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

Di ec eu De la publica ion / Publica ion di ec o : Gilles Bloch
P ésiden du Muséum na ional d’His oi e na u elle
éDac ice en che / edi o -in-chie : Lau e Desu e -G andcolas
assis an e De éDac ion / assis an edi o : Anne Mabille ([email p o ec ed])
Mise en page / Page layou : Anne Mabille
coMi é scien i ique / scien i ic boa d :
Nes ine Akka i (Na u his o isches Museum, Vienne, Au iche)
Ma ia Ma a Cigliano (Museo de La Pla a, La Pla a, A gen ine)
Se ge Go as (Uni e sidad de Málaga, Málaga, Espagne)
Syl ain Hugel (CNRS, Uni e si é de S asbou g, F ance)
Ma co Isaia (Uni e si à degli S udi di To ino, Tu in, I alie)
Ra ael Ma quez (CSIC, Mad id, Espagne)
Jose Ch is ophe E. Mendoza (Lee Kong Chian Na u al His o y Museum, Singapou )
Annema ie Ohle (MNHN, Pa is, F ance)
Jean-Y es Rasplus (INRA, Mon e ie -su -Lez, F ance)
Wanda M. Weine (Polish Academy o Sciences, C aco ie, Pologne)
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Fe nandes M. R. 2024. — T ipho idae (Gas opoda) om Ma inique sampled by he MADIBENTHOS expedi ion,
wi h no es on shallow-wa e species om Guadeloupe. Zoosys ema 46 (18): 457-503. h ps://doi.o g/10.5252/zoosys-
ema2024 46a18. h p://zoosys ema.com/46/18
ABSTRACT
This s udy aims o e alua e he axonomic composi ion o ipho ids om Ma inique ob ained by
he MADIBENTHOS expedi ion, which comp ised mo e han 500ma ine sampling e en s. In ad-
di ion, some iden i ica ions o shallow-wa e ipho ids om Guadeloupe (based on ecen wo ks)
a e co ec ed. A e he analysis o 1615 specimens, 33 ipho id species we e ound in he shallow
wa e s o Ma inique, o which Inella spinosa n.sp., Iso ipho a pa dus n.sp. and Me axia discus n.sp.
a e desc ibed as new species. New eco ds a e gi en o he Lesse An illes, such as Chei odon a du-
pliniana (Olsson, 1916), S obilige a inaudi a (Rolán & Lee, 2008) and T ipho a ma ii Rolán &
Fe nández-Ga cés, 1995. The axonomy o some species is discussed (mainly on Nanapho a Lase on,
1958), and new gene ic alloca ions a e p oposed: Ma shallo a abacoensis (Rolán & Red e n, 2008) n.
comb., Ma shallo a a mandoi (Espinosa & O ea, 2020) n. comb., Ma shallo a g enadensis (Rolán &
Lee, 2008) n. comb., Ma shallo a mon ei oi (Rolán & Fe nández-Ga cés, 2015) n. comb. The indi-
idual a e ac ion cu e o ipho ids om MADIBENTHOS is nea ly sa u a ed bu did no each
he asymp o e. Se en species (all wi h plank o ophic de elopmen ) we e conside ed abundan o
much abundan , whe eas eigh species (wi h plank o ophic o non-plank o ophic de elopmen )
we e ep esen ed by single ons o double ons. B ushing and suc ion we e e icien me hods o ob ain
ipho ids, and some d edging e en s eco e ed up o 13species (mainly deepe han 50m). Mo e
species we e sampled in he A lan ic side o Ma inique, bu hype -di e se s a ions we e widesp ead
along he island. The abundance o ipho ids and species composi ion change in Ma inique along
he dep h g adien ; Ini o is u is homae (Hol en, 1802) is he mos abundan ipho id be ween
0-30m, bu Cosmo ipho a melanu a (C.B.Adams, 1850) p edomina es be ween 31-85m.
Mau ício Romulo FERNANDES
Depa amen o de Zoologia, Ins i u o de Biociências,
Uni e sidade Fede al do Es ado do Rio de Janei o (UNIRIO),
A enida Pas eu , 458, U ca, Rio de Janei o (B azil)
[email p o ec ed]
Submi ed on 26 July 2023 | Accep ed on 10 Janua y 2024 | Published on 17 July 2024
T ipho idae (Gas opoda) om Ma inique
sampled by he MADIBENTHOS expedi ion,
wi h no es on shallow-wa e species om Guadeloupe
KEY WORDS
Biodi e si y,
Lesse An illes,
la al de elopmen ,
dep h zones,
new combina ions,
new species.
458 ZOOSYSTEMA • 2024 • 46 (18)
Fe nandes M.R.
RÉSUMÉ
Les T ipho idae (Gas opoda) de Ma inique échan illonnés lo s de l’expédi ion MADIBENTHOS, a ec
des ema ques su les espèces des eaux peu p o ondes de Guadeloupe.
Ce e é ude ise à é alue la composi ion axonomique des ipho idés de la Ma inique collec és
pendan l’expédi ion MADIBENTHOS, qui comp enai plus de 500 é énemen s de collec es. De
plus, ce aines iden i ica ions des ipho idés peu p o onds de Guadeloupe (basées su des a aux
écen s) son co igées. Ap ès l’analyse de 1 615 spécimens, 33 espèces de ipho idés on é é dénom-
b ées dans les eaux peu p o ondes de la Ma inique, don Inella spinosa n.sp., Iso ipho a pa dus
n.sp. e Me axia discus n.sp. déc i s comme nou elles espèces. De nou eaux signalemen s son
p oposés pou les Pe i es An illes, comme Chei odon a dupliniana (Olsson, 1916), S obilige a inau-
di a (Rolán & H.G.Lee, 2008) e T ipho a ma ii Rolán & Fe nández-Ga cés, 1995. La axonomie
de ce aines espèces es discu ée (p incipalemen pou Nanapho a Lase on, 1958), e de nou elles
alloca ions géné iques son p oposées: Ma shallo a abacoensis (Rolán & Red e n, 2008) n. comb.,
Ma shallo a a mandoi (Espinosa & O ea, 2020) n. comb., Ma shallo a g enadensis (Rolán & Lee,
2008) n.comb., Ma shallo a mon ei oi (Rolán & Fe nández-Ga cés, 2015) n. comb. La cou be de
a é ac ion indi iduelle des ipho idés de MADIBENTHOS es p esque sa u ée mais n’a pas a ein
l’asymp o e. Sep espèces ( ou es à dé eloppemen planc o ophe) son considé ées comme abondan es
ou ès abondan es, andis que hui espèces (à dé eloppemen planc o ophe ou non-planc o ophe)
son ep ésen ées pa des “single ons” ou des “double ons”. Le b ossage e l’aspi a ion se son é élés
e icaces pou écol e des ipho idés, e ce ains d agages on pe mis de écol e jusqu’à 13espèces
(p incipalemen à des p o ondeu s in é ieu es à 50m). Plus d’espèces on é é échan illonnées su
la açade a lan ique de la Ma inique, mais des s a ions hype -di e si iées é aien épandues le long
de l’île. L’abondance des ipho idés e la composi ion des espèces changen en Ma inique le long
du g adien de p o ondeu ; Ini o is u is homae (Hol en, 1802) es le ipho idé le plus abondan
en e 0 e 30m, alo s que Cosmo ipho a melanu a (C.B. Adams, 1850) p édomine en e 31 e 85m.
INTRODUCTION
T ipho idae G ay, 1847 is ega ded as one o he “Big Fi e” ami-
lies o ma ine gas opods in e ms o species ichness (Bouche
e al. 2002; Albano e al. 2011), which may be a consequence
o hei pa asi ic/mic o-p eda o y eeding on sponges, i.e., he
specializa ion on pa icula sponge hos s possibly igge ed
specia ion and high di e si y (Ma shall 1983; Fe nandes&
Pimen a 2020; Nü zel 2021). In ac , many axonomic p ob-
lems a e de i ed om his g ea di e si y, e en mo e when
only emp y (and o en wo n) shells a e a ailable (Bouche &
S ong 2010). The disco e y o Ca ibbean ipho ids began
in he ea ly 19 h cen u y (Hol en 1802), bu eached a highe
g ound a e he s udy o species om Jamaica (C. B. Adams
1850). Since hen, se e al au ho s inc eased he knowledge
abou Ca ibbean ipho ids a e gene al ca alogues o mol-
luscs om pa icula locali ies (e.g. Dall& Simpson 1901;
Wa mke& Abbo 1962; Nowell-Us icke 1969; Vokes&
Vokes 1983; De Jong& Coomans 1988; Me lano& Hegedus
1994; Hewi & an Leeuwen 2017) o s udies ocused on he
desc ip ion o new species (e.g. Moolenbeek& Fabe 1989;
Fabe & Moolenbeek 1991; Rolán& Luque 1999). Recen
axonomic e isions (wi h good illus a ions) o T ipho idae
om Ca ibbean si es include ma e ial om Cuba (e.g. Rolán&
Fe nández-Ga cés 2007, 2008) and Bahamas (Red e n 2013).
Based on Rosenbe g (2009), he ipho ids o Guadeloupe
and Ma inique we e un ouched un il he desc ip ion o
T ipho a guadaloupensis Rolán& Fe nández-Ga cés, 2008,
endemic om Guadeloupe. Fu he , Rolán& Fe nández-
Ga cés (2015) s udied he shallow-wa e ipho ids om
Guadeloupe, sampled by he KARUBENTHOS expedi ion
(he ea e , KARUBENTHOS 1). Mos o hei iden i ica ions
we e ollowed by Lamy& Poin ie (2018), which also included
new ma e ial om Guadeloupe and Ma inique; o he la e
si e, only Inella ise ialis (Dall, 1881), Ini o is u is homae
(Hol en, 1802), Ma shallo a nig ocinc a (C. B.Adams, 1839),
Monopho us oli aceus (Dall, 1889) and No o ipho a deco a a
(C. B. Adams, 1850) we e eco ded. These wo ks g ea ly
imp o ed he knowledge abou ipho ids om he F ench
An illes, bu he e we e mis akes on some o hese axonomic
iden i ica ions, equi ing co ec ions.
The MADIBENTHOS expedi ion was o ganized by he
MNHN (Muséum na ional d’His oi e na u elle, F ance) eam
and conduc ed in Ma inique be ween Sep embe -Oc obe
2016, comp ising 503 sampling e en s in he sho e and sea,
such as d edging, ap lines, b ushing o ha d subs a a and
suc ion sampling (Bouche e al. 2019). Sampling s a ions
we e pe o med down o 100m, co e ing all po ions o he
island (Bouche e al. 2019). Se e al new species o gas opods
we e ecen ly desc ibed om ma e ial ob ained by he expe-
di ion (e.g. Espinosa& O ea 2017; O ea& Buske 2018;
Ga igues& Lamy 2019; Cecalupo& Pe ugia 2020), and
o he s a e s ill o come.
The objec i e o his s udy is o e alua e he axonomic
composi ion o ipho ids ob ained by he MADIBENTHOS.
In addi ion, some iden i ica ions o shallow-wa e ipho ids
om Guadeloupe (Rolán& Fe nández-Ga cés 2015; Lamy&
Poin ie 2018) a e co ec ed. The deep-sea species om Gua-
deloupe, ob ained by he KARUBENTHOS 2 expedi ion,
will be ea ed in ano he s udy.
MOTS CLÉS
Biodi e si é,
Pe i es An illes,
dé eloppemen la ai e,
zones de p o ondeu ,
combinaisons nou elles,
espèces nou elles.
459
T ipho idae om Ma inique
ZOOSYSTEMA • 2024 • 46 (18)
MATERIAL AND METHODS
Sampling and axonomic p ocedu eS
S a ions in which ipho ids we e ob ained a e lis ed unde
each species, ollowing he i e main geog aphic zones wi hin
Ma inique p oposed by Bouche e al. (2019); hei coo di-
na es, dep hs and sampling me hods a e lis ed in Appendix1.
All shallow-wa e species om Guadeloupe examined by he
au ho a e also lis ed, ob ained by KARUBENTHOS 1 (Rolán&
Fe nández-Ga cés 2015) and KARUBENTHOS 2 expedi ions.
A ew shells om Guadeloupe we e illus a ed; in such cases
as e isks (*) a e gi en in he pla es. Li e ipho ids om Gua-
deloupe we e s o ed in e hanol du ing MNHN expedi ions,
and hese will be s udied in an ongoing molecula phylogeny
o T ipho oidea (unpublished da a); hose om Ma inique
we e mainly s o ed in d y capsules, wi h a ew excep ions
s o ed in e hanol. Du ing expedi ions, li e specimens we e
pho og aphed unde a Canon EOS 50D (mac o lens 65mm
o 90mm) came a by he MNHN eam. Li e ipho ids om
Guadeloupe a e also ma ked wi h as e isks (*) in he pla es.
A he labo a o y, shells we e pho og aphed unde a Nikon
D5000 came a coupled o a s e eomic oscope Leica MZ16.
