large spine and setae on ventral surface. Propodus stout and curved,
propodus. Auxiliary claws present, about half as long as main claw.
SABROUX R. et al., Pycnogonida from the Madibenthos Expedition to Martinique
27
Fig. 5. Comparison between Tanystylum boucheti sp. nov. (A–H) and T. hummelincki Stock, 1954 (I–
J). A–H I–J A. Body, dorsal view.
B. Body, lateral view (same scale as A). C. Palp. D. Third leg. E. Cement gland tube. F. Tarsus and
propodus of third leg. G. Oviger. H. Oviger strigilis. I. Body, dorsal view. J. Body, lateral view. I and J
same scale as A.
European Journal of Taxonomy 851: 1–141 (2022)
28
(mm). Trunk 0.61; abdomen 0.20; proboscis 0.33; chelifore 0.12; coxa 1 0.12; coxa 2
0.26; coxa 3 0.10; femur 0.42; tibia 1 0.40; tibia 2 0.45; tarsus 0.05; propodus 0.35; main claw 0.21.
Sexual dimorphism
Female unknown.
Remarks
The closest species to Tanystylum boucheti sp. nov. is T. isthmiacum Stock, 1955; we distinguish this
margin of coxae 1 rather than rounded, and its more pronounced dorsal tubercles on the margin of
the lateral processes (Müller & Krapp 2009). It may also be worth noting that the original description
of T. isthmiacum from Panama mentioned 5-articled palps, while many specimens (often regarded as
belonging to the subspecies Stock, 1966) are generally recorded with 6-articled
palps in the Atlantic (e.g., Fage & Stock 1966; Stock 1966a, 1979; Müller & Krapp 2009). This may
be a whole distinct species, morphologically closer to T. boucheti
criteria mentioned above). It is therefore possible that T. bouchetiT. isthmiacum or
in the past.
Another similar species is Tanystylum hummelincki Stock, 1954 (Fig. 5I–J), with which the present
material was mixed in Sabroux et al. (2019b). Indeed, the two species show similar appendages and
T. boucheti
T. hummelincki by the length of the proboscis, the height and shape of the ocular
tubercle, and by the higher abdomen, which reaches beyond the ocular tubercle. The CO1 p-distances
between the T. boucheti T. hummelincki range from
0.104 to 0.106 (see Appendix).
The species also strongly resembles Tanystylum evelinae Marcus, 1940, described and only recorded
from Saõ Paulo State, Brazil (Marcus 1940). Tanystylum bouchetiT. evelinae by the
presence of a posterior tubercle on the ocular tubercle, by the shape of the proboscis, which is conical
rather than barrel-shaped, by the orientation of the abdomen, and by the size of the cement gland tube,
which is longer in T. boucheti.
Tanystylum relatives using the following combination
tubercles on the lateral processes, the conical shape of the proboscis, and the shape of the ocular tubercle.
A single specimen of the species was collected on the Atlantic coast of Martinique.
Distribution
Only known from the type locality.
Depth range
0–2 m.
Tanystylum duospinum Hilton, 1939
Tanystylum duospinum Hilton, 1939: 33.
Tanystylum oculospinosum Hilton, 1942b: 70.
Tanystylum tubirostre
SABROUX R. et al., Pycnogonida from the Madibenthos Expedition to Martinique
29
Tanystylum mexicanum –h.
Tanystylum oculospinum — Stock 1954a: 122; 1955: 248–249; 1966b: 390.
Tanystylum oculospinosum – Child & Hedgpeth 1971Child 1979: 34; 1992a: 2–3, 23–24;
1992b2009Stock 1994: 18, 38.
Tanystylum tubirostrum – Stock 1975a: 984.
Tanystylum tubirostrum – Stock
Tanystylum duospinum –
Sabroux
et al. 2019Wang et al. 2020León-Espinosa et al. 2021
Ramírez-Tello et al. 2022: 154, tab. 1.
Type material
Tanystylum duospinum Hilton, 1939. Holotype: USNM 81531 (not examined). Type locality: Baja
Tanystylum oculospinum Hilton, 1942. Holotype: USNM 81518 (not examined). Type locality: Todos
Tanystylum tubirostre Stock, 1954. Holotype: NL ZMA.PYC.P.1520 (not examined). Type locality: Punt
Vierkant, Bonaire.
Tanystylum mexicanum Child, 1979. USNM 170650 (not examined). Type locality: Gulf of California,
Material examined
MARTINIQUE • depth 10 m; 19 Sep. 2016; st. AB195;
depth 23 m; 25 Sep. 2016;
st. AB405; MNHN-IU-2016- depth
0–16 m; 18 Sep. 2016; st. AS255; MNHN-IU-2016-1177/MK411071.
Remarks
We follow the suggestion of Bamber et al. (2022) in considering T. oculospinosum, T. tubirostrum and
T. mexicanum as junior synonyms of T. duospinum
Distribution
Panama, Galápagos, Ecuador, Peru), and West Atlantic (Bermuda, Gulf of Mexico, Caribbean). Other
records from Kenya, Taiwan, Indonesia and Australia (Carpentaria Gulf). It is worth noting that the
distribution range nominatively for T. duospinum
that the widest distribution for the species has been recorded under the synonym T. oculospinosum
. Tanystylum tubirostrum
T. mexicanum was
Depth range
Intertidal to 54 m.
European Journal of Taxonomy 851: 1–141 (2022)
30
Tanystylum hummelincki Stock, 1954
Fig. 5I–J
Tanystylum hummelincki
Tanystylum hummelincki – Stock–21,
et al. 2019b
Type material
Holotype: NL ZMA.PYC.P.1485 (not examined). Type locality: La Pecha, Los Frailes Islands, Venezuela.
Material examined
collection data as for preceding; MNHN-IU-2016-1398/MK411111 • 1 juv.; same collection data as
12 Sep. 2016; st. AM012; MNHN-IU-2016-1309.
Remarks
This is another rare species of Tanystylum
abundantly sampled here in this genus. Most specimens were collected on the Atlantic coast of
Martinique, except MNHN-IU-2016-1309, which was sampled near the southern cape of the Island.
Distribution
Caribbean (Venezuela, Colombia, Martinique).
Depth range
0–3 m. We extend the known range to 23 m.
Tanystylum ingrallis sp. nov.
urn:lsid:zoobank.org:act:2CBAF7F7-5414-4617-80C9-809155ADA7FE
Fig. 6
Tanystylum sp. 3 – Sabroux et al.
Material examined
Holotype
MARTINIQUE •
MNHN-IU-2016-1055/MK411038.
Paratype
MARTINIQUE • depth 80 m; 24 Sep. 2016; st. AD261; MNHN-
IU-2016-867/MK411178.
Etymology
Contraction of ‘in grallis’, in + plural accusative of the Latin gralla, -ae (1st decl, fem.): ‘on stilts’,
referring to the long legs of this species, relatively to most instars of the genus.
SABROUX R. et al., Pycnogonida from the Madibenthos Expedition to Martinique
31
Description
Trunk completely unsegmented, discoidal; cuticle granular. No dorsomedian ornamentation.
eyes, one acute terminal spur anteriorly and 2 small lateral sense organs. Cephalon with 2 small lateral
tubercles at anterior margin. 1st
as long as wide; all lateral processes jointed; 1st lateral process without ornamentation, 2nd, 3rd and 4th
lateral processes with one anterior tubercle on distal margin, carrying one seta.
Proximal half broad and rounded, distal half tubular, blunt at tip, bent ventrally, about as
long as distance between cephalon distal margin and abdomen base.
th
One dorsal tubercle in distal part, carrying setae.
1-articled, composed of one small knob with one distal spine.
6-articled, with setae mostly on last 2 distal articles. 1st article shortest, wider than length. 2nd
article about 2.5 times as long as wide. 3rd article as long as 1st article. 4th article longest, about 1.7 times
as long as 2nd article, about 3.5 times as long as wide, with cannula (gland?) on outer side. 5th article
about 0.2 times as long as wide. 6th article about 3.5 times as long as wide.
10-articled, with scarce setae. 1st article as long as wide. 2nd article more than 1.5 times as long
as wide. 3rd article about twice as long as wide. 4th and 5th articles subequal and longest, 4th about 3.5
times as long as wide, 5th about 4 times as long as wide. 6th article about twice as long as wide. 7th article
about as long as wide, presenting one distal low spur carrying long setae. 8th article about as long as
wide. 9th article about twice as long as wide. 10th article about 1.5 times as long as wide. Strigilis spines
Rather long for the genus, 1st leg pair longer than others. Coxa 1 shorter than wide, with 4 tubercles
mounted with seta on distal margin of 1st and 2nd legs: one dorsomedian, two dorsolateral and one
ventrolateral on posterior side; or with 3 tubercles on 3rd and 4th legs: one dorsomedian, one dorsolateral
and one ventrolateral on posterior side. Coxa 2 longer than coxa 1 or 3, with one large distal tubercle
on anterior side and 2 or 3 distal tubercles on posterior side, all mounted with one seta at tip. Coxa 3 as
long as broad, with ventral setae. Femur straight, with long dorsal spur at distal margin with one lateral
cement gland tube on inner side, rounded at base and tubular distally. Tibia 1 shorter than femur, with
two low dorsal knobs mounted by long setae and distal dorsomedian spur. Tibia 2 longer than tibia 1 or
femur, with scarce setae, and 3 low knobs on dorsal surface mounted with long setae. Tarsus trapezoid,
short, about as long as wide, carrying one large spine and setae on ventral surface. Propodus gently
curved, with low heel carrying 2 large heel spines, plus one smaller spine more distally. Main claw
curved, about half as long as propodus. Auxiliary claws present, about 0.6 times as long as main claw.
(mm). Trunk 0.84; abdomen 0.55; proboscis 0.80; coxa 1 0.22 (1st leg), 0.24 (2nd leg); coxa
2 0.46 (1st leg), 0.36 (2nd leg); coxa 3 0.27 (1st leg), 0.28 (2nd leg); femur 1.25 (1st leg), 0.82 (2nd leg); tibia 1
1.17 (1st leg), 0.78 (2nd leg); tibia 2 1.40 (1st leg), 1.04 (2nd leg); tarsus 0.10; propodus 0.50; main claw
0.28.
Sexual dimorphism
No female currently available.
European Journal of Taxonomy 851: 1–141 (2022)
32
Fig. 6. Tanystylum ingrallis A. Body, dorsal view.
B. Body, lateral view (same scale as A). C. Palp. D. Oviger. E. Oviger strigilis. F. Second leg. G. Tarsus
and propodus of second leg. H. Femur of third leg, dorsal view (same scale as F). I. Close-up of distal
spur on femur of second leg, and cement gland tube. J. First leg (same scale as F).
SABROUX R. et al., Pycnogonida from the Madibenthos Expedition to Martinique
33
Individual variability
On the other available specimen, the chelifores are almost not visible in dorsal view.
Remarks
Tanystylum ingrallis sp. nov. is readily discriminated from most other described species by the peculiarly
long and slender legs and the absence of nodules on legs. Most species of Tanystylym have their propodus
conspicuously longer than half the tibia 2, while here the propodus is just half as long as tibia 2 for the
T. lamonti Staples, 2019, T. tiara
Staples, 2019, T. zuydtorpi Arango, 2009 and T. paramexicanum Müller & Krapp, 2009 have long legs
as well. Tanystylum lamonti, T. tiara and T. zuydtorpi T. ingrallis, due for
example to the shape of their proboscis (long and barrel shape in the two former species, pyriform in
the latter), and their longer chelifores. On the other hand, T. paramexicanum shares some morphological
characters with this new species: the shape of the proboscis, the strong spur on the anterior margin of
the ocular tubercle and the presence of 3 or 4 tubercles on coxae 1. However, the two species do not
the shorter size of coxae 2 and the bent ocular tubercle tip in T. paramexicanum, but also by the longer
abdomen, the presence of tubercles on coxae 2 and the presence of a high tubercle on the distal margin of
tibia 1 in T. ingrallisT. paramexicanum.
Regrettably, only females are known for T. paramexicanum, so it is not possible to compare the cement
gland opening of the two species. Indeed, the lateral opening of the cement gland tube in T. ingrallis is
another strong diagnostic character.
The morphologically closest species to T. ingrallis sp. nov. is probably T. duospinum Hilton, 1939,
which presents a similar proboscis, and has the same palp structure. These two species are, however,
T. ingrallis extends beyond
T. duospinum; (ii) in T. ingrallis, the abdomen is oriented diagonally, while it is almost vertical in
T. duospinum; (iii) the anterior tubercle-ornamented coxa 1 is bifurcated in T. duospinum
(Müller & Krapp 2009), but not in T. ingrallis; (iv) the dorsodistal spur of the legs is longer in T. ingrallis
compared to T. duospinum; (v) knobs dorsally positioned along the legs are smaller and more spaced in
T. ingrallis compared to T. duospinum; (vi) tibia 2 is longer relative to tibia 1 in T. ingrallis compared
to T. duospinum. Two specimens of the species were collected on the Caribbean and northern coasts of
Martinique.