Shells we e no coa ed p io o SEM (Scanning Elec on Mi-
c oscope) images in o de o p ese e hei colo ; he SEM
equipmen used was a JEOL 6490-LV. In legend cap ions,
shell size e e s o leng h. Desc ip ions a e solely based on
he ma e ial examined, ollowing p ocedu es and e minol-
ogy on Fe nandes& Pimen a (2015, 2019a). Emp y shells
and li e specimens a e espec i ely indica ed in he ma e ial
examined by ‘sh’ and ‘spm’. Mos species a e ea ed in he
axonomic sec ion b ie ly, wi h educed da a on synonymic
lis s and no desc ip ions o lis s o geog aphic dis ibu ion; o
hose species, see p e ious s udies (e.g. Fe nandes& Pimen a
2020) o comple e da a.
abb e ia ionS
ANSP The Academy o Na u al Sciences o D exel Uni-
e si y, Philadelphia;
BMSM Bailey-Ma hews Na ional Shell Museum, Sanibel;
FLMNH Flo ida Museum o Na u al His o y, Gaines ille;
MNCN Museo Nacional de Ciencias Na u ales, Mad id;
MNHN Muséum na ional d’His oi e na u elle, Pa is;
MNRJ Museu Nacional, Uni e sidade Fede al do Rio de
Janei o, Rio de Janei o;
NHMUK Na u al His o y Museum, London;
USNM/NMNH
Na ional Museum o Na u al His o y, Washing on
D.C.;
ZMA.MOLL Mollusca collec ion, Na u alis Biodi e si y Cen e ,
Leiden.
ecological analySeS
The indi idual a e ac ion cu e and he abundance dis ibu-
ion o ipho ids om MADIBENTHOS we e ob ained in
he so wa e PAST 4.02, ega ding he o al numbe o shells/
specimens, as well as indices o dominance, Shannon’s di e -
si y, Pielou’s equi abili y and Buzas& Gibson’s e enness (all
wi h 9999 boo s ap eplica ions). The es ima i e o species
ichness (i.e., o e alua e he numbe o missing species) was
calcula ed wi h a Chao1 es ima o (100 andomiza ion uns;
bias-co ec ed) in he so wa e Es ima eS 9.1. The so wa e
Excel was used o simple g aphs, such as he numbe and
p opo ion o species sampled by each sampling me hod
(AB,b ushing; AD, d edging; AM, isual a in e idal; AR,
isual a di ing; AS, suc ion), ollowing he codes in Bouche
e al. (2019); s a ions AN ( ap lines) we e excluded because
hey did no con ain ipho ids. Shells iden i ied as Nanapho a
c . e be nei (Moolenbeek& Fabe , 1989) we e included
wi hin N. e be nei in all ecological analyses.
A he compa ison be ween species ound in Ma inique s
o he si es in he Wes A lan ic, only species om he con-
inen al shel we e conside ed, i.e., wi h a leas one eco d
shallowe han 150m; he eco ds o Inella ise ialis and
“Inella” a . ha yleei Rolán& Fe nández-Ga cés, 2008 espec-
i ely om Ma inique and Guadeloupe (Lamy& Poin ie
2018) we e ega ded as om he deep-sea, e en hough he
minimum dep h was indica ed as 55m (see Discussion). In
addi ion, some mo phs wi hou o mal names we e consid-
e ed (e.g. h ee undesc ibed species o Ma shallo a Bouche ,
1985 in B azil – Fe nandes e al. 2021), bu nomina dubia
such as Me axia icina (C. B. Adams, 1850), eco ded o he
A uba-Bonai e-Cu açao Is. (he ea e ABC Is.) (De Jong&
Coomans 1988), we e igno ed. To e alua e whe he ipho ids
om Ma inique a e endemic o ha e a wide ange, a species
is he e conside ed o belong o he en i e Wes A lan ic (and
no es ic ed o he Lesse An illes o he NW A lan ic) i
c ossing he Amazon il e . Fo example, Iso ipho a icingu-
la a Rolán& Fe nández-Ga cés, 2015 occu s in pa o he
Ca ibbean and in Amapá s a e (No h B azil) (Fe nandes&
Pimen a 2020), bu i is absen below he Amazon mou h,
hus i is ega ded as a NW A lan ic species.
Fo ba hyme ic compa isons, only shells no much wo n
we e conside ed, o a oid bias o pos -mo em dislodgemen .
Shells we e assigned o each o i e dep h zones (0-10m; 11-
20m; 21-30m; 31-60m; 61-85m). When wide dep h anges
a e gi en in a s a ion, he p edominan ange was conside ed
(e.g. in a dep h o 8-16m, hus ange 11-20m was selec ed)
o only he minimum dep h, i dis ibu ed in equal pa s (e.g.
in a dep h o 8-12m, hus ange 0-10m was selec ed). Two
o dina ions o nMDS (non-me ic mul idimensional scaling)
we e calcula ed in PAST 4.02 o he dep h and geog aphic
zones wi hin Ma inique (as p oposed by Bouche e al. 2019:
217), one ega ding he ela i e abundance (pe cen age) o
each species pe zone (using a B ay-Cu is index a e an
‘a csin squa e oo ’ ans o ma ion), he o he ega ding he
p esence/absence o each species pe zone (using a Jacca d
index); dep h/geog aphic zones wi h less han 20 ipho id
shells we e excluded in o de o educe sampling bias.
RESULTS
A e he analysis o 1615shells/specimens, 33 ipho id spe-
cies we e sampled in he shallow wa e s o Ma inique by he
MADIBENTHOS expedi ion, o which h ee a e desc ibed as
new species. The axonomic indings a e ea ed indi idually
in he nex sec ion (Figs1-17), also comp ising shallow-wa e

460 ZOOSYSTEMA • 2024 • 46 (18)
Fe nandes M.R.
eco ds om Guadeloupe (mos ly KARUBENTHOS 1, wi h
a ew eco ds om KARUBENTHOS 2), and li e specimens
a e illus a ed u he (Figs18-20). The ecological analyses
(Figs21-26) a e ea ed in he Discussion.
Family ipho idae G ay, 1847
Genus Chei odon a Ma shall, 1983
Chei odon a dupliniana (Olsson, 1916)
(Fig.1A)
T ipho a dupliniana Olsson, 1916: 138, pl.3 ig.8.
ma e ial examined.— Guadeloupe. KARUBENTHOS 1 • 1sh
[iden i ied by Rolán& Fe nández-Ga cés (2015): ig.4L as Ma -
shallo a modes a]; s a. GS22; MNHN • 1sh [p e iously iden i ied
as Sageno ipho a osclausum]; s a. GB25; MNHN • 1sh [idem];
s a. GB34; MNHN.
Ma inique. MADIBENTHOS • 1sh; ‘No d A lan ique’; s a.
AD244; MNHN • 1sh; ‘No d Ca aibe’; s a. AM040; MNHN • 2sh;
‘No d Ca aibe’; s a.AS570; MNHN • 1sh; s a. AS576; MNHN.
ba hyme ic diS ibu ion.— Reco ded dep h in Guadeloupe:
10-45m. Reco ded dep h in Ma inique: 0-59m. P e ious eco ded
dep h in he Wes A lan ic: 6-183m (Fe nandes& Pimen a 2020).
Genus Co iopho a Lase on, 1958
Co iopho a no em (Nowell-Us icke, 1969)
(Figs1B; 18A, B)
T ipho a no em Nowell-Us icke, 1969: 12, pl.2 ig.403.
ma e ial examined.— Guadeloupe. KARUBENTHOS 1 • 1sh;
s a. GB06; MNHN • 1sh; s a. GS07; MNHN • 2sh; s a. GB08;
MNHN • 1 spm; s a. GM08; MNHN • 1sh; s a. GB09; MNHN
• 1sh; s a. GD10; MNHN • 2sh; s a. GR13; MNHN • 5sh;
s a. GS13; MNHN • 1sh; s a. GB20; MNHN • 1sh; s a. GS24;
MNHN • 1sh; s a. GD25; MNHN • 1sh [ju enile]; s a. GD31;
MNHN • 2sh; s a. GD58; MNHN • 1sh; s a. GD59; MNHN.—
KARUBENTHOS 2 • 1sh [wo n]; s a. DW4545; MNHN.
Ma inique. MADIBENTHOS. ‘No d A lan ique’ • 1sh; s a. AB197;
MNHN • 1spm; s a. AD254; MNHN • 1sh; s a. AB260; MNHN
• 2sh; s a. AD260; MNHN • 3sh; s a. AD261; MNHN • 4sh; s a.
AD263; MNHN • 3sh; s a. AD275; MNHN • 3sh; s a. AB350;
MNHN • 2sh; s a. AB562; MNHN • 3sh; s a. AS565; MNHN.
— ‘Sud A lan ique’ • 2sh; s a. AS081; MNHN • 2sh; s a. AB181;
MNHN • 1sh [ju enile]; s a. AB183; MNHN • 1sh; s a. AS255;
MNHN • 1sh; s a. AB400; MNHN • 2sh; s a. AB401; MNHN •
1sh; s a. AB405; MNHN • 1sh [wo n]; s a. AB419; MNHN.—‘No d
Ca aibe’ • 1sh; s a. AB161; MNHN • 2sh; s a. AD280; MNHN •
4sh; s a. AD290; MNHN • 1sh; s a. AB362; MNHN • 1sh; s a.
AB460; MNHN • 2sh; s a. AS574; MNHN • 2sh; s a. AS576;
MNHN. ‘Sud Ca aibe’ • 3sh; s a. AB058; MNHN • 1sh; s a. AB060;
MNHN • 4sh; s a. AS071; MNHN • 1sh; s a. AB123; MNHN •
1sh [ju enile]; s a. AB155; MNHN • 1sh; s a. AB169; MNHN •
1sh [ju enile]; s a. AD218; MNHN • 1sh [ju enile]; s a. AB358;
MNHN • 2sh; s a. AB369; MNHN • 1sh; s a. AD616; MNHN.—
‘Baie de Fo -de-F ance’ • 1sh; s a. AB398; MNHN.
ba hyme ic diS ibu ion.— Reco ded dep h in Guadeloupe:
1-160m (li e specimen a 1m). Reco ded dep h in Ma inique:
4-80m (li e specimen a 62m). P e ious eco ded dep h in he
Wes A lan ic: 2-110m (Fe nandes& Pimen a 2020).
Genus Cosmo ipho a Olsson& Ha bison, 1953
Cosmo ipho a a noldoi Fabe & Moolenbeek, 1991
(Fig.1C)
Cosmo ipho a a noldoi Fabe & Moolenbeek, 1991: 81, igs1-2.
ma e ial examined.— Ma inique. MADIBENTHOS. ‘No d
A lan ique’ • 1sh; s a. AD260; MNHN.
ba hyme ic diS ibu ion.— Reco ded dep h in Ma inique:
78m. P e ious eco ded dep h in he Wes A lan ic: 20-155m
(Fe nandes& Pimen a 2020).
Cosmo ipho a melanu a (C. B. Adams, 1850)
(Figs1D; 18C-F)
Ce i hium melanu a C. B. Adams, 1850: 117.
ma e ial examined.— Guadeloupe. KARUBENTHOS 1 •
1sh; s a. GB01; MNHN • 3sh, 1 spm; s a. GB02; MNHN • 1sh,
1spm; s a. GB03; MNHN • 3sh; s a. GS04; MNHN • 2sh; s a.
GS05; MNHN • 2sh, 2 spm; s a. GB06; MNHN • 4sh; s a. GS06;
MNHN • 2sh; s a. GS07; MNHN • 2sh; s a. GB08; MNHN •
2sh [ju eniles]; s a. GM08; MNHN • 11sh; s a. GS08; MNHN
• 8sh; s a. GB09; MNHN • 3sh; s a. GS09; MNHN • 3sh; s a.
GD10; MNHN • 3sh; s a. GR10; MNHN • 14sh; s a. GB11;
MNHN • 1sh; s a. GS11; MNHN • 1sh; s a. GB12; MNHN •
1sh; s a. GS12; MNHN • 6; s a. GB13; MNHN • 25sh; s a. GS13;
MNHN • 1sh; s a. GB14; MNHN • 1sh; s a. GR14; MNHN
• 2sh; s a. GS14; MNHN • 1sh; s a. GB15; MNHN • 1sh; s a.
GS15; MNHN • 13sh; s a. GB16; MNHN • 1sh; s a. GS16;
MNHN • 3 spm; s a. GR17; MNHN • 3sh, 1 spm; s a. GS17;
MNHN • 2sh; s a. GB18; MNHN • 3sh; s a. GS18; MNHN •
1sh; s a. GB19; MNHN • 3sh; s a. GR19; MNHN • 6sh, 1 spm;
s a. GB20; MNHN • 1sh; s a. GD20; MNHN • 5sh; s a. GS20;
MNHN • 1sh; s a. GD21; MNHN • 1sh; s a. GR21; MNHN
• 4sh; s a. GB22; MNHN • 2sh; s a. GS22; MNHN • 1sh; s a.
GB23; MNHN • 2sh; s a. GS23; MNHN • 1sh; s a. GB24;
MNHN • 1sh; s a. GD24; MNHN • 2sh; s a. GS24; MNHN
• 2sh; s a. GS25; MNHN • 1sh; s a. GB26; MNHN • 5sh; s a.
GS29; MNHN • 2sh; s a. GB30; MNHN • 10sh; s a. GB31;
MNHN • 1sh; s a. GD31; MNHN • 1sh; s a. GS32; MNHN •
1sh; s a. GD33; MNHN • 3sh; s a. GB34; MNHN • 4sh; s a.
GS34; MNHN • 1sh; s a. GB36; MNHN • 3sh; s a. GR36; MNHN
• 4sh; s a. GS36; MNHN • 1sh; s a. GS37; MNHN • 1sh; s a.
GD52; MNHN • 1sh; s a. GD55; MNHN • 2sh; s a. GD58;
MNHN • 1sh; s a. GD59; MNHN • 1sh; s a. GD69; MNHN.
— KARUBENTHOS 2 • 9sh; s a. DW4545; MNHN • 1sh;
s a. DW4546; MNHN • 1sh; s a. DW4550; MNHN • 1sh; s a.
DW4555; MNHN • 1sh; s a. DW4586; MNHN • 1sh [ju enile];
s a. DW4587; MNHN • 1sh [ju enile]; s a. DW4599; MNHN.
Ma inique. MADIBENTHOS. ‘No d A lan ique’ • 4sh; s a.
AB197; MNHN • 2sh; s a. AD244; MNHN • 2sh; s a. AD248;
MNHN • 1sh [ju enile]; s a. AD250; MNHN • 2sh; s a. AD254;
MNHN • 2sh; s a. AD257; MNHN • 1sh; s a. AB260; MNHN •
7sh; s a. AD260; MNHN • 7sh; s a. AD261; MNHN • 10sh; s a.
AD263; MNHN • 7sh; s a. AD275; MNHN • 1sh; s a. AB350;
MNHN • 1sh; s a. AS557; MNHN • 3sh; s a. AB562; MNHN
• 3sh; s a. AS563; MNHN • 13sh; s a. AS565; MNHN. — ‘Sud
A lan ique’ • 4sh; s a. AS078; MNHN • 1sh; s a. AS081; MNHN
• 6sh; s a. AB181; MNHN • 2sh; s a. AB189; MNHN • 2sh; s a.
AB193; MNHN • 1sh; s a. AD222; MNHN • 2sh; s a. AD224;
MNHN • 1sh; s a. AD232; MNHN. — ‘No d Ca aibe’. 2sh; s a.
AS054; MNHN • 1sh; s a. AB062; MNHN • 2sh; s a. AB108;
MNHN • 1sh; s a. AD109; MNHN • 6sh; s a. AB159; MNHN
461
T ipho idae om Ma inique
ZOOSYSTEMA • 2024 • 46 (18)
• 3sh; s a. AB183; MNHN • 2sh; s a. AD276; MNHN • 3sh; s a.
AD280; MNHN • 2sh; s a. AD289; MNHN • 12sh; s a. AD290;
MNHN • 3sh; s a. AD298; MNHN • 1sh; s a. AD299; MNHN
• 2sh; s a. AR308; MNHN • 3sh; s a. AS363; MNHN • 2sh; s a.
AB372; MNHN • 1sh; s a. AS373; MNHN • 7sh; s a. AS375;
MNHN • 1sh; s a. AB377; MNHN • 1sh; s a. AS378; MNHN •
3sh; s a. AB386; MNHN • 1sh; s a. AB460; MNHN • 5sh; s a.
AB463; MNHN • 2sh; s a. AB567; MNHN • 3sh; s a. AS570;
MNHN • 2sh; s a. AS572; MNHN • 3sh; s a. AS574; MNHN •
2sh; s a. AS576; MNHN • 3sh; s a. AD612; MNHN. ‘Sud Ca aibe’
ABCD E
FGHI J
Fig. 1. — T ipho idae G ay, 1847 om Ma inique: A, Chei odon a dupliniana (Olsson, 1916), MNHN, s a. AS576, 5.4 mm; B, Co iopho a no em (Nowell-Us icke,
1969), MNHN, s a. AD280, 5.0 mm; C, Cosmo ipho a a noldoi Fabe & Moolenbeek, 1991, MNHN, s a. AD260, 2.6 mm; D, Cosmo ipho a melanu a (C. B. Ad-
ams, 1850), MNHN, s a. AB123, 4.1 mm; E, Eu ipho a be mudensis (Ba sch, 1911), MNHN, s a. AS257, 4.2 mm; F, Ini o is gudeliae Rolán & Fe nández-Ga cés,
2009, MNHN, s a. AS403, 4.7 mm; G, Ini o is pseudo homae Rolán & Fe nández-Ga cés, 1993, MNHN, s a. AB062, 7.0 mm; H, Ini o is u is homae (Hol en,
1802), MNHN, s a. AS075, 5.7 mm; I, Iso ipho a icingula a Rolán & Fe nández-Ga cés, 2015, MNHN, s a. AD263, 4.9 mm; J, Iso ipho a sp., MNHN, s a. AS403,
2.6 mm. Scale ba s: 1 mm.
462 ZOOSYSTEMA • 2024 • 46 (18)
Fe nandes M.R.
• 2sh; s a. AS057; MNHN • 1sh; s a. AB058; MNHN • 1sh; s a.
AD067; MNHN • 3sh; s a. AS068; MNHN • 4sh; s a. AS071;
MNHN • 2sh; s a. AS075; MNHN • 3sh; s a. AS092; MNHN •
1spm; s a. AS096; MNHN • 1sh, 1spm; s a. AB123; MNHN •
6sh; s a. AB126; MNHN • 2sh; s a. AB150; MNHN • 5sh; s a.
AB152; MNHN • 1sh; s a. AS154; MNHN • 1sh; s a. AB163;
MNHN • 4sh; s a. AB169; MNHN • 1sh; s a. AB173; MNHN
• 1sh; s a. AB177; MNHN • 1sh, 1spm; s a. AD214; MNHN •
3sh; s a. AD216; MNHN • 5sh; s a. AD218; MNHN • 1sh; s a.
AD220; MNHN • 1sh; s a. AD271; MNHN • 3sh; s a. AB354;
MNHN • 1sh [ju enile]; s a. AB358; MNHN • 1sh; s a. AR359;
MNHN • 1sh; s a. AB360; MNHN • 2sh; s a. AB369; MNHN
• 1sh; s a. AS370; MNHN • 1sh; s a. AB578; MNHN • 2sh; s a.
AS579; MNHN • 2sh; s a. AD615; MNHN • 1sh; s a. AD616;
MNHN. — ‘Baie de Fo -de-F ance’ • 1sh; s a. AB392; MNHN
• 1sh; s a. AB398; MNHN.
ba hyme ic diS ibu ion.— Reco ded dep h in Guadeloupe:
0-150m (li e specimens a 3-23m); some in ac adul shells ob-
ained by KARUBENTHOS 2 be ween 204-482m, bu hey mus
be ega ded wi h cau ion. Reco ded dep h in Ma inique: 0-80m
(li e specimens a 4-70m). P e ious eco ded dep h in he Wes
A lan ic: 0-160m, wi h disc epan eco ds down o 480m (Fe -
nandes& Pimen a 2020).
Genus Eu ipho a Co on& God ey, 1931
Eu ipho a be mudensis (Ba sch, 1911)
(Fig.1E)
T ipho is be mudensis Ba sch, 1911: 305, pl.28, igs2, 4.
ma e ial examined.— Guadeloupe. KARUBENTHOS 1 • 1sh
[p e iously iden i ied as Ini o is u is homae]; s a. GS05; MNHN •
1sh [idem]; s a. GS06; MNHN • 1sh [idem]; s a. GB08; MNHN •
1sh [idem]; s a. GS18; MNHN • 1sh [idem]; s a. GB26; MNHN.
Ma inique. MADIBENTHOS. ‘Sud A lan ique’ • 1sh; s a. AS253;
MNHN • 1sh; s a. AS257; MNHN. — ‘No d Ca aibe’ • 1sh; s a.
AB161; MNHN. ‘Sud Ca aibe’: • 1sh; s a. AB169; MNHN. —
‘Baie de Fo -de-F ance’ • 1sh; s a. AB392; MNHN.
ba hyme ic diS ibu ion.— Reco ded dep h in Guadeloupe:
17-49m. Reco ded dep h in Ma inique: 13-50m. P e ious eco ded
dep h in he Wes A lan ic: 4-56m (Rolán& Fe nández-Ga cés 1995).
ema k
The single shell iden i ied by Rolán& Fe nández-Ga cés
(2015: ig.4T) as E. be mudensis om Guadeloupe, and
ollowed by Lamy& Poin ie (2018: pl.91, ig.6), ac ually
e e s o T ipho a ma ii Rolán& Fe nández-Ga cés, 1995.
Genus Inella Bayle, 1879
Inella spinosa n.sp.
(Fig.4)
u n:lsid:zoobank.o g:ac :42468F50-0E06-472E-8F1D-5738EF4F29B5
ype
ma e ial
. — Holo ype. Ma inique. MADIBENTHOS •
sh; Macouba, egion ‘No d A lan ique’; s a. AD275, 14°55’00”N,
61°08’54”W; dep h 80m; MNHN-IM-2000-38595.
Pa a ypes. Ma inique. MADIBENTHOS. ‘No d A lan ique’ •
2sh; s a. AD263; MNHN-IM-2000-38596.
ype locali y.— Ma inique: Macouba, egion ‘No d A lan-
ique’; MADIBENTHOS expedi ion s a. AD275, 14°55’00”N,
61°08’54”W; 80m.
o he ma e ial examined.— Guadeloupe. KARUBENTHOS 2
• 1sh; s a. DW4545; MNHN.
Ma inique. MADIBENTHOS. ‘No d A lan ique’ • 1sh [ju enile,
wo n]; s a. AD260; MNHN.
e ymology.— F om La in, spinosus. The species is named due o
he spiny appea ance o he spi al co ds o eleoconch.
diagnoSiS.— Spi ally-elonga ed, poin ed nodules in median and
abapical co ds, gi ing a spiny appea ance o he eleoconch p o ile;
adapical spi al co d ini ially e y na ow, ne e eaching he same
size o o he co ds; shell wi h a c eam backg ound.
geog aphic diS ibu ion.— Guadeloupe and Ma inique.
ba hyme ic diS ibu ion.— Reco ded dep h in Guadeloupe:
60-82m. Reco ded dep h in Ma inique: 78-80m.
deSc ip ion
Shell sinis al, conical/ usi o m, ec ilinea p o ile, up o
4.4mm long, 1.1mm wide, leng h/wid h a io 3.5-3.9;
adul shells each a leas 2.7mm in leng h. P o oconch
paucispi al wi h unca ed apex, wi hou clea di e en ia ion
om he eleoconch, o 2.75-3.0 who ls, 0.57-0.61mm
long, 0.50-0.52mm wide; he small nucleus ises in an
adapical di ec ion and u he goes abapical in an oblique
descen ; wo smoo h, keeled spi al co ds o equal size, si u-
a ed a 26-34% and 66-71% o las who l heigh , in addi-
ion o a na ow and smoo h subsu u al co d. Teleoconch
wi h up o 8.5 who ls; wo main spi al co ds (median and
abapical) a he beginning, con inuous o hose o p o o-
conch, assuming a keeled-shape along he eleoconch, wi h
median co d always mo e p ominen , excep on he body
who l whe e bo h co ds ha e nea ly he same size; adapical
spi al co d ini ially e y na ow, g adually s eng hening
along he eleoconch bu ne e eaching he same size o
o he co ds o o ming dis inc nodules; a smoo h su u al
co d is also e iden , g adually s eng hening and eaching
he same size o adapical co d in la ge shells; spi ally-
elonga ed, poin ed nodules in median and abapical co ds,
gi ing a spiny appea ance o he eleoconch p o ile; nine
o en o hocline axial ibs on se en h eleoconch who l;
su u e dis inc ; smoo h subpe iphe al co d, one na ow,
smoo h basal co d; sup anume ical co ds absen ; ellip i-
cal ape u e, 0.51-0.67mm long, 0.37-0.44mm wide,
leng h/wid h a io 1.4-1.5; open, e y sho an e io canal,
0.05-0.12mm long, 0.18-0.27mm wide, leng h/wid h
a io 0.3-0.5; pos e io canal absen . Shell wi h a c eam
backg ound, whi e nodules on eleoconch, in e nodula
spaces ligh b own.
ema kS
The spiny appea ance o he spi al co ds o he eleoconch,
combined wi h he colo pa e n and small shell size ( each-
ing only 4.4mm long), makes I. spinosa n.sp. unique among
Inella and o he Wes A lan ic ipho ids. Because o i s
463
T ipho idae om Ma inique
ZOOSYSTEMA • 2024 • 46 (18)
p obable leci ho ophic de elopmen and ba hyme ic ange
es ic ed o he uppe 100m (absen om deepe wa e s o
Guadeloupe, based on samples om KARUBENTHOS 2),
I.spinosa n.sp. p obably has a na ow geog aphic ange in he
Lesse An illes. The single shell ob ained om Guadeloupe
(Fig.4D) has he axial sculp u e much weake han shells
om Ma inique (Fig.4A-C), a enua ing he spiny appea -
ance o he spi al co ds.
ABCDE
FGHI J
Fig. 2
. — T ipho idae G ay, 1847 om Ma inique - con .: A, La i ipho a albida (A. Adams, 1854), MNHN, s a. AB149, 4.0 mm; B, Ma shallo a c . modes a (C. B. Ad-
ams, 1850), MNHN, s a. AB150, 4.3 mm; C, Me axia excelsa Fabe & Moolenbeek, 1991, MNHN, s a. AB123, 3.6 mm; D, Me axia ugulosa (C. B. Adams, 1850),
MNHN, s a. AD265, 6.3 mm; E, Me axia aeniola a (Dall, 1889), MNHN, s a. AD261, 5.5 mm; F, Monopho us oli aceus (Dall, 1889), MNHN, s a. AS576, 9.5 mm;
G, No o ipho a deco a a (C. B. Adams, 1850), MNHN, s a. AB390, 6.8 mm; H, Sageno ipho a osclausum (Rolán & Fe nández-Ga cés, 1995), MNHN, s a. AS565,
3.4 mm; I, Similipho a in e media (C. B. Adams, 1850), MNHN, s a. AD067, 5.6 mm; J, Similipho a sp., MNHN, s a. AS552, 5.1 mm. Scale ba s: 1 mm.