Distribution
Only known from Martinique.
Depth range
40–80 m.
Tanystylum orbiculare Wilson, 1878
Tanystylum orbiculare
Tanystylum orbiculare
Sumner et al. 1913
29–
1975a: 985; 1986: tab. 1; Clark 1963 (?):
European Journal of Taxonomy 851: 1–141 (2022)
34
1987: 43–Krapp & Kraeuter 1976Munilla & de Haro 1981
11; 1984: 531, 533–535, tabs 3–Bremec et al.Child 1992b: 8 (key),
215, tabs 2–Gillespie & Bain 2006: 309, 316–et al. 2006: tabs
Arango 2007: 917, 919–920. Krapp et al.
Masta et al. 2010 (mitogenome): 61–62,
et al. 2011a: 321; 2011b: 345, 347; et al.
–
2–et al.
Sabroux et al. 2017: 15, appendices 1–Dietz et al. 2018
et al. 2019Ramírez-Tello et al. 2022: tab. 1.
Clotenia orbiculare – Bouvier 1923aGiltay 1929: 175.
non Tanystylum orbiculare – Stock 1954b: 145 (= Tanystylum conirostre (Dohrn, 1881))
nec Tanystylum conirostre (Dohrn, 1881) – Sabroux et al. 2019
Type material
Holotype: USNM 37781 (not examined, illustrated in Child 1992b). Type locality: Long Island Sound,
Material examined
Remarks
Tanystylum conirostre in Sabroux et al. (2019b), as the examined
T. orbiculare given by Krapp (1973) by having slenderer legs
and a straight abdomen. However, T. orbiculare can be distinguished from T. conirostre by the number
of palp articles (5 rather than 4) and the proboscis shape, which is more barrel-shaped in T. orbiculare.
Although this species is very common in the Atlantic, it was not yet recorded from Martinique.
CO1 p-distances to previously barcoded material (Arango & Wheeler 2007; Masta et al. 2010;
respectively GenBank accession numbers DQ390064 from Mar del Plata, Argentina, and GU370064
from Massachusetts) are high (0.172 and 0.11–0.14, respectively; see Appendix). All specimens were
collected at a single station in the Baie de Fort-de-France.
It should be noted that this species is mostly amphi-Atlantic, but two records were provided from New
South Wales by Stock (1954b) and Clark (1963). However, Stock regarded T. orbiculare as synonymous
to T. conirostre and explicitly stated that he found the material similar to the type material of Clotenia
conirostris ( = T. conirostre), so this record should be regarded as T. conirostre. In turn, it is possible that
T. orbiculare following Stock.
Distribution
Widely recorded in the Atlantic (New England, Massachusetts, Virginia, South Carolina, Georgia,
Brazil, Argentina, Senegal, Congo, Angola, Cap Verde); also found in the Caribbean, Gulf of Mexico,
SABROUX R. et al., Pycnogonida from the Madibenthos Expedition to Martinique
35
occidental and oriental Mediterranean Sea, as well as the Bosphorus. It was also recorded in New
South Wales (Clark 1963), though this record is questionable considering the otherwise amphi-Atlantic
distribution of this species.
Depth range
0.5–58 m.
Tanystylum tayronae Müller & Krapp, 2009
Tanystylum tayronae
Tanystylum tayronae – Sabroux et al.
Tanystylum sp. – Müller 1990a: 280.
Type material
Holotype: SMF1111 (not examined). Type locality: Arreçifes near Punta el Diamante, Santa Marta,
Colombia.
Material examined
MARTINIQUE • depth 2 m; 12 Sep. 2016; st. AD230;
depth
depth 15 m; 10 Sep. 2016; st. AB062; MNHN-IU-2016-1300/MK411097.
Remarks
as Tanystylum sp., and was formally described later by Müller & Krapp (2009) from Colombia. The
characteristic tubercle near the abdomen base is quite variable in size, but it is generally lower in the
studied material compared to Müller & Krapp’s drawing. This tubercle is the only conspicuous character
T. tayronae from T. geminum
DNA barcodes are needed to make sure the two species are not synonymous.
All specimens were collected on the Atlantic and Caribbean coasts of Martinique.
Distribution
So far only known from the Caribbean (Colombia and Martinique).
Depth range
0–2 m, extended to 15 m.
Family Pallenopsidae Fry, 1978
Genus Pallenopsis Wilson, 1881
Type species
Pallenopsis (Pallenopsis) (Krøyer, 1844), by subsequent designation (Child 1998a).
European Journal of Taxonomy 851: 1–141 (2022)
36
Pallenopsis candidoi Mello-Leitão, 1949
Pallenopsis candidoi Mello-Leitão, 1949: 299–307, pls. 9–10.
Pallenopsis candidoi –
et al.
Pallenopsis (Pallenopsis) candidoi – Stock 1975a: 1018 (key), 1030; 1992a: 130, 139.
Type material
Type locality:
Florianópolis, Santa Catarina, Brazil.
Material examined
MARTINIQUE • depth 11–14 m; 6 Oct. 2016; st. AS565;
MNHN-IU-2016-814/MK411136.
Remarks
distribution of this species, already recorded from the northwest Atlantic, Surinam and Brazil. The
specimen was collected on the Atlantic coast.
Distribution
West Atlantic (Brazil, Surinam, Caribbean and US Georgia).
Depth range
14–102 m.
Pallenopsis schmitti Hedgpeth, 1943
Pallenopsis schmitti Hedgpeth, 1943: 44.
Pallenopsis schmitti – Hedgpeth
Müller & Krapp 2009: 121 (key), 122, 137,
et al. 2019b
Pallenopsis (Pallenopsis) schmitti – –
Type material
Material examined
MARTINIQUE • depth 74 m; 2 Oct. 2016; st. AD290;
MNHN-IU-2016-813/MK411135.
Remarks
coast. This species was already recorded in the Caribbean from Colombia, Cuba and Florida.
SABROUX R. et al., Pycnogonida from the Madibenthos Expedition to Martinique
37
Distribution
Tropical West Atlantic (Gulf of Mexico, Caribbean, French Guiana, Brazil).
Depth range
15–600 m.
Family Ascorhynchidae Hoek, 1881
Genus Ascorhynchus Sars, 1877
Type species
Ascorhynchus abyssi Sars, 1877, by original designation.
Ascorhynchus castellioides Stock, 1957
Ascorhynchus castellioides
Ascorhynchus castellioides – Bayer et al. 1975a: 968;
Birkeland et al.Arango & Wheeler 2007:
Müller & Krapp 2009: 10, 55 (key),
57–58, 130–et al.