470 ZOOSYSTEMA • 2024 • 46 (18)
Fe nandes M.R.
ema kS
Lamy& Poin ie (2018: pl.92, ig.16) illus a ed he
holo ype o I. icingula a unde he name T ipho asp. 5,
p o iding a dep h eco d o 50m, con a y o he ue dep h
eco d o he ype locali y (i.e., 85m in s a ion GD31 o
KARUBENTHOS 1, no 81m, as eco ded in he o iginal
desc ip ion). I dis ega d he dep h eco d o 50m p o ided
by Lamy& Poin ie (2018), due o he unce ain y o which
shells we e s udied.
Iso ipho asp.
(Fig.1J)
m
a e ial
examined
.— Ma inique.
MADIBENTHOS.
‘Sud
A lan ique’ • 1sh; s a. AS403; MNHN.
ba hyme ic diS ibu ion.— Only known om 11m.
Genus La i ipho a Ma shall, 1983
La i ipho a albida (A. Adams, 1854)
(Fig.2A)
T ipho is albidus A. Adams, 1854: 278.
ma e ial examined.— Guadeloupe. KARUBENTHOS 1 • 1sh;
s a. GS17; MNHN • 1sh; s a. GB36; MNHN.
Ma inique. MADIBENTHOS. ‘No d A lan ique’ • 2sh; s a. AB562;
MNHN. — ‘Sud A lan ique’ • 1sh; s a. AB117; MNHN • 2sh;
s a. AB149; MNHN. — ‘No d Ca aibe’ • 1sh, 1spm; s a. AB386;
MNHN. ‘Sud Ca aibe’ • 1sh [wo n]; s a. AM033; MNHN • 1sh;
s a. AS075; MNHN • 1sh [ju enile]; s a. AB126; MNHN • 1sh; s a.
AD216; MNHN. ‘Baie de Fo -de-F ance’ • 1sh; s a. AB390; MNHN.
ba hyme ic diS ibu ion.— Reco ded dep h in Guadeloupe:
3-16m. Reco ded dep h in Ma inique: 2-26m (li e specimen
a 17m). P e ious eco ded dep h in he Wes A lan ic: 0-107m
(Fe nandes& Pimen a 2020).
ema k
Rolán& Fe nández-Ga cés (2015: ig.4S) illus a ed a shell
o No o ipho a deco a a (C. B. Adams, 1850) unde he name
L. albida, which was ollowed by Lamy& Poin ie (2018:
pl.91, ig.15), and he e we e o he 14 MNHN lo s o N.
deco a a om Guadeloupe p e iously iden i ied as L. albida;
only wo shells o L. albida we e co ec ly iden i ied.
Genus Ma shallo a Bouche , 1985
Ma shallo a c . modes a (C. B. Adams, 1850)
(Fig.2B)
Ce i hium modes um C. B. Adams, 1850: 117.
ma e ial examined.— Guadeloupe. KARUBENTHOS 1 • 2spm;
s a. GM01; MNHN • 1sh, 3 spm; s a. GM02; MNHN • 2spm;
s a. GM03; MNHN • 2sh; s a. GM11; MNHN • 1sh; s a. GM29;
MNHN • 1sh; s a. GS30; MNHN • 3sh; s a. GD39; MNHN •
1sh; s a. GD49; MNHN • 1sh; s a. GD51; MNHN. — Ma i-
nique. MADIBENTHOS. ‘Sud A lan ique’ • 1sh; s a. AM042;
MNHN • 2sh [one wi h he mi c ab]; s a. AM043; MNHN • 1sh;
s a. AM325; MNHN. ‘Sud Ca aibe’ • 1sh; s a. AB150; MNHN.
‘Baie de Fo -de-F ance’ • 1sh; s a. AM034; MNHN • 8sh; s a.
AB390; MNHN • 1sh; s a. AB392; MNHN • 3sh; s a. AB394;
MNHN • 1sh; s a. AD603; MNHN • 1sh; s a. AD607; MNHN
• 1sh; s a. AD623; MNHN.
ba hyme ic diS ibu ion.— Reco ded dep h in Guadeloupe:
0-15m (li e specimens a 1m). Reco ded dep h in Ma inique:
0-13m. P e ious eco ded dep h in he Wes A lan ic: 0-20m
(Rosenbe g 2009).
ema kS
As abo emen ioned, he illus a ion o M. modes a om Gua
-
deloupe by Rolán& Fe nández-Ga cés (2015: ig.4L) ac ually
e e s o C. dupliniana, whe eas hei eco d o Ma shallo a
nig ocinc a (C. B. Adams, 1839) mus be e used, because his
species is known wi h ce ain y only om Canada and USA,
based on DNA e idences and some shell ea u es (Fe nandes
e al. 2021). Acco dingly, he eco d o M. nig ocinc a om
Guadeloupe and Ma inique by Lamy& Poin ie (2018) is
ejec ed. The ma e ial om he F ench An illes will be s udied
unde an in eg a i e axonomic basis, oge he wi h new sam-
ples om he Wes A lan ic (unpublished da a). Meanwhile,
shells a e he ein iden i ied as M. c . modes a, ega ding he
exis ence o a leas wo Ca ibbean species in his complex
(Fe nandes e al. 2021).
Ma shallo a mon ei oi
(Rolán& Fe nández-Ga cés, 2015) n. comb.
(Figs7; 8; 20I-J)
“T ipho a” mon ei oi Rolán& Fe nández-Ga cés, 2015: 47, pl.3.
T ipho a po o icensis non Rolán& Red e n, 2008 – Lamy& Poin ie
2018: 289, pl.92, ig.11.
ype ma e ial. — Holo ype. Guadeloupe • sh; Basse-Te e, Baie
de Baille-A gen ; KARUBENTHOS 1 s a. GD21, 16°15’33”N,
61°48’48”W; dep h 40m; MNHN-IM-2000-30473.
ype locali y.— Guadeloupe: Basse-Te e, Baie de Baille-A gen ;
KARUBENTHOS 1 s a. GD21, 16°15’33”N, 61°48’48”W; 40m.
ma e ial examined.— Guadeloupe. KARUBENTHOS 1 • 1sh
[p e iously iden i ied as Ma shallo a modes a – his shell is om he
ype locali y]; s a. GD21; MNHN.
Ma inique. MADIBENTHOS. ‘No d A lan ique’ • 1sh; s a.
AD263; MNHN • 1sh; s a. AD275; MNHN • 1sh; s a. AB350;
MNHN. — ‘Sud A lan ique’ • 1sh; s a. AS252; MNHN. — ‘No d
Ca aibe’ • 10sh, 1spm; s a. AD283; MNHN • 1sh; s a. AD290;
MNHN • 1sh; s a. AS482; MNHN.
ba hyme ic diS ibu ion.— Reco ded dep h in Guadeloupe:
40m (including a li e specimen). Reco ded dep h in Ma inique:
10-80m (li e specimen a 37-40m).
geog aphic diS ibu ion.— Guadeloupe and Ma inique.
emended deSc ip ion
Shell sinis al, conical o sligh ly pupoid, ec ilinea o sligh ly
con ex p o ile, up o 4.6mm long, 1.2mm wide, leng h/wid h

471
T ipho idae om Ma inique
ZOOSYSTEMA • 2024 • 46 (18)
a io 2.4-3.9; adul shells each a leas 2.0mm in leng h. P o-
oconch sub-columna , sligh ly in la ed, o 2.75-3.0 who ls,
0.44-0.51mm long, 0.41-0.48mm wide, weak dis inc ion
be ween p o oconch and eleoconch; i s smoo h, globose,
bu on subsequen who ls axial ibs soon appea ing, wi h hei
adapical pa eme ging on he abapical pa o he smoo h
who l abo e, assuming an i egula shape, o en conside ably
sigmoid on second who l, bu usually sligh ly sigmoid o
nea ly o hocline (some imes opis hocline) owa ds he end
o p o oconch, whe e i numbe s 20-22 ibs; one main spi al
co d eme ging soon a e he axial ibs, keel-shaped and o en
smoo h (bu w inkled in some shells), si ua ed a 47-60% o
las who l heigh . Teleoconch wi h up o nine who ls; wo
spi al co ds (adapical and abapical) a he beginning, he
abapical one con inuous o ha o p o oconch; median spi al
co d eme ging a he beginning o second o he end o hi d
who l, eaching he same size han o he co ds a e one o
wo who ls; a he body who l, he h ee co ds a e equidis-
an and ha e nea ly he same size; 22-24 nea ly o hocline
axial ibs on he six h who l; ounded nodules o medium
size; dis inc su u e, wi h a su u al co d; sligh ly nodulose
o smoo h subpe iphe al co d, wo smoo h, well-de eloped
basal co ds; no sup anume ical co ds; ape u e o a e (wi h
a minu e no ch in he pos e io end), 0.50-0.81mm long,
0.40-0.61mm wide, leng h/wid h a io 1.3-1.5; an e io
canal sho , open, 0.13-0.25mm long, 0.14-0.28mm wide,
leng h/wid h a io 0.8-0.9. P o oconch b ownish, c eam,
golden o whi e; eleoconch da k o ligh b own, a ely beige;
head- oo whi e, anslucen .
ema kS
This species was p e iously known only om he holo ype,
which has a ligh b own eleoconch and whi e p o oconch
(Rolán& Fe nández-Ga cés 2015). An addi ional shell
om he ype locali y in Guadeloupe is illus a ed (Fig.7J),
wi h a da ke eleoconch, which is he common pa e n in
shells o Ma shallo a mon ei oi n. comb. om Ma inique
(Fig.7). An excep ion is a shell wi h a g adual (no ab up )
colo ansi ion be ween he ligh e p o oconch and he
c eam/beige eleoconch (Fig.7K), which is mo e simila
o he plank o ophic species Ma shallo a os en a Rolán&
Fe nández-Ga cés, 2008, and i pe haps does no belong
o M. mon ei oi n. comb. The colo o he p o oconch is
a iable among shells o M. mon ei oi n. comb., om whi e
o b ownish (Fig.7). The e a e also sligh a ia ions in he
p o oconch sculp u e, wi h a spi al co d usually smoo h and
si ua ed a he mid po ion o he who l, bu some imes
w inkled o si ua ed a a lowe po ion, in addi ion o he
a iable axial ibs (Fig.8). Rema kably, he adul shell leng h
anges om 2.0 o 4.6mm.
AB
CD
EF
GH
Fig. 8. — Ma shallo a mon ei oi (Rolán & Fe nández-Ga cés, 2015) n. comb.: A, MNHN, s a. AD283 (Fig. 7A); B, MNHN, s a. AS482 (Fig. 7B); C, MNHN, s a. B350
(Fig. 7C); D, MNHN, s a. AD275 (Fig. 7D); E, MNHN, s a. AD283 (Fig. 7E); F, MNHN, s a. AD283 (Fig. 7F); H, MNHN, s a. AS252 (Fig. 7I); H, MNHN, s a. AD263
(Fig. 7K). Scale ba s: 100 μm.
472 ZOOSYSTEMA • 2024 • 46 (18)
Fe nandes M.R.
This species is placed unde he A lan ic genus Ma shal-
lo a owing o he b own eleoconch, smoo h basal co ds
(Fig.7L-M), absence o sup anume ical co ds (Fig.7N) and
whi e head- oo (Fig.20I-J), among o he ea u es. The e
a e o he Ca ibbean species wi h paucispi al p o oconchs
cu en ly alloca ed unde he ca ch-all axon “T ipho a” ha ,
o he wise, i in o he concep o Ma shallo a, ollowing he
abo e-men ioned ea u es [including he whi e head- oo o
one species – Red e n (2013: ig.356B)]. These species a e
he ein ans e ed o Ma shallo a, pending u u e ana omi-
cal and gene ic s udies o es hese alloca ions: Ma shallo a
abacoensis (Rolán& Lee, 2008) n. comb., Ma shallo a a man-
doi (Espinosa& O ea, 2020) n. comb., Ma shallo a cal a
(Fabe & Moolenbeek, 1991) [al eady ci ed in his genus by
Fe nandes& Pimen a 2020] and Ma shallo a g enadensis (Ro-
lán& Lee, 2008) n.comb. The la e , only known om he
holo ype, is he mos simila species o M. mon ei oi n.comb.,
ega ding sligh di e ences in he colo and sculp u e o he
p o oconch (Rolán& Fe nández-Ga cés 2015), and possibly
ha ing a less in la ed p o oconch and sligh ly la e eme gence
o he median spi al co d (eme ging na owly a he end o
ou h eleoconch who l – Rolán& Fe nández-Ga cés 2008:
ig.30D), equi ing addi ional ma e ial o u he compa isons.
Genus Me axia Mon e osa o, 1884
Me axia discus n.sp.
(Fig.9A-F)
u n:lsid:zoobank.o g:ac :B613849C-897F-4E0A-BE99-EBD074B89222
ype ma e ial. — Holo ype. Ma inique. • sh; No h o P esqu’Île
de la Ca a elle, egion ‘No d A lan ique’; MADIBENTHOS
expedi ion; s a. AD245; 14°53’06”N, 60°3’24”W; dep h 60m;
MNHN-IM-2000-38601.
Pa a ype. Ma inique. MADIBENTHOS • 1sh; ype locali y;
MNHN-IM-2000-38602.
ype locali y.— Ma inique: No h o P esqu’Île de la Ca a elle,
egion ‘No d A lan ique’; MADIBENTHOS expedi ion s a. AD245,
14°53’06”N, 60°3’24”W; 60m.
e ymology.— F om La in, discus. The species is named due o
he discoid shape o he p o oconch.
diagnoSiS.— P o oconch wi h 2.5much con ex who ls, i s
who l discoid, b oad, wi h same wid h han second one, and wi h
wo spi al co ds o a ying exp ession; shell whi e.
geog aphic diS ibu ion.— Ma inique ( his s udy).
ba hyme ic diS ibu ion.— Only known om 60m.
deSc ip ion
Shell dex al, conical/ usi o m, ec ilinea p o ile, up o 3.2mm
long, 0.8mm wide, leng h/wid h a io 3.5-4.1. P o oconch
paucispi al, o 2.5much con ex who ls, 0.41-0.44mm long,
0.34-0.37mm wide; i s who l discoid, b oad, wi h same wid h
han second one, wi h wo spi al co ds on he adapical hal o he
who l, ha ing a ying exp ession, he abapical co d mo e p omi
-
nen and ini ially undula ing (zigzag) o u he nea ly s aigh ,
and he adapical co d undula ing (zigzag); on he second who l,
he wo spi al co ds ha ing nea ly he same wid h and being no
longe es ic ed o he adapical po ion o he who l, si ua ed
espec i ely a c.39% and c.71% o who l heigh , wi h small
nodules, wi h incomple e, o hocline o sligh ly p osocline axial
ible s loca ed abo e adapical co d. Teleoconch wi h up o se en
who ls; ou equidis an spi al co ds, wi h he i s (adapical)
co d being much smalle and he hi d co d he mos p ominen ,
in addi ion o a much educed, smoo h su u al co d; ellip ical
nodules o small size; c.15 weak, o hocline axial ibs on he
six h eleoconch who l; smoo h and na ow subpe iphe al co d,
no dis inc basal co d; ape u e ounded (bu pa ially b oken),
wi h small an e io no ch. Shell whi e.
ema kS
Me axia discus n.sp. supe icially esembles he mo ph named
Me axiasp. 1 (Fig.9G-H), om Ba bados (Rolán& Fe nán-
dez-Ga cés 2008). The eleoconch o Me axiasp. 1has h ee
main spi al co ds, simila ly o Me axia excelsa (Fabe & Mo-
lenbeek, 1991) o Me axia aeniola a (Dall, 1889), whe eas
M. discus n.sp. has ou main spi als in la e who ls (i.e., he
uppe co d is s onge han ha o Me axiasp. 1, which seems
a weak subsu u al co d), simila ly o Me axia ugulosa (C. B.
Adams, 1850). The p o oconch o Me axiasp. 1 is illus a ed
he e (Fig.9H), and, despi e sligh ly wo n, is mo e simila o
M. ugulosa han o M. discus n.sp. (see compa isons below).
Despi e ha ing a nea ly iden ical eleoconch, M. discus
n.sp. is di e en ia ed om he widesp ead Wes A lan ic
species M. ugulosa by he discoid and unca ed shape o
he p o oconch, o en wi h undula ing spi al co ds (Fig.9D),
whe eas he p o oconch o M. ugulosa is always con ex, wi h a
dome-shaped i s who l and s aigh spi al co ds in he second
who l (Rolán& Red e n 1996; Fe nandes& Pimen a 2011).
Me axia discus n.sp. has a p o oconch simila o Me axia
quad a a Fabe , 2010, which is known only by he holo ype,
om A uba. Howe e , he p o oconch o M. quad a a seems
o ha e h ee spi al co ds on he i s who l ( s wo co ds in
M. discus n.sp.), wi h a mo e accen ua ed zigzag pa e n han
in M. discus n.sp., al hough he second who l o p o oconch
has a nea ly ec ilinea adapical spi al co d (bu mo e undula -
ing in M. discus n.sp.). In addi ion, he shell o M. quad a a
is yellowish b own and i has a weake axial sculp u e (Fabe
2010) s en i ely whi e in M. discus n.sp.
Me axia excelsa Fabe & Moolenbeek, 1991
(Figs2C; 18J, L-O)
Me axia excelsa Fabe & Moolenbeek, 1991: 83.
ma e ial examined.— Guadeloupe. KARUBENTHOS 1 •
2sh; s a. GB02; MNHN • 1spm; s a. GB03; MNHN • 3sh; s a.
GB06; MNHN • 1sh, 1spm; s a. GS06; MNHN • 2sh, 1spm;
s a. GB08; MNHN • 1sh; s a. GB11; MNHN • 1sh; s a. GS11;
MNHN • 1spm; s a. GB12; MNHN • 1sh; s a. GB13; MNHN
• 15sh, 3spm; s a. GS13; MNHN • 1sh; s a. GB14; MNHN
• 1spm; s a. GS15; MNHN • 4sh; s a. GB16; MNHN • 1sh;
s a. GS17; MNHN • 4sh; s a. GS18; MNHN • 1sh; s a. GS19;
MNHN • 1sh; s a. GS22; MNHN • 1sh; s a. GB23; MNHN •
473
T ipho idae om Ma inique
ZOOSYSTEMA • 2024 • 46 (18)
1sh; s a. GS23; MNHN • 1sh; s a. GB25; MNHN • 1sh; s a.
GB36; MNHN • 1sh; s a. GD68; MNHN. — KARUBENTHOS
2 • 1sh; s a. DW4545; MNHN.
Ma inique. MADIBENTHOS. ‘No d A lan ique’ • 1spm; s a.
AB197; MNHN • 1sh [wo n]; s a. AD255; MNHN • 2sh; s a.
AD260; MNHN • 2sh; s a. AD263; MNHN • 3sh; s a. AD267;
MNHN • 5sh; s a. AD275; MNHN • 2sh; s a. AS565; MNHN.
— ‘Sud A lan ique’ • 1sh; s a. AD222; MNHN • 1sh; s a. AB401;
MNHN. — ‘No d Ca aibe’ • 1sh; s a. AB159; MNHN • 3sh,
1spm; s a. AB161; MNHN • 2sh; s a. AD280; MNHN • 1sh; s a.
AD289; MNHN • 3sh; s a. AD290; MNHN • 1spm; s a. AB362;
MNHN • 1sh; s a. AS363; MNHN • 3sh; s a. AB372; MNHN •
1sh; s a. AS375; MNHN • 1sh; s a. AB382; MNHN • 4sh; s a.
AB386; MNHN • 1sh; s a. AB463; MNHN • 1sh; s a. AB567;
MNHN • 1sh; s a. AS572; MNHN • 3sh; s a. AS576; MNHN. ‘Sud
Ca aibe’ • 1sh; s a. AB058; MNHN • 1spm; s a. AB123; MNHN
AB CDE
F
GH
Fig. 9
. — A-F, Me axia discus n. sp. A, B, D, F, MNHN-IM-2000-38601, holo ype, 3.2 mm; C, E, MNHN-IM-2000-38602, pa a ype, s a. AD245, 2.7 mm; G, H, Me axia
sp. 1 (mo ph named by Rolán & Fe nández-Ga cés [2008]), USNM 87301, 4.2 mm ( s 5.0 mm, acco ding o Rolán & Fe nández-Ga cés [2008]); c edi s: Yolanda
Villacampa (NMNH). Scale ba s: A-C, G, 1 mm; D, E, 100 μm; F, H, 200 μm.
474 ZOOSYSTEMA • 2024 • 46 (18)
Fe nandes M.R.
• 2sh; s a. AB150; MNHN • 2sh; s a. AB152; MNHN • 2sh; s a.
AB169; MNHN • 1sh; s a. AB173; MNHN • 2sh; s a. AB177;
MNHN • 1sh; s a. AD203; MNHN • 2sh; s a. AD216; MNHN •
1sh [ju enile]; s a. AD271; MNHN • 1sh; s a. AB354; MNHN •
1sh; s a. AS370; MNHN • 1sh; s a. AD617; MNHN. ‘Baie de Fo -
de-F ance’ • 1sh; s a. AB392; MNHN • 1sh; s a. AB394; MNHN.
ba hyme ic diS ibu ion.— Reco ded dep h in Guadeloupe:
3-60m (li e specimens a 14-50m). Reco ded dep h in Ma inique:
2-80m (li e specimens a 4-25m). P e ious eco ded dep h in he
Wes A lan ic: 1-147m (Fe nandes& Pimen a 2020).
Me axia ugulosa (C. B. Adams, 1850)
(Fig.2D)
Ce i hium ugulosum C. B. Adams, 1850: 121-122.
ma e ial examined.— Guadeloupe. KARUBENTHOS 1 • 1sh;
s a. GS13; MNHN • 1sh; s a. GB14; MNHN • 1sh; s a. GB30;
MNHN. — KARUBENTHOS 2 • 1sh; s a. DW4586; MNHN.
Ma inique. MADIBENTHOS. ‘No d A lan ique’ • 1sh; s a.
AD244; MNHN • 1sh; s a. AD260; MNHN • 2sh; s a. AD261;
MNHN • 2sh; s a. AD263; MNHN • 1sh; s a. AD265; MNHN •
4sh; s a. AD275; MNHN. — ‘Sud A lan ique’ • 1sh; s a. AB120;
MNHN • 1sh; s a. AB185; MNHN. — ‘No d Ca aibe’ • 1sh; s a.
AD277; MNHN • 1sh; s a. AD280; MNHN • 1sh; s a. AS363;
MNHN • 1sh; s a. AS576; MNHN. ‘Sud Ca aibe’ • 1sh; s a.
AD067; MNHN • 1sh; s a. AB150; MNHN • 1sh; s a. AD218;
MNHN. ‘Baie de Fo -de-F ance’ • 1sh; s a. AB398; MNHN.
ba hyme ic diS ibu ion.— Reco ded dep h in Guadeloupe:
16-50m; one in ac adul shell ob ained by KARUBENTHOS 2
be ween 204-251m, bu i mus be ega ded wi h cau ion. Reco ded
dep h in Ma inique: 2-80m. P e ious eco ded dep h in he Wes
A lan ic: 0-198m (Fe nandes& Pimen a 2020).
ema kS
The shell om Guadeloupe illus a ed by Rolán& Fe nández-
Ga cés (2015: ig.4G) and Lamy& Poin ie (2018: pl.92,
ig.3) as M. ugulosa ac ually e e s o M. aeniola a (Dall,
1889).
Me axia aeniola a (Dall, 1889)
(Figs2E; 19A, B)
Ce i hiopsis me axae a . aeniola a Dall, 1889: 256.
ma e ial examined.— Guadeloupe. KARUBENTHOS 1 • 1sh
[p e iously iden i ied as M. ugulosa]; s a. GB08; MNHN • 1sh
[idem]; s a. GD10; MNHN • 1sh [idem]; s a. GD31; MNHN.
Ma inique. MADIBENTHOS. ‘No d A lan ique’ • 1spm; s a.
AD261; MNHN • 1sh; s a. AD275; MNHN • 1sh; s a. AS552;
MNHN. — ‘No d Ca aibe’ • 1sh; s a. AS572; MNHN.
ba hyme ic diS ibu ion.— Reco ded dep h in Guadeloupe:
17-85m. Reco ded dep h in Ma inique: 18-80m (li e specimen
a 80m). P e ious eco ded dep h in he Wes A lan ic: 4-101m
(Fe nandes& Pimen a 2020).
Me axiasp.
(Fig.10)
m
a e ial
examined
.— Ma inique. MADIBENTHOS. ‘Sud
A lan ique’ • 1sh; s a. AB181; MNHN.
geog aphic diS ibu ion.— Only known om Ma inique.
ba hyme ic diS ibu ion.— Only known om 22m.
AB
CD
Fig. 10. — Me axia sp. MNHN, s a. AB181, 2.6 mm. Scale ba s: A, B, 1 mm; C, 200 μm; D, 100 μm.
475
T ipho idae om Ma inique
ZOOSYSTEMA • 2024 • 46 (18)
A
B
CD E
FG
HI
Fig. 11
. — A-E, Nanapho a decolla a (Rolán & Fe nández-Ga cés, 1994), MNHN, s a. GB09 (KARUBENTHOS 1), 2.9 mm; a weak mic o-sculp u e on he eleoconch
is shown in B; F-I, Nanapho a c . e be nei (Moolenbeek & Fabe , 1989); F, G, MNHN, s a. AB130, bo h 3.0 mm; H, I, MNHN, s a. AB149, 2.8 mm, 3.1 mm. Scale
ba s: A, F-I, 1 mm; B, E, 100 μm; C, D, 200 μm.

476 ZOOSYSTEMA • 2024 • 46 (18)
Fe nandes M.R.
ema kS
The single a ailable shell o Me axiasp. has a ain axial sculp-
u e and a p o oconch wi h 2.5 who ls (measu ing 0.32mm
long, 0.31mm wide), and i s emb yonic shell has a dis inc ,
abapical zigzag spi al co d and a smalle , adapical spi al co d
bo de ing he adapical po ion o he who l (Fig.10D). I
could be an a ypical shell o M. ugulosa, ega ding he wide
a ia ion in he emb yonic shell o his species (Rolán&
Red e n 1996), al hough Ca ibbean shells o M. ugulosa
a e usually whi e and ha e a coa se axial sculp u e (Rolán&
Fe nández-Ga cés 2007, 2008; Lee 2009). Mos B azilian
shells o M. ugulosa a e b own (Fe nandes& Pimen a 2011,
2020), simila ly o Me axiasp. (Fig.10A, B), which demands
u he s udies – desi ably wi h gene ic e idence. Me axia
p opinqua Rolán& Fe nández-Ga cés, 2008, om sou he n
USA, has shells en i ely whi e o wi h a whi e p o oconch and
ligh b own eleoconch. The whi e pa a ype o M. p opinqua
illus a ed by Rolán& Fe nández-Ga cés (2008: ig.1E-G)
seems a ypical shell o M. ugulosa, con a y o he bicolo
holo ype.