Krapp & Viquez 2011: 205, tab. 1Sabroux et al. 2017
2019Wagner et al. 2017
Lucena et al. 2019
Type material
Holotype: ZMH-A0000719 (not examined). Type locality: Puerto Cabello, Venezuela.
Material examined
depth 2 m; 3 Oct. 2016; st. AB382; MNHN-IU-2016-583.
Remarks
coast).
Distribution
Tropical West Atlantic: Caribbean (Venezuela, Panama, Florida, Columbia, Barbados, Martinique,
Bonaire, Curaçao) and Brazil (Pernambuco and Paraíba).
Depth range
0–30 m.
European Journal of Taxonomy 851: 1–141 (2022)
38
Ascorhynchus horologium Child, 1992
Ascorhynchus horologium
Ascorhynchus horologium – Child 2002: 1814 (key); 2009et al. 2019b: tab. 1.
Type material
Material examined
MARTINIQUE • depth 90 m; 14 Sep. 2016; st. AD231;
MNHN-IU-2016-1199 • 1322.
Remarks
are morphologically very similar to Child’s description. Unfortunately,
negative for the two DNA extracts.
Distribution
Only recorded from West Florida (Gulf of Mexico) and Martinique.
Depth range
73–90 m.
Ascorhynchus iguanarum sp. nov.
urn:lsid:zoobank.org:act:AF5026C5-94E5-4B01-BBC1-684199D91CBE
Fig. 7
Ascorhynchus sp. 5 – Sabroux et al. 2019
Material examined
Holotype
MNHN-IU-2016-1047/MK411035.
Paratype
MNHN-IU-2016-1190/MK411077.
Etymology
From Latin iguana, -ae, 1st decl. (fem.), genitive plural. Named after the green iguanas of Fort Saint-
dorsomedian trunk tubercles of this new species.
Description (holotype, MNHN-IU-2016-1047)
Trunk stout, smooth, completely segmented. Trunk segments 1 to 3 bearing tall dorsomedian
pointing tubercles, slightly curved anteriorly. Ocular tubercle shorter than dorsomedian tubercles, about
SABROUX R. et al., Pycnogonida from the Madibenthos Expedition to Martinique
39
2.5 times as tall as wide, with acute tip, bearing four pigmented eyes distally. Lateral margin of cephalon
processes about as long as wide, 4th pair shortest, well separated by less than half their own diameter,
distally bearing a dorsomedian pointing tubercle, without any seta or spine at tip.
Horizontal, long, extending beyond coxa 2 of 4th
Abdomen segmented at base.
2-articled, shorter than proboscis. Scape 1-articled, carrying seta at mid-length and 3
additional setae at distal margin. Chela not functional, bearing one lateral seta. Finger oriented ventrally.
10-articled, with setae mostly on ventral side of last 5 articles. 1st article shorter than wide, carrying
dorsal pointy tubercle. 2nd article shortest, shorter than width. 3rd article subequal to 5th article and longer
than other articles, with scarce spines. 4th article slightly longer than wide. 5th article with one long lateral
seta on outer surface near proximal end, plus several small spines near distal end. 6th article bent, as long
as wide, with setae on ventral surface. 7th and 8th articles subequal, about 1.5 times as long as wide. 9th
and 10th articles subequal, about as long as wide.
10-articled, with scarce setae. 1st article shortest, as long as wide. 2nd article about twice as long
as wide. 3rd article 1.5 times as long as wide, glabrous. 4th and 5th articles subequal in length, 5th article
curved. 6th and 7th articles subequal. 6th article twice as long as wide. 7th article longest within strigilis,
8th, 9th and 10th articles subequal. Strigilis with 2 rows of compound spines. Terminal claw short, with
blunt tip.
Slender with scarce setae. Coxa 1 shorter than wide, dorsodistally with 2 tubercles, each
surmounted with spine. Coxa 2 slightly longer than maximal width, with ventral seta. Coxa 3 as long as
1 or femur, with many long setae on dorsal surface and shorter setae on ventral surface. Tarsus medium-
sized, trapezoid. Propodus straight, about 0.6 times as long as tibia 2, sole with 8 small spines. Main
claw curved, about 0.4 times as long as propodus. No auxiliary claw.
(mm). Trunk 1.13; abdomen 0.39; proboscis 0.73; chelifore 0.21; coxa 1 0.14; coxa 2
0.29; coxa 3 0.18; femur 0.57; tibia 1 0.57; tibia 2 0.71; tarsus 0.09; propodus 0.38; main claw 0.15.
Sexual dimorphism
No female currently available.
Individual variability
The paratype is a juvenile, which presents developed chelae as expected, and is overall smaller in size.
Remarks
The high, curved, conical dorsomedian tubercles of this species is a strong diagnostic character for this
species. Only Ascorhynchus athernus Child, 1982 shows dorsomedian tubercles comparable in size
and longer than the ocular tubercle, but it is otherwise very dissimilar to A. iguanarum sp. nov. This
species is morphologically close to A. castelli (Dohrn, 1881), which can be found in the same region,
from which it can be discriminated not only by the higher dorsomedian tubercles, but also by the larger
European Journal of Taxonomy 851: 1–141 (2022)
40
Fig. 7. Ascorhynchus iguanarum sp. nov., h A. Body, dorsal view.
B. Body, lateral view (same scale as A). C. Palp. D. Oviger, with close-up of strigilis spines. E. Third leg.
F. Tarsus and propodus of third leg.
SABROUX R. et al., Pycnogonida from the Madibenthos Expedition to Martinique
41
tubercles on the lateral processes, the presence of small pointy tubercles on the 1st article of the palp and
at the base of palp insertion, the absence of a spur on the dorsodistal margin of tibiae 1 and the smaller
distance between the lateral processes.
The two specimens were collected on the Atlantic coast of Martinique.
Distribution
Only known from Martinique.
Depth range
14–23 m.
Ascorhynchus latipes (Cole, 1906)
Barana latipes Cole, 1906: 217–222, pls 1–2.
Ascorhynchus latipes
et al.2009:
et al. 2019bet al. 2021: tab. 1. Ramírez-Tello
et al. 2022: tab. 1.
Ascorhynchus cf. latipes – Krapp & Viquez 2011: 204, tab. 1.
Type material
Holotype: MCZ CRU-6947 (not examined). Type locality: Sweeting’s Village, Great Abaco, Bahamas.
Material examined
depth 4–9 m; 27 Sep. 2016; st. AR461; MNHN-IU-2016-871/MK411182.
Remarks
southern coasts, although one specimen was found in Pointe Baham (southern Atlantic coast).
Distribution
Tropical, amphi-Atlantic (Gulf of Mexico, Caribbean, Senegal).