Ano he ema kably simila species o Me axiasp. is Me axia
quad a a Fabe , 2010, desc ibed om A uba, based on a
single (ju enile) shell. This species was desc ibed as yellow-
ish b own, and he axial sculp u e o he eleoconch is much
ain , as obse ed in Me axiasp. (Fabe 2010). The appa en
single di e ence be ween hem is ha M. quad a a has h ee
spi al co ds ( s wo in Me axiasp.) on he i s p o oconch
who l. Because Me axia species a e mainly di e en ia ed by
sub le di e ences in p o oconch mo phology, mo e ma e ial
om he Lesse An illes is equi ed o p ope ly e alua e he
axonomy o his mo ph om Ma inique.
Genus Monopho us G illo, 1877
Monopho us oli aceus (Dall, 1889)
(Figs2F; 19C-F)
T i o is deco a a a . oli acea Dall, 1889: 244.
ma e ial examined.— Guadeloupe. KARUBENTHOS 1 •
1sh, 1spm; s a. GB03; MNHN • 1sh; s a. GS06; MNHN • 1sh;
s a. GS07; MNHN • 3sh; s a. GS08; MNHN • 3sh; s a. GD10;
MNHN • 2sh; s a. GB11; MNHN • 1sh; s a. GB12; MNHN
• 15sh, 1spm; s a. GS13; MNHN • 5sh; s a. GB14; MNHN •
2sh; s a. GD15; MNHN • 1sh; s a. GS16; MNHN • 2sh; s a.
GS18; MNHN • 2sh [ju eniles]; s a. GB19; MNHN • 1spm;
s a. GD21; MNHN • 1sh; s a. GS21; MNHN • 3sh; s a. GB22;
MNHN • 1sh; s a. GS22; MNHN • 1sh; s a. GB24; MNHN •
1sh; s a. GB25; MNHN • 2sh; s a. GS26; MNHN • 1sh; s a.
GS29; MNHN • 2sh [ju eniles]; s a. GR34; MNHN • 1sh; s a.
GB36; MNHN • 1sh; s a. GD61; MNHN.
Ma inique.
MADIBENTHOS.
‘No d A lan ique’ • 1sh; s a.
AR143; MNHN • 1sh; s a. AB197; MNHN • 1sh; s a. AB199;
MNHN • 1sh; s a. AD254; MNHN • 1sh [ju enile]; s a. AD260;
MNHN • 1sh; s a. AD263; MNHN • 6sh; s a. AB556; MNHN
• 8sh; s a. AS557; MNHN • 3sh; s a. AB559; MNHN • 1sh; s a.
AB562; MNHN • 7sh; s a. AS565; MNHN. — ‘Sud A lan ique’
• 1sh; s a. AD084; MNHN • 1spm; s a. AB185; MNHN • 1sh;
s a. AS255; MNHN • 3sh [ju eniles]; s a. AB405; MNHN. —
‘No d Ca aibe’ • 5sh; s a. AB108; MNHN • 8sh; s a. AB159;
MNHN • 2sh; s a. AB161; MNHN • 1sh, 1spm; s a. AD283;
MNHN • 1sh; s a. AD294; MNHN • 4sh; s a. AR308; MNHN
• 1sh; s a. AB362; MNHN • 1sh; s a. AS365; MNHN • 1sh; s a.
AB372; MNHN • 2sh; s a. AS373; MNHN • 10sh; s a. AS375;
MNHN • 7sh; s a. AB377; MNHN • 2sh; s a. AS378; MNHN
• 2sh; s a. AB386; MNHN • 1sh; s a. AB388; MNHN • 1spm;
s a. AR461; MNHN • 1sh [ju enile]; s a. AB510; MNHN •
5sh; s a. AS570; MNHN • 2sh; s a. AS572; MNHN • 1sh;
s a. AS574; MNHN • 5sh; s a. AS576; MNHN. ‘Sud Ca aibe’
• 1sh; s a. AS066; MNHN • 1sh; s a. AS071; MNHN • 1sh;
AR100; MNHN • 1sh; s a. AR122; MNHN • 1sh; s a. AB126;
MNHN • 6sh; s a. AB150; MNHN • 1spm; s a. AB152; MNHN
• 1sh; s a. AB155; MNHN • 3sh; s a. AB177; MNHN • 1sh;
s a. AD271; MNHN. ‘Baie de Fo -de-F ance’ • 1sh; s a. AB394;
MNHN • 2sh; s a. AS581; MNHN.
ba hyme ic diS ibu ion.— Reco ded dep h in Guadeloupe:
9-80m (li e specimens a 22-50m). Reco ded dep h in Ma inique:
2-80m (li e specimens a 4-40m). P e ious eco ded dep h in he
Wes A lan ic: 0-120m, wi h disc epan eco ds down o 260m
(Fe nandes& Pimen a 2020).
Genus Nanapho a Lase on, 1958
Nanapho a decolla a
(Rolán& Fe nández-Ga cés, 1994)
(Fig.11A-E)
Chei odon a decolla a Rolán& Fe nández-Ga cés, 1994: 20, igs19-
21, 23-24, 30CD; 2007: 20, pl.1, igs6-11.— Zhang 2011: 99,
ig.289.— Red e n 2013: 127, ig.361.— Lamy& Poin ie 2018:
284, pl.91, ig.2.
Nanapho a decolla a – Fe nandes& Pimen a 2015: 502.— Rolán&
Fe nández-Ga cés 2015: 53, ig.4V.
ype ma e ial.— Holo ype. Cuba • sh; MNCN 15.05/11142.
Pa a ypes. See he o iginal desc ip ion.
ype locali y.— Cuba: Ma ianao beach, La Habana.
ma e ial examined.— Guadeloupe. KARUBENTHOS 1 • 1sh;
s a. GB09; MNHN.
geog aphic diS ibu ion.— Bahamas (Red e n 2013); Cuba
(Rolán& Fe nández-Ga cés 1994); Cayman ( his s udy); An igua
(Zhang 2011); Guadeloupe (Rolán& Fe nández-Ga cés 2015;
Lamy& Poin ie 2018; his s udy).
ba hyme ic diS ibu ion.— Reco ded dep h in Guadeloupe:
6m. P e ious eco ded dep h in he Wes A lan ic: 2m (Rolán&
Fe nández-Ga cés 1994) o 14m (Red e n 2013).
ema kS
Nanapho a decolla a was o iginally desc ibed in Chei odon a
Ma shall, 1983, a genus wi h only wo con i med species,
i.e., he ype species Chei odon a pallescens (Je eys, 1867)
om he Eas A lan ic and Chei odon a dupliniana (Olsson,
1916) om he Wes A lan ic, plus a hi d species [Chei-
odon a labia a (A. Adams, 1854), om Aus alia] wi h a
en a i e alloca ion (Ma shall 1983; Fe nandes& Pimen a
2015). The A lan ic species o Chei odon a ha e adula
wi h mul icuspid ee h, o which he ma ginal a e much
elonga ed (Bouche & Guillemo 1978; Bouche 1985;
Fe nandes& Pimen a 2019b). In ac , he e a e signi ican
477
T ipho idae om Ma inique
ZOOSYSTEMA • 2024 • 46 (18)
simila i ies be ween he adula o N. decolla a (Rolán&
Fe nández-Ga cés 1994) and hose o Chei odon a, mainly
in he mo phology o he la e al ee h, wi h he cen al and
ma ginal ee h o N. decolla a being also simila o hose o C.
dupliniana, bu sligh ly di e en om hose o C. pallescens,
which shows cen al oo h wi h a dias ema and ma ginal
ee h wi h cusps es ic ed o he dis al pa o he basal
pla e. The p o oconch o N. decolla a is also sligh ly simila
o ha o Chei odon a, bea ing g anules on he emb yonic
shell and wo spi al co ds on he la al shell (Fig.11E), al-
hough he o me p o oconch is sho e and i s who ls a e
mo e con ex. The mos disc epan di e ences be ween N.
decolla a and Chei odon a ely on eleoconch ea u es, wi h
he o me showing a small and o oid shell shape, nodules
much la ge (including he subpe iphe al and basal co ds)
and a much educed su u e (Fig.11A). These condi ions
a e also obse ed in o he Wes A lan ic species, which,
pe haps p ema u ely, led Fe nandes& Pimen a (2015) o
include hem in Nanapho a, a p obably non-monophyle ic
genus which shows species wi h di e en pa e ns o em-
b yonic and la al shell sculp u es, and whose ype species
has a paucispi al p o oconch (hinde ing compa isons) and
unknown adula (Ma shall 1983; Fe nandes& Pimen a
2015). Ci ing Fe nandes& Pimen a (2015: 502), “ he a -
ini y among he gene a Nanapho a, Opimapho a Lase on,
1958 and Chei odon a makes necessa y a axonomic e ision
o hem o achie e a p ecise delimi a ion o each genus”,
including a u he molecula phylogeny.
Two unusual eleoconch ea u es o N. decolla a we e indica ed
by Rolán& Fe nández-Ga cés (1994) and Red e n (2013),
and a e con i med he e. The p esence o mic o-sculp u e
(Fig.11B) is no widesp ead in T ipho idae (Ma shall 1983;
Fe nandes& Pimen a 2020), bu some species cu en ly al-
loca ed in Nanapho a may show i (Ma shall 1983; see also
he nex species). Ano he ema kable ea u e o N. decolla a
is he eme gence o he median spi al co d no be ween he
adapical and abapical co ds, as obse ed in mos ipho ids,
bu de i ed om a spli o he adapical co d on he body
who l (Fig.11C), a e he adapical co d had become axially
elonga ed – i does no seem a sup anume ical co d (Bouche
1985; Fe nandes& Pimen a 2015). The spli ing o one spi al
co d in o wo co ds is obse ed in some species o Ce i hiop-
sidae H. Adams& A. Adams, 1853, and i was epo ed in a
T ipho idae species om Ascension Island (Bakke & Swin-
nen 2021), al hough wi h a di e en o ma ion and majo
di e gences in he shell.
Rosenbe g (2009) ci ed wo new locali ies o he Ca ibbean
ange o Nanapho a e be nei (Moolenbeek& Fabe , 1989),
i.e., Cayman and G enada, based on ma e ial om he ANSP
collec ion. A e checking images o he single lo s o such
locali ies (ANSP 200078, G and Cayman Is., 2m - 2.8mm
long, wi h p o oconch; ANSP 296542, P ickly Bay, G enada,
0-1m - 3.5mm long, wi hou p o oconch), he ma e ial om
Cayman ac ually e e s o N. decolla a, whe eas ha om G e-
nada is wo n and p ecludes u he iden i ica ion (al hough i
seems N. decolla a). The main di e ence be ween bo h species
is he spi al sculp u e o he p o oconch, since N. decolla a
has always wo spi al co ds (Fig.11E). Shells wi hou apex
may be much simila owing o he o oid shape and bi u ca -
ing adapical spi al co d o eleoconch ( o N. e be nei, see
Fig.13B and Moolenbeek& Fabe 1989: ig.8). Rolán&
Fe nández-Ga cés (1994) a gued ha he whi e band is only
seen in he adapical co d o N. decolla a, bu i can be also
p esen in he abapical co d in he penul ima e who l o N.
e be nei, which is seen in mos shells om Ma inique wi hou
apex (Fig.11F-I); e en hough, i is p e e ed o name hese
decolla e shells as N. c . e be nei, and so a N. decolla a is
absen om Ma inique.
AB
CD
Fig. 12
. — Types o Nanapho a e be nei (Moolenbeek & Fabe , 1989); c edi s o MSc. Hannco Bakke ; A, ZMA.MOLL.136613, holo ype; B-D, ZMA.MOLL.136655,
pa a ypes. Scale ba s: 1 mm.
478 ZOOSYSTEMA • 2024 • 46 (18)
Fe nandes M.R.
Nanapho a e be nei (Moolenbeek& Fabe , 1989)
(Figs12; 13)
T ipho a e be nei Moolenbeek& Fabe , 1989: 77, igs6-8.
T ipho a exiguum non C. B. Adams, 1850 – De Jong& Coomans
1988: 49.
Chei odon a e be nei – Rolán& Fe nández-Ga cés 1994: 20,
igs17-18, 22, 30 CV.
Cosmo ipho a e be nei – Rolán& Fe nández-Ga cés 2007: 20,
pl.1, igs17-18.
Nanapho a e be nei – Fe nandes& Pimen a 2015: 500 (new gene ic
alloca ion, bu he species was misiden i ied – see below).
ype ma e ial.— Holo ype. Cu açao • sh; ZMA.MOLL.136613.
Pa a ypes. Cu açao • 19sh; ype locali y; ZMA.MOLL.136655.
ype locali y.— Cu açao: Boca Labade a, San a Ca a ina, beach.
ma e ial examined.— Guadeloupe. KARUBENTHOS 1 • 1sh
[wo n – p e iously iden i ied as Nanapho a decolla a]; s a. GM06;
MNHN • 1sh [idem]; s a. GM11; MNHN • 1sh [ju enile –
p e iously iden i ied as Co iopho a no em]; s a. GS13; MNHN.
— KARUBENTHOS 2 • 1sh; s a. DW4545; MNHN • 1sh
[ju enile]; s a. DW4551; MNHN.