Depth range
0–19 m.
Ascorhynchus sp. 4
Fig. 8
Ascorhynchus sp. 4 – Sabroux et al. 2019
Material examined
MARTINIQUE • depth 78–80 m; 24 Sep. 2016; st. AD263;
MNHN-IU-2016-1039/MK411031.
European Journal of Taxonomy 851: 1–141 (2022)
42
Remarks
The only available specimen is a juvenile that is rather poorly preserved. The ovigers are broken. In
these circumstances, it seems unwise to name this species, although it is probably new to science.
Ascorhynchus by the following character combination:
ocular tubercle present, with pigmented eyes positioned below the ocular tubercle tip; slender
dorsomedian tubercles on trunk segments; lateral processes less than twice as long as wide, separated
by about their own diameter, carrying median tubercle; ocular tubercle near anterior cephalon boarder;
chelifore scapes monosegmented. These characters are shared with A. okai Nakamura & Child, 1983
from Japan, A. cactoides A. pudicus Stock, 1970
from the Eastern Atlantic and Mediterranean (Munilla and Soler-Membrives 2014). Ascorhynchus sp. 4
A. okai by the longer distance between the insertions of the ovigers and palps (Nakamura &
Child 1983), from A. cactoides in having the abdomen straight and from A. pudicus in having no
dorsomedian tubercle on the 4th trunk segment.
The specimen was collected in Northern Martinique.
Depth range
78–80 m.
Genus Eurycyde Schiødte, 1857
Type species
Zetes hispidus Krøyer, 1844 by monotypy.
Eurycyde clitellaria Stock, 1955
Eurycyde clitellaria
Eurycyde clitellaria –
Stock
et al.
2019bRamírez-Tello et al. 2022: tab. 1.
Type material
Holotype: NHMD 110654 (not examined). Type locality: sound between St John and St James (Virgin
Islands).
Material examined
MARTINIQUE • depth 70 m; 9 Sep. 2016; st. AD214;
depth 19 m; 7 Sep.
depth 65 m; 11 Sep. 2016; st. AD224; MNHN-IU-2016-825/MK411144 same collection
data as for preceding; MNHN-IU-2016-579
depth
depth 65 m; 11 Sep. 2016; st. AD222; MNHN-IU-2016-547 •
depth 28 m; 7 Sep. 2016; st. AB157; MNHN-
IU-2016-549 • Vétiver; depth 25 m; 8 Sep. 2016; st. AB161; MNHN-
SABROUX R. et al., Pycnogonida from the Madibenthos Expedition to Martinique
43
Fig. 8. Ascorhynchus sp. 5, juvenile, MNHN-IU-2016-1039. A. Body, dorsal view. B. Body, lateral view
(same scale as A). C. Palp. D. First leg. EF. Chela.
European Journal of Taxonomy 851: 1–141 (2022)
44
depth 72 m; 8 Oct. 2016; st.
depth
37–40 m; 1 Oct. 2016; st. AD283; MNHN-IU-2016-1053 • 5 juvs; same collection data as for preceding;
MNHN-IU-2016-1289.
Remarks
A noticeable feature of this species compared to other representatives of Eurycyde is the strong bending
of the abdomen about mid-length in a right angle. This feature is illustrated by Müller & Krapp (2009)
and Stock (1955), although the bending seems to be only very slight in Child (1992b).
the Caribbean coast. Records of E. clitellaria outside the Caribbean Sea are actually rare, with only one
North Carolina (McCloskey 1967).
Distribution
Depth range
Intertidal to 100 m.
Eurycyde kaiouti sp. nov.
urn:lsid:zoobank.org:act:8B0E31E8-5FB3-47DF-8BED-F63E014DA6B7
Fig. 9A–J
nec Eurycyde raphiaster Loman, 1912 – Sabroux et al.
Material examined
Holotype
MNHN-IU-2016-1129/MK411058.
Paratypes
SABROUX R. et al., Pycnogonida from the Madibenthos Expedition to Martinique
45
st. AB173; MNHN-IU-2019-3392.
Etymology
Apposition of the Kalinago (Caribbean) word ‘kaiouti’, ‘which has spines’, in reference to the long
tubercles of coxae 1.
Description
ornamentation. Ocular tubercle tall, carrying four pigmented eyes and seven thick setae around ocular
tubercle tip. Oviger base almost touching lateral processes. Lateral processes about 1.5 times as long as
wide, 4th pair shortest, well separated by less than half their own diameter, distally bearing a dorsomedially
pointing tubercle, without seta or spine at tip.
Bipartite, anterior part as petiole, posterior part pyriform, about as long as body, anterior and
posterior parts articulated together.
European Journal of Taxonomy 851: 1–141 (2022)
46
Long, extending beyond coxa 1 of 4th
articled with trunk, movable, tip conical.
3-articled, reaching beyond proximal petiole of proboscis. Scape 2-articled, 1st article
longest, with distal dorsomedian thick seta. 2nd article carrying many thick setae dorsally. Chela residual,
st article about as long as wide,
carrying one dorsal pointy tubercle. 2nd article shortest, shorter than wide. 3rd article longest, with distal
thick setae. 4th article slightly longer than wide. 5th article 0.7 times as long as 3nd, carrying several thick
setae. 6th article about twice as long as wide, 6th and 10th articles subequal in length, shorter than 7th. 7th,
8th and 9tharticles subequal in length.
10-articled, with scarce setae. 1st article shortest, as short as wide. 2nd and 3rd articles subequal.
4th and 5th articles subequal in length, 5th article curved. 6th articles 3 times as long as broad, longer
than 7th article. 7th article longest within strigilis. 8th, 9th and 10th articles subequal. Compound spines of
strigilis in 2 rows. Terminal claw shorter than 10th article.
Slender with many thick setae. Coxa 1 shortest, about as long as wide, with 2 long dorsolateral
pointy tubercles on distal margin, anterior one about as long as 1st coxa, posterior one at least twice as
long as anterior one and carrying long thick seta at base for 1st to 3rd legs. Coxa 2 slightly longer than
broad. Coxa 3 as long as wide. Femur with cement gland tube at about mid-length on anterior side and
many thick setae distally. Tibia 1 13 times as long as wide, about 1.3 times as long as femur, thinner,
with many dorsal thick setae at about mid-length. Tibia 2 subequal to tibia 1. Tarsus medium-sized,
trapezoid. Propodus gently curved, about half as long as tibia 2; sole carrying many small spines. Main
(mm). Trunk 1.05; abdomen 0.55; proboscis 1.42; chelifore 1.05; coxa 1 0.23; coxa 2
0.27; coxa 3 0.16; femur 0.72; tibia 1 0.92; tibia 2 0.90; tarsus 0.10; propodus 0.48; main claw 0.16.
Sexual dimorphism
Females with shorter oviger articles; anterior dorsolateral pointy tubercle of coxa 1 often shorter than
in males.
Individual variability
Size of pointy dorsolateral tubercles of coxa 1 variable among specimens, posterior one always
longer (or rarely as long) than length of coxa 1, anterior one always shorter, posterior tubercle always
conspicuously longer than anterior tubercle. In holotype specimen, posterior dorsolateral pointy tubercle
of coxa 1 curved anteriorly, but may curve posteriorly in other specimens.
Remarks
Eurycyde kaiouti sp. nov. can be discriminated from most other species of Eurycyde by the following
combination of characters: (i) the ocular tubercle presents long thick setae (the number of these setae
may vary with loss and we do not recommend using it as a criterion); (ii) the coxa 1 carries dorsaly a
(iii) lateral processes bear one pointy tubercle dorsally. These characters are only shared with
E. raphiaster Loman, 1912, to which it is very close morphologically. Comparing with the illustration
in Bouvier (1917) of the type material, the two tubercles of coxae 1 in E. raphiaster are subequal in
length and less long than coxa 1 (shorter in females). Instead, E. kaiouti presents a posterior coxa 1
tubercle that is much longer than the anterior one: the anterior tubercle is shorter than coxa 1, the
SABROUX R. et al., Pycnogonida from the Madibenthos Expedition to Martinique
47
Fig. 9. Comparison between Eurycyde kaiouti sp. nov. (A–J) and E. raphiaster Loman, 1912 (K).
A–JKA. Body, dorsal view. B. Body,
lateral view (same scale as A). C. Oviger. D. Oviger strigilis. E. Pointed teeth of oviger strigilis. F. Third
leg. G. Tarsus and propodus of third leg. H. Chelifore. I. Chela bud. J. Palp. K. Body, dorsal view (same
scale as A).
European Journal of Taxonomy 851: 1–141 (2022)
48
E. raphiaster and E. kaiouti is recovered
also in juveniles. Furthermore, we observe that, in Martinique material, the tibia 1 in E. kaiouti is
shorter than in E. raphiaster, e.g., 13 times as long as wide for the E. kaiouti holotype against 15 times
for E. raphiaster specimen MNHN-IU-2016-818. The two species can be distinguished based on CO1
barcodes (E. kaiouti E. raphiaster =
0.173–0.187; see Appendix).
One specimen recorded in Sabroux et al. (2019b), MNHN-IU-2016-819, is morphologically very
similar to the holotype of Eurycyde kaiouti sp. nov., but falls in a barcode cluster on its own that is very
divergent from the E. kaiouti cluster (p-distance = 0.182–0.188, see Appendix). Because it is the only
discriminate it as another species relying on morphological characters. For this reason, we choose to
keep this specimen as Eurycyde sp. Further material from NWTA Eurycyde should be barcoded and
studied to resolve the status of this specimen.
Eurycyde kaiouti was widely collected on both Atlantic and Caribbean coasts.
Distribution
Only known from Martinique.
Depth range
1–90 m.
Eurycyde raphiaster Loman, 1912
Fig. 9K
Eurycyde raphiaster Loman, 1912: 13.
Eurycyde raphiaster –
et al.Child 1979: 21; 1982a: 360; 1988a: 8 (key); 1992b: 8
Arango
Nakamura et al.
32–et al.Sabroux et al. 2017 : appendices 1–2,
2019
Type material
Holotype: MOM INV-0001360 (not examined). Type locality: 4 miles southwest of Boa Vista, Cap
Verde.
Material examined
1 juv.; same collection data as for preceding; MNHN-IU-2016-548 • 1 juv.; same collection data as for
SABROUX R. et al., Pycnogonida from the Madibenthos Expedition to Martinique
49
22 m; 9 Sep. 2016; st. AB163; MNHN-IU-2016-546
depth 63 m; 21 Sep. 2016; st. AD257; MNHN-IU-2016-
depth
depth 25 m; 8 Sep. 2016; st. AB161; MNHN-IU-2016-
MNHN-IU-2016-1305.
Remarks
Unlike Müller & Krapp (2009), we have never observed an absence of the lateral process tubercle in
females.
Eurycyde raphiaster and E. kaiouti sp. nov. are very similar species that may have been mistaken in the
past. After revising literature illustrations, we agree with the determinations of Müller & Krapp (2009)
(at least the illustrated male), while Hedgpeth (1948) and Child (1992b) may have illustrated E. kaiouti
rather than E. raphiaster.
that it is less common in Martinique than E. clitellaria or E. kaiouti. The species was most often collected
Caravelle on the Atlantic coast.
Distribution
Bahamas, Gulf of Mexico, Caribbean, Cap Verde, Guyana and Senegal.
Depth range
0–91 m.
Family Callipallenidae Hilton, 1942
Genus Callipallene Flynn, 1929
Type species
Pallene brevirostris Johnston, 1837, by original designation.
Callipallene brevirostris (Johnston, 1837)
Pallene brevirostris
Pallene empusa
Pallene brevirostris
1884a: 649;
–
–Schimkewitsch 1909:
–
et al.
European Journal of Taxonomy 851: 1–141 (2022)
50
144.
Pallene sp. – Verrill 1873: 415.
Pallene empusa – Wilson
Callipallene brevirostris ––
1954a: 115; 1954b: 43––
–622; 1974: 59 (key), 60–
– et al. 1978:
94– –2009: 819 (list).
Arango & Wheeler 2007: appendix
et al. 2007Nakamura et al. 2007Montoya Bravo
et al. 2009Müller & Krapp 2009: 11, 67 (key), 67–69, 72, 137 (list), tabs 1–
34–35.
Brenneis et al. 2011a: 321;
et al.et al.
–
et al. 2017: appendices 1–
(phylogeny); 2019bLehmann et al. 2021León-Espinosa et al.
2021Ramírez-Tello et al. 2022: tab. 1.
Callipallene brevirostris brevirostris – Stock 1970a: 9.
Callipallene brevirostrum – Stock 1975a: 1010.
Callipallene brevirostris ? – Child 2004: 151.
nec ?Phoxichilus spinosus Montagu, 1808 –
Type material
Pallene brevirostris Johnston, 1837. Type(s): unknown. Type locality: Berwickshire, southeast Scotland.
Pallene empusa Wilson, 1878. Syntypes: USNM 37777 (not examined). Type locality: Vineyard Sound
or Long Island Sound, or Noank (Connecticut, USA).