Ma inique. MADIBENTHOS. ‘No d A lan ique’ • 1sh; s a. AD261;
MNHN • 1sh; s a. AD275; MNHN. — ‘No d Ca aibe’ • 1sh;
s a. AD280; MNHN. ‘Sud Ca aibe’ • 1sh; s a. AD115; MNHN.
m
a e ial
examined
o
N
aNapho a
c
.
e be Nei
(Figs1F-I;1I).—
Ma inique. ‘Sud A lan ique’ • 4sh; s a. AB130; MNHN • 2sh,
1spm; s a. AB149; MNHN.
geog aphic diS ibu ion.— Cuba (Rolán& Fe nández-Ga cés
1994); Pue o Rico (Moolenbeek& Fabe 1989); Guadeloupe and
Ma inique ( his s udy); Bonai e and Cu açao (Moolenbeek&
Fabe 1989).
ba hyme ic diS ibu ion.— Reco ded dep h in Guadeloupe:
0-82m. Reco ded dep h in Ma inique: 11-80m. Reco ded dep h
o N. c . e be nei in Ma inique: 1-2m (li e specimen a 2m).
P e ious eco ded dep h in he Wes A lan ic: 0-90m (Moolen-
beek& Fabe 1989).
emended deSc ip ion
Shell sinis al, cy oconoid, con ex p o ile, up o 3.9mm long,
1.4mm wide, leng h/wid h a io 2.8-3.0. P o oconch mul-
ispi al, conical/columna , o 5.0-5.5 sligh ly con ex who ls,
0.54-0.57mm long, 0.42-0.44mm wide; emb yonic shell
dome-shaped, co e ed by ounded g anules; la al shell ini ially
wi h wo spi al h eads bu adapical one soon disappea ing,
wi h one spi al co d (abapical) emaining un il he eme gence
o adapical co d in he penul ima e who l, he wo co ds si u-
a ed a 32-36% and 61-68% o las who l heigh ; c.27 nea ly
ec ilinea o sligh ly sigmoid axial ibs. Teleoconch wi h up
o nine who ls; wo spi al co ds (adapical and abapical) a he
beginning, abapical one con inuous o ha o p o oconch;
median spi al co d eme ging na owly a six h o se en h
who l, bo de ing close he adapical co d, eaching he same
size o abapical co d (adapical one is sligh ly mo e p ominen
on body who l) a e 1.5-2.0 who ls; ounded nodules o la ge
size; 17-19 opis hocline axial ibs on se en h who l; shallow
su u e, wi h small su u al co d; spi al mic o-sculp u e p esen
in all eleoconch; subpe iphe al co d nodulose, adapical basal
co d sligh ly nodulose o wa y, nea ly smoo h abapical basal
co d; sup anume ical co ds no disce nible (bu pe is ome
ne e in ac ); nea ly ci cula ape u e, 0.55-0.57mm long,
0.50-0.58mm wide, leng h/wid h a io 0.9-1.1; an e io canal
almos closed, c ossed in i s base by p ojec ion o ou e lip,
sligh ly cu ed backwa d, 0.34-0.50mm long, 0.22-0.26mm
wide, leng h/wid h a io 1.3-2.2; pos e io canal as a small hole,
de ached om ape u e. Ligh b own o o ange p o oconch;
eleoconch ligh b own o nea ly o ange, wi h adapical spi al
co d o en whi ish on body who l, and some shells also show
abapical co d ligh e han adapical co d on mid who ls.
ema kS
The au ho ini ially alloca ed his species in Co iopho a due o
he spi al sculp u e o he p o oconch (Fig.13H, K), combined
wi h a mic o-sculp u e on eleoconch (Fig.13G, J), shell shape,
opis hocline axial ibs, shallow su u e and la e eme gence o
median spi al co d o eleoconch (Fig.13A-E) – ega ding he e
a e subs an ial di e gences be ween he single species om he
Wes A lan ic (C. no em) and he ype species o Co iopho a
(Fe nandes& Pimen a 2020). Howe e , a e e iewing de ails
on he p o oconch sculp u e o Nanapho a e be nei in he
li e a u e om he Ca ibbean (Moolenbeek& Fabe 1989;
Rolán& Fe nández-Ga cés 1994), an e o was de ec ed.
Fe nandes& Pimen a (2015) in e p e ed he emb yonic shell
o he Ca ibbean species as e icula ed ( ollowing shells om
B azil), owing o he inadequa e SEM images p e iously a ail-
able, despi e being desc ibed wi h hemisphe ic ube cles in
he ma e ial om Cuba (Rolán& Fe nández-Ga cés 1994).
Fe nandes& Pimen a (2015) also conside ed ha N. e be -
nei had always wo spi al co ds in he la al shell, ollowing
shells om B azil and he dubious image o a shell om Cuba
(Rolán& Fe nández-Ga cés 1994: ig.22, which is b igh in
pa o he p o oconch, hampe ing a p ope isualiza ion), bu
hey equi ocally con es ed he desc ip ion o he la al shell
as ha ing ini ially one bu u he wo spi al co ds (Moolen-
beek& Fabe 1989: ig.7). In ac , he Ca ibbean N. e be nei
has ounded g anules on he emb yonic shell and he pa e n
2-1-2-(1) spi al co ds on he la al shell (Fig.13H, K). The
B azilian mo ph, despi e showing a much simila eleoconch
mo phology, is no N. e be nei, as indica ed by disc epan
ea u es on he adula and on he colo o li ing specimens
(Fe nandes& Pimen a 2019b), and i will be discussed unde
he ollowing species. Acco dingly, he eco d o N. e be nei
om An igua (Zhang 2011) is ela ed o he nex species, and
he eco ds om Cayman and G enada (Rosenbe g 2009) a e
in alid (see ema ks o N. decolla a).
The iden i ica ion o N. e be nei om Guadeloupe by Rolán&
Fe nández-Ga cés (2015), and ollowed by Lamy& Poin ie
(2018), is also inco ec (see nex species); he eal N. e be nei
was hidden unde he names o wo o he species (see ma e ial
examined). Because E. Rolán was one e iewe o he manu-
sc ip ha esul ed in Fe nandes& Pimen a (2015), nea ly a
he same ime o he publica ion o Rolán& Fe nández-Ga cés
(2015), i may ha e in luenced he w ong iden i ica ion o
shells om Guadeloupe. Compa ing he ma e ial om Gua-
479
T ipho idae om Ma inique
ZOOSYSTEMA • 2024 • 46 (18)
ABCDE
FG H
IJ K
Fig. 13. — Nanapho a e be nei (Moolenbeek & Fabe , 1989): A, MNHN, s a. GM11 (KARUBENTHOS 1), 3.8 mm; B, MNHN, s a. AD115, 3.9 mm; C, MNHN,
s a. AD261, 3.7 mm; D, F-I, MNHN, s a. AD280, 3.9 mm; E, J, K, MNHN, s a. DW4545 (KARUBENTHOS 2 expedi ion), 3.8 mm. A weak mic o-sculp u e on he
eleoconch is shown in G, J. Scale ba s: A-E, 1 mm; F, 500 μm; G-H, J-K, 100 μm; I, 200 μm.
486 ZOOSYSTEMA • 2024 • 46 (18)
Fe nandes M.R.
ma e ial examined.— Guadeloupe. KARUBENTHOS 1 • 1sh;
s a. GD35; MNHN.
Ma inique. MADIBENTHOS. ‘Sud Ca aibe’ • 1sh; s a. AD218;
MNHN.
ba hyme ic diS ibu ion.— Reco ded dep h in Guadeloupe:
66m. Reco ded dep h in Ma inique: 60m. P e ious eco ded
dep h in he Wes A lan ic: 58-163m (Fe nandes& Pimen a 2020).
ema k
This species was iden i ied as “T ipho a”sp. by Rolán&
Fe nández-Ga cés (2015: ig.4U) and as Iso ipho asp. 1 by
Lamy& Poin ie (2018: pl.91, ig.14).
Genus “T ipho a” Blain ille, 1828 s.l.
T ipho a ellyae De Jong& Coomans, 1988
(Figs3B; 20G-H)
T ipho a ellyae De Jong& Coomans, 1988: 50, pl.34, ig.242.
ma e ial examined.— Guadeloupe. KARUBENTHOS 1 • 2sh
[p e iously iden i ied as Similipho a in e media]; s a. GB05; MNHN
• 1sh, 1spm [idem]; s a. GB06; MNHN • 1sh [idem]; s a. GB11;
MNHN • 1sh [iden i ied as Monopho us a e albus by Rolán&
Fe nández-Ga cés (2015: ig.4X)]; s a. GD13; MNHN • 1sh [p e-
iously iden i ied as Similipho a in e media]; s a. GS13; MNHN.
Ma inique. MADIBENTHOS. ‘No d A lan ique’ • 1sh; s a. AD260;
MNHN • 1sh; s a. AD263; MNHN • 1sh; s a. AB350; MNHN
• 1sh; s a. AB556; MNHN • 2sh; s a. AS557; MNHN. — ‘No d
Ca aibe’ • 2sh; s a. AB159; MNHN • 1sh; s a. AD283; MNHN
• 1sh; s a. AD290; MNHN • 1sh; s a. AD299; MNHN • 1spm;
s a. AB308; MNHN • 1sh; s a. AS375; MNHN • 3sh; s a. AB463;
MNHN • 1sh; s a. AS570; MNHN • 2sh; s a. AS572; MNHN. ‘Sud
Ca aibe’ • 1sh; s a. AB062; MNHN • 1sh; s a. AB169; MNHN.
ba hyme ic diS ibu ion.— Reco ded dep h in Guadeloupe:
5-50m (li e specimen a 23m). Reco ded dep h in Ma inique:
5-80m. P e ious eco ded dep h in he Wes A lan ic: 3-110m
(Fe nandes& Pimen a 2020).
T ipho a ma ii Rolán& Fe nández-Ga cés, 1995
(Fig.3C)
“T ipho a” ma ii Rolán& Fe nández-Ga cés, 1995: 16, igs39-42.
ma e ial examined.— Guadeloupe. KARUBENTHOS 1 • 1sh
[iden i ied as Eu ipho a be mudensis by Rolán& Fe nández-Ga cés
(2015: ig.4T)]; s a. GS18; MNHN • 1sh [p e iously iden i ied
as Ini o is u is homae]; s a. GD59; MNHN.
Ma inique. MADIBENTHOS. ‘Sud A lan ique’ • 1sh; s a.
AS252; MNHN.
ba hyme ic diS ibu ion.— Reco ded dep h in Guadeloupe:
49-88m. Reco ded dep h in Ma inique: 10-12m. P e ious eco ded
dep h in he Wes A lan ic: 20-40m (Rolán& Fe nández-Ga cés 1995).
T ipho a po o icensis Rolán& Red e n, 2008
(Figs16; 17)
“T ipho a” po o icensis Rolán& Red e n in Rolán& Fe nández-Ga cés
2008: 158, ig.32A-E.
Ini o issp. – Red e n 2001: 66, pl.33, ig.278A, B.
“T ipho a” po o icensis – Red e n 2013: 127, ig.358A, B.—
Fe nandes& Pimen a 2015: 505, ig.6.— Fe nandes& Pimen a
2020: 168.
ype ma e ial.— Holo ype. Pue o Rico • sh; FLMNH 363895.
Pa a ype. Bahamas • 1sh; Abaco; 26°44’00”N, 77°09’00”W;
9.IX.1987; C. Red e n leg.; dep h 52m; BMSM 55395.
ype locali y.— Pue o Rico.
ma e ial examined.— Ma inique. MADIBENTHOS. ‘No d
A lan ique’ • 3sh; s a. AB350; MNHN. ‘Sud A lan ique’ • 1sh; s a.
AB183; MNHN •1sh; s a. AB191; MNHN • 1sh; s a. AB193;
MNHN • 1sh; s a. AS255; MNHN • 5sh; s a. AB405; MNHN.
geog aphic diS ibu ion.— Bahamas (Rolán& Fe nández-Ga cés
2008; Red e n 2013); Pue o Rico (Rolán& Fe nández-Ga cés
2008); Ma inique ( his s udy); B azil. Rio G ande do No e o
Rio de Janei o (Fe nandes& Pimen a 2015).
ba hyme ic diS ibu ion.— Reco ded dep h in Ma inique:
14-23m. P e ious eco ded dep h in he Wes A lan ic: 23-100m
(Fe nandes& Pimen a 2015, 2020).
emended deSc ip ion
Shell sinis al, conical o sligh ly pupoid, ec ilinea o sligh ly
con ex p o ile, eaching 3.6mm long, 1.3mm wide, leng h/
wid h a io 2.6-3.1; adul shells each a leas 1.9mm in
leng h. P o oconch sub-columna , sligh ly in la ed, o 2.75-
3.25 who ls, 0.40-0.58mm long, 0.41-0.51mm wide, weak
dis inc ion be ween p o oconch and eleoconch; ini ial 0.5
who l smoo h, bu axial ibs soon appea , wi h hei adapi-
cal pa eme ging on he abapical pa o he smoo h who l
abo e, assuming an i egula shape, o en a ow-shaped ini-
ially bu a ying om sigmoid, nea ly o hocline, s ongly
opis hocline o s ongly p osocline on he las who l, whe e
i numbe s 20-34 ibs; one main spi al co d o en occupying
he mid po ion o he who l (some imes he abapical po -
ion), si ua ed a 44-61% o las who l heigh , keel-shaped,
in addi ion o a small su u al co d. Teleoconch wi h up o
se en who ls; wo spi al co ds (adapical and abapical) a he
beginning, he abapical one con inuous o ha o p o oconch;
median spi al co d eme ging a he end o second who l o
a he hi d who l, eaching he same size han o he co ds
a e c.1.5 who l; a he body who l, he h ee co ds being
equidis an and nea ly he same size; 20-21 nea ly o hocline
axial ibs on he i h who l; ounded nodules o medium
size, wi h squa e in e spaces among nodules; dis inc su u e,
wi h a su u al co d; sligh ly nodulose o wa y subpe iphe al
co d, one o wo smoo h basal co ds, o en na ow, bu some-
imes well-de eloped; no dis inc sup anume ical co ds, bu
a minu e spi al h ead some imes appea ing be ween abapi-
cal and subpe iphe al co ds p io o he pe is ome; ape u e
ounded o sligh ly homboid (wi h a disc e e no ch in he
pos e io end), 0.47-0.83mm long, 0.43-0.59mm wide,
leng h/wid h a io 1.1-1.4; an e io canal e y sho , open,
0.07-0.15mm long, 0.13-0.20mm wide, leng h/wid h a io
0.4-0.7. Whi e o ligh c eam shell, wi h adapical spi al co d
and base sligh ly da ke .

487
T ipho idae om Ma inique
ZOOSYSTEMA • 2024 • 46 (18)
AB
C
D
M
EF
GH
I
K
L
O
N
J
Fig. 18
. — Li e ipho ids om Ma inique and Guadeloupe (ma ked wi h as e isk): A, B, Co iopho a no em (Nowell-Us icke, 1969), MNHN, s a. GM08 (A),
AD254 (B); C-F, Cosmo ipho a melanu a (C.B.Adams, 1850), MNHN, s a. GB03 (C), AS096 (D), GS17 (E), AD214 (F); G-I, K, Ini o is u is homae (Hol en, 1802),
MNHN, s a. AS075 (G-H), AM005 (I), GB07 (K); J, L-O, Me axia excelsa Fabe & Moolenbeek, 1991, MNHN, s a. AB123 (J), GB12 (L, O), AB362 (M), AB197 (N).
C edi s: MNHN eam.
488 ZOOSYSTEMA • 2024 • 46 (18)
Fe nandes M.R.
ema kS
The shell om Guadeloupe iden i ied by Lamy& Poin ie
(2018: pl.92, ig.11) as T ipho a po o icensis is ac ually he
holo ype o Ma shallo a mon ei oi n. comb. Some shells o
T. po o icensis om Ma inique (Fig.16) esemble one shell
en a i ely iden i ied as M. mon ei oi n. comb. (Fig.7K), bu
he o me mo ph has a sligh ly da ke adapical spi al co d o
eleoconch ( s spi al co ds wi h a mo e homogeneous colo
in M. mon ei oi n. comb.) and basal co ds (Fig.17A, E) o en
conside ably hinne han M. mon ei oi n. comb. (Fig.7L-M).
Shells o T. po o icensis om Ma inique a y in he adul
shell leng h (1.9-3.6mm) and in he axial sculp u e o he
p o oconch, bu he la ges shells a e simila o he holo ype
om Pue o Rico (4.4mm o eigh eleoconch who ls s up
o se en who ls in shells om Ma inique). In con as , shells
om Bahamas and B azil a e en i ely whi e (Red e n 2013;
Fe nandes& Pimen a 2015); hose om B azil ha e a di e -
en eleoconch sculp u e in la e who ls, wi h a mo e dis an
adapical spi al co d (no disce nible in Bahamian shells, only
known by ju eniles), and each up o 10mm o 13 eleoconch
who ls (Fe nandes& Pimen a 2015). This sugges s ha he
cu en concep o T. po o icensis is ac ually a species com-
plex masked by he simila p o oconch mo phology, which
could be de i ed om con e gence o pa allelism.
T ipho a c . scylla Fe nandes& Pimen a, 2015
(Fig.3D)
T ipho a scylla Fe nandes& Pimen a, 2015: 509, ig.8.
A
B
C
D
E
F
GHI
C
Fig. 19. — Li e ipho ids om Ma inique and Guadeloupe (ma ked wi h as e isk): A, B, Me axia aeniola a (Dall, 1889), MNHN, s a. AD261; C-F, Monopho us
oli aceus (Dall, 1889), MNHN, s a. GD21 (C), AB185 (D), AR461 (E, F); G, H, Nanapho a leei Fe nandes & Pimen a, 2015, MNHN, s a. AB400 (G), AB419 (H);
I, Nanapho a c . e be nei (Moolenbeek & Fabe , 1989), MNHN, s a. AB149. C edi s: MNHN eam.
489
T ipho idae om Ma inique
ZOOSYSTEMA • 2024 • 46 (18)
ma e ial examined.— Ma inique. MADIBENTHOS. ‘No d
A lan ique’ • 1sh [ju enile]; s a. AD275; MNHN • 1sh [ju enile];
s a. AB562; MNHN.
ba hyme ic diS ibu ion.— Reco ded dep h in Ma inique:
14-80m. P e ious eco ded dep h in he Wes A lan ic: 20-150m
(Fe nandes& Pimen a 2020).
DISCUSSION
A e mo e han 500 collec ion e en s and 33 ipho id
species sampled om he shallow wa e s o Ma inique,
he MADIBENTHOS expedi ion signi ican ly imp o ed
ou knowledge abou he di e si y o T ipho idae om he
Ca ibbean. Besides he desc ip ion o h ee new species
A
B
C
D
EFG
HI J
Fig. 20. — Li e ipho ids om Ma inique and Guadeloupe (ma ked wi h as e isk): A-D, No o ipho a deco a a (C.B. Adams, 1850), MNHN, s a. GR10 (A), AS253
(B), AB382 (C), AB161 (D); E, F, Sageno ipho a osclausum (Rolán & Fe nández-Ga cés, 1995), MNHN, s a. AB191; G, H, T ipho a ellyae De Jong & Coomans,
1988, MNHN, s a. AB308; I, J, Ma shallo a mon ei oi (Rolán & Fe nández-Ga cés, 2015) n. comb., MNHN, s a. AD283 (I), GD21 (J). C edi s: MNHN eam.
490 ZOOSYSTEMA • 2024 • 46 (18)
Fe nandes M.R.
(all non-plank o ophic), he e we e some new eco ds o
plank o ophic species o he Lesse An illes. Chei odon a
dupliniana and S obilige a inaudi a we e epo ed along
he No h and Sou h po ions o he Wes A lan ic, bu
nei he we e p e iously eco ded om any Ca ibbean island
(Fe nandes& Pimen a 2020). Despi e he abo emen ioned
No. species
No. shells
No. s a ions
No. species
Plank o ophic Plank o ophic
Non-plank o ophic Non-plank o ophic
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 o 2 1 o 2
3 o 5 3 o 5
6 o 20 6 o 2021 o 50 21 o 5051 o 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
Ini o is u is homae
Cosmo ipho a melanu a
Fig. 21
. — A, Indi idual a e ac ion cu e o ipho ids om MADIBENTHOS; blue cu es indica e he 95% con idence in e al; B, abundance dis ibu ion o
ipho ids om MADIBENTHOS; C, D, numbe o species s numbe o shells (C) o numbe o s a ions (D), ega ding plank o ophic (black ba s) and non-
plank o ophic (g ey ba s) species.
491
T ipho idae om Ma inique
ZOOSYSTEMA • 2024 • 46 (18)
issues o iden i ica ion, Nanapho a leei is now con i med o
he Ca ibbean, and no es ic ed o B azil; he same may
apply o T ipho a c . scylla, pending addi ional ma e ial.
T ipho a ma ii was p e iously known om Cuba (Rolán&
Fe nández-Ga cés 1995; Espinosa e al. 2007; Ga cía&
Capo e 2013). On he o he hand, no shell o Inella ise-
ialis was sampled by MADIBENTHOS, con i ming his
is a deep-sea species; he iden i ica ion om Ma inique
is unce ain (Lamy& Poin ie 2018), ega ding he poo
condi ion o ypes, om he deep sea o he Yuca án S ai
(Rolán& Fe nández-Ga cés 2008; Fe nandes& Pimen a
2019a).
The indi idual a e ac ion cu e o ipho ids om MADIB-
ENTHOS (Fig.21A) is nea ly sa u a ed bu did no each he
asymp o e, sugges ing mo e han 33 species exis in Ma i-
nique. Acco dingly, Chao1 es ima o sugges ed he occu ence
o 38 species, i.e., i e species om Ma inique we e possibly
missed (no sampled) by MADIBENTHOS. This numbe is
conside ably in e io han he o al ipho id species ound
in simila e o s by he MNHN eam in he Indo-Paci ic,
such as New Caledonia (174 species, 42 s a ions – Bouche
e al. 2002) and Vanua u (259 species, 566 s a ions – Albano
e al. 2011). Simila ly, Ma shall (1983) indica ed he exis -
ence o a leas 80 ipho id species in a single sand sample
No. species
AB
B ushing (AB): 68 s a ions
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%)
D edging (AD): 126 s a ions
Visual a in e idal (AM): 51 s a ions
Visual a di ing (AR): 204 s a ions Suc ion (AS): 42 s a ions
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, numbe o ipho id species ob ained in MADIBENTHOS by each sampling me hod (AB, b ushing; AD, d edging; AM, isual a in e idal; AR, isual
a di ing; AS, suc ion); B, numbe o species sampled s p opo ion o s a ions acco ding o each sampling me hod in MADIBENTHOS.

492 ZOOSYSTEMA • 2024 • 46 (18)
Fe nandes M.R.
om NE Aus alia. These con as ing numbe s e lec he
known highe di e si y o T ipho idae in he Indo-Paci ic
(Albano e al. 2011).
The abundance dis ibu ion o ipho ids (Fig.21B) shows
wo species wi h skewed dis ibu ions, i.e., Ini o is u is homae
and Cosmo ipho a melanu a. This is e lec ed in he low e en-
ness (0.276; 0.258-0.294), al hough he dominance was also
low (D: 0.209; 0.192-0.226), whe eas Shannon’s di e si y
(H: 2.208; 2.141-2.274) and Pielou’s equi abili y (J: 0.631;
0.612-0.650) we e mode a ely high. Se en species (all wi h
plank o ophic de elopmen ) we e conside ed abundan (51-
100shells) o much abundan (>100shells) and we e sampled
in mo e han 20 s a ions, whe eas eigh species (wi h plank o-
ophic o non-plank o ophic modes) we e much a e (i.e.,
ep esen ed by single ons o double ons) and sampled in one
o wo s a ions (Fig.21C, D). No all plank o ophic species
Wes A lan ic
(18 spp.; 54.5%)
NW A lan ic
(8 spp.; 24.2%)
Lesse An illes
(2 spp.; 6.1%)
Ma inique
(5 spp.; 15.2%)
No. species
To al Sha ed species wi h Ma inique
2922
Ca ibbean A lan ic
W
es
A
l
an
ic
(
18 spp.; 54.5%
)
NW
A
l
a
n
ic
(
8 s
pp
.; 24.2%
)
Lesse
A
n
il
les
(
2 spp.; 6.1%
)
Ma iniqu
e
(
5 s
pp
.; 15.2%
)
N
o. s
p
ec
i
es
To a
l
S
ha ed species wi h Ma iniqu
e
2
9
22
C
a ibbean
A
l
a
n i
c
No. species
26
30
25
15
5
30
30
0
No d A lan ique Sud A lan ique No d Ca aïbe Sud Ca aïbe Baie de Fo -de-
F ance
22
20
18
11
A
C
D
EF
B5960
50
40
30
20
10
0
B azil CubaMa inique 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, Geog aphic ange o ipho ids om Ma inique, whe he endemic s p esen in o he si es in Lesse An illes o beyond in he No hwes o Wes
A lan ic; B, o al shallow-wa e ipho ids eco ded in each well-sampled locali y, including he numbe o sha ed species wi h Ma inique; e e ences om B azil
a e hose o M. Fe nandes and colleagues, om Cuba hose o E. Rolán (o J. Espinosa) and colleagues, om A uba, Bonai e and Cu açao (ABC Is.) a e De Jong &
Coomans (1988), Moolenbeek & Fabe (1989), Fabe & Moolenbeek (1991) and Fabe (2010), and Red e n (2013) om Abaco, Bahamas; C, numbe o ipho id
species pe geog aphic zone; D, numbe o ipho id species pe Ca ibbean o A lan ic side in Ma inique; E, F, nMDS o geog aphic and ba hyme ic zones e-
ga ding ipho id species, based on B ay-Cu is (E) and Jacca d (F) indexes: da k g een, No d A lan ique; ligh g een, Sud A lan ique; da k blue, No d Ca aibe;
ligh blue, Sud Ca aibe; ed, Baie de Fo -de-F ance; do s, 0-10 m; squa es, 11-20 m; iangles, 21-30 m; X, 31-60 m; + - 61-85 m.
493
T ipho idae om Ma inique
ZOOSYSTEMA • 2024 • 46 (18)
we e abundan o p esen in many s a ions, because he e migh
be bio ic and abio ic es ic ions o la al se lemen and adul
su i o ship (such as he p esence o pa icula sponge hos s).
Simila highe abundances o plank o ophic ipho ids we e
also obse ed in a su ey om Vanua u, SW Paci ic (Albano
e al. 2011), sugges ing ha non-plank o ophic ipho ids a e
demog aphically a e owing o ce ain biological (e.g. sho o
e en null la al dispe sion) o ecological es ic ions. Two hi ds
(22 o 33) o he ipho id species om MADIBENTHOS
a e plank o ophic, a p opo ion highe han he nea ly 50%
ound o ipho ids om B azil (Fe nandes& Pimen a 2020)
and sou he n Aus alia (Ma shall 1983). Howe e , he s udy
o deep-sea ma e ial om Ma inique will ce ainly al e his
scena io by adding mo e non-plank o ophic species.
The e was a bias in he sampling me hods on MADIB-
ENTHOS, wi h 42 suc ion (AS) e en s, 