Material examined
MARTINIQUE • depth 15 m; 10 Sep. 2016; st. AB062; MNHN-
depth 2 m; 12 Sep. 2016; st.
depth 2 m;
24 Sep. 2016; st. AB452; MNHN-IU-2016-
IU-2016-1312.
Remarks
specimens). The species was found on both the Atlantic and Caribbean coasts.
Distribution
Widespread in both sides of the Atlantic: North Sea (Norway, Denmark, Scotland, England and the
Netherlands), West Scotland, Irish Sea, West Ireland, Scilly Islands, English Channel, Gulf of Biscay,
SABROUX R. et al., Pycnogonida from the Madibenthos Expedition to Martinique
51
Portugal, Mediterranean, Black Sea, Bay of Fundy, Massachusetts, Virginia, West Florida, Caribbean,
French Guiana). Also recorded from Pakistan, Thailand and Indonesia.
Depth range
0–316 m.
Callipallene cinto Müller & Krapp, 2009
Callipallene cinto
Callipallene cinto – Sabroux et al.
Type material
Holotype: SMF1497 (not examined). Type locality: Bahía Cinto in Tayrona National Park, Colombia.
Material examined
st. AB189; MNHN-IU-2016-880/MK411190.
Remarks
Atlantic coast.
Distribution
Caribbean Colombia and Martinique.
Depth range
1–3.5 m, extended to 16 m.
Callipallene longicoxa Stock, 1955
Callipallene brevirostris longicoxa
Callipallene belizae
Callipallene longicoxa ––11, 13, 67 (key), 69,
Sabroux et al. 2019b
Type material
Callipallene brevirostris longicoxa Stock, 1955. Holotype: NHMD 110621 (not examined). Type
Callipallene belizae Child, 1982. Holotype: USNM 171035 (not examined). Type locality: Carrie Bow
Cay, Belize.
Material examined
MARTINIQUE • depth 28 m;
data as for preceding; MNHN-IU-2016-1277.
European Journal of Taxonomy 851: 1–141 (2022)
52
Remarks
Grande Anse d’Arlets. Comparatively to other specimens, collected between 10 and 30 meters, the
single specimen collected by Stock (1986) was found strikingly deep into the Strait of Florida (512–
549 m).
Distribution
Caribbean and Strait of Florida.
Depth range
10–512 m.
Genus Pallenoides Stock, 1951
Type species
Pallenoides magnicollis Stock, 1951, by original designation.
ICZN code recommandations for genera ending in -oides.
Pallenoides cf. amazonicus Stock, 1975
Fig. 10
Pallenoides amazonica
Pallenoides sp. nr amazonica – Arnaud & Child 1988: 142–143.
Pallenoides amazonica – Stock
Müller & Krapp 2009: 77.
Pallenoides sp. 1 – Sabroux et al.
Type material
Pallenoides amazonicus Stock, 1975. Holotype: NL ZMA.PYC.P.1356 (not examined). Type locality:
Material examined
MARTINIQUE • depth 66 m; 7 Oct. 2016; st. AD612;
MNHN-IU-2016-866/MK411177.
Remarks
This unique specimen is very close to Stock (1975a)’s original description; however, we note a few
specimen (about half as long as in the holotype) and its conspicuous pointy tubercle on the dorsal
surface of the chelifores. In the absence of a CO1 barcode sequence for P. amazonicus, we refrain from
describing this species as new.
SABROUX R. et al., Pycnogonida from the Madibenthos Expedition to Martinique
53
Fig. 10. Pallenoides cf. amazonicus A. Body, dorsal view.
B. Oviger. C. First coxa of third leg, frontal view. D. Proboscis, ventral view. E. Second leg. F. Tarsus and
propodus of second leg. G. Oviger. H. Oviger strigilis, with close-up of some compound teeth. I. Chela.
European Journal of Taxonomy 851: 1–141 (2022)
54
Distribution
Pallenoides amazonicus is known from Brazil, with one juvenile that has been considered as a possible
P. amazonicus collected in South Africa (Arnaud & Child 1988), though this record seems very doubtful.
Depth range
Pallenoides amazonicus is known to dwell between 17 and 136 m. The present specimen was sampled
at 66 m.
Pallenoides spinulosus Stock, 1955
Pallenoides (?) spinulosa
Pallenoides (?) spinulosa – Capriles 1970: 105.
Pallenoides spinulosa –Müller & Krapp 2009: 13, 67 (key), 77,
Pallenoides spinulosum – Sabroux et al. 2019b et al. 2021:
et al. 2022: tab. 1.
Type material
Holotype: NHMD 110703 (not examined). Type locality: between St Jan and St Thomas, Virgin Islands.
Material examined
MARTINIQUE •
depth
depth 4–10 m; 14 Sep. 2016; st. AB123; MNHN-IU-2016-848/
depth 3 m; 15 Sep. 2016; st. AB126;
depth
depth
depth 10 m; 19 Sep. 2016; st. AB195; MNHN-IU-2016-1405.
Remarks
Distribution
Caribbean, Yucatán Channel.
Depth range
1.5–37 m.
Genus Parapallene Carpenter, 1892
Type species
Pallene australiensis Hoek, 1881, by original designation
SABROUX R. et al., Pycnogonida from the Madibenthos Expedition to Martinique
55
Parapallene bermudensis Lebour, 1949
Parapallene bermudensis
Parapallene bermudensis – Child 1982a: 366––
Sabroux et al. 2019b
Parapallene bermudensis? –
Type material
Material examined
MARTINIQUE • depth 24 m; 30 Sep. 2016; st.
AR413; MNHN-IU-2016-1221/MK411083.
Remarks
Parapallene bermudensis thanks
to the characteristic pectinated spines on tibia 2 and the ventrodistal extremity of the tarsus, and on the
(Caribbean coast).
Distribution
Depth range
16.8–54 m.
Family Nymphonidae Wilson, 1878
Genus Nymphon Fabricius, 1794
Type species
Nymphon grossipes O. Frabricius, 1780, by subsequent designation (Child 1998a).
Nymphon aemulum Stock, 1975
Nymphon aemulum
Nymphon aemulum 2009Müller
& Krapp 2009: 12––et al. 2012: 1356
et al.
Nymphon aemulon – Varela 2012: 3–
nec–
Type material
Holotype: NL ZMA.PYC.P.1267 (not examined). Type locality: Dickinson Bay, Antigua.
European Journal of Taxonomy 851: 1–141 (2022)
56
Material examined
same collection data as for preceding; MNHN-
same collection data
as for preceding;
24 m; 30 Sep. 2016; st. AR413; MNHN-IU-2016-1221.
Remarks
Like all Nymphon but one found in Madibenthos material (i.e., Nymphon timons sp. nov.), this species
is part of the aequidigitatumet al. (2012)
(iii) bifurcate chela teeth, (iv) propodus main claw as long as or shorter than auxiliaries, and (v) one or
all propodal claws bearing endal rugosities or setules.
it was found to be rather common.
Distribution
Tropical West Atlantic (Gulf of Mexico, Caribbean, Surinam).
Depth range
0.5–37 m.
Nymphon dorlis sp. nov.
urn:lsid:zoobank.org:act:8B1908A6-04E4-4B2E-B142-8E6E9F0990AD
Fig. 11
Nymphon sp. 1 – Sabroux et al. 2019b
Material examined
Holotype
AB189; MNHN-IU-2016-879/MK411189.
Paratypes
same collection data as for holotype;
st. AR413; MNHN-IU-2016-1221.
SABROUX R. et al., Pycnogonida from the Madibenthos Expedition to Martinique
57
Etymology
Noun in apposition. Named after the Dorlis, also called ‘night husband’, a malevolent, invisible character
on the ocular tubercle and large setose chelae; and only males were found.
Description
Species rather small and stout, trunk completely segmented, cuticle smooth. No dorsomedian
tubercle. Ocular tubercle rounded, as tall as wide, positioned anteriorly to 1st lateral processes, with
2 conspicuous pointy lateral sense organs beside tip and four pigmented eyes. Preocular neck short,
shorter than wide. Oviger base jointing 1st lateral process. Lateral processes without ornamentation, well
separated by less than their own diameter.
Rounded and large, reaching distal margin of chelifore scape in lateral view.
Medium-sized, not reaching beyond lateral processes, carrying several setae at about mid-
length, directed diagonaly. No basal segmentation.
2-articled, reaching far beyond proboscis. Scape 1-articled, 4.5 times as long as wide,
5-articled. 1st article shortest, shorter than wide. 2nd article longest, about 7 times as long as wide,
with distal setae. 3rd article about 0.8 times as long as 2nd, carrying ventrodistal setae. 4th article about
0.6 times as long as 5th, about 3 times as long as wide, with many long ventral setae along its length. 5th
article 0.8 times as long as 3rd, carrying many ventral setae.
10-articled. 1st article shorter than wide. 2nd article about 1.5 times as long as wide. 3rd article
about twice as long as wide. 4th article about 4 times as long as wide, with a few distal setae. 5th article
longest, about twice as long as 4th, straight, strongly swollen distally, with light constriction at base.
6thth, with many setae along its length. 7th and 10th articles subequal, about
¾ of 6th
teeth, proximalmost shortest and spatulated to oblong, median spines lancelolate, distalmost spines
spatulated to oblong, longer than proximalmost. Intermediary position spines roughly intermediary in
shape. Distalmost spines of 9th and 10th
length of 10th article.
Slender, setose. Coxa 1 no longer than broad. Coxa 2 about 3 times as long as maximal width,
longer than coxae 1 and 3 together. Coxa 3 about as long as wide. Femur about 8 times as long as wide
and more than twice as long as coxa 2, ventrally carrying one row of hardly conspicuous cement glands
and cement gland pores, 44 glands counted on 3rd leg. No cement gland tube. Tibia 1 gently curved,
slender, about as long as femur. Tibia 2 even slenderer, longest, about 1.5 times length of femur or tibia
1, with many tall setae along and two ventral spines on distal margin. Tarsus medium-sized, trapezoid,
one spine distally on ventral surface. Propodus straight, slender, about 3 times as long as tarsus, and ¼
of tibia 2 length. Main claw short, about ¼ of propodal length, well curved. Auxiliary claws present,
well curved, as long as main claw. Both main claw and auxiliary claws with small teeth on inner surface.
(mm). Trunk 1.13; abdomen 0.22; proboscis 0.61; chelifore scape 0.61; chela palm
tarsus 0.15; propodus 0.36; main claw 0.12; auxillary claw 0.12.
European Journal of Taxonomy 851: 1–141 (2022)
58
Fig. 11. Nymphon dorlisA. Body, dorsal view. B. Body,
lateral view (same scale as A). C. Palp. D. Oviger. E. Pointed teeth of oviger. F. Chela. G. Third leg. H.
Tarsus and propodus of third leg.
SABROUX R. et al., Pycnogonida from the Madibenthos Expedition to Martinique
59
Sexual dimorphism
No female available.
Individual variability
Strigilis formula more or less with one compound spine per article in investigated specimens, the highest
Remarks
Nymphon dorlis sp. nov. is a new species in the aequidigitatum-group (Child 1988b). It can be
discriminated from most of its relatives by the following three criteria: (i) preocular neck short, (ii) tarsus
this species can be distinguished from N. biformidens Stock, 1974 from Madagascar by the larger
N. dorlis, as well as its conspicuous horn-like lateral sense organs
(Stock 1974). It can be distinguished from N. aequidigitatum Flynn, 1919 from Australia by the shorter
N. dorlis against ½ in N. aequidigitatum) and the prominent lateral organs
of N. dorlisN. draconis
no conspicuous range of cement gland tubes on the ventral side of the femorae, a longer main claw
N.
draconis; N. megacheles Child, 1988 by the larger space between
the lateral processes, the longer 5th palp segment, the shorter main claw relatively to the propodus and
the slenderer leg articles (Child 1988a).
Among Martinique species, Nymphon dorlis sp. nov. could be mistaken for N. macabou Müller, 1990.
N. dorlis and curved in N. macabou; and the
tibiae 2, which are slender and 1.5 times as long as the femorae, while in N. macabou tibiae 2 are only
slightly longer than the femorae (Müller 1990a).
The species was only collected on the Atlantic coast.
Depth range
Only known from type locality. 16–63 m.
Nymphon ludovici sp. nov.
urn:lsid:zoobank.org:act:53F8FAE5-3380-479C-A5AE-8C6B30CAC9A5
Figs 12–13
Nymphon sp. 3 – Sabroux et al. 2019b
Material examined
Holotype
Sep. 2016; st. AR050; MNHN-IU-2016-835/MK411150.
Paratypes
15 m; 9 Sep. 2016; st. AB058; MNHN-IU-2016-837/MK411152 • 1 ind. (sex indet.); Passe du Marin;
European Journal of Taxonomy 851: 1–141 (2022)
60
same collection data as for preceding; MNHN-IU-2016-562 • 1 juv.; same collection data as for
preceding; MNHN-IU-2016-1274/MK411094 • 1 juv.; same collection data as for preceding; MNHN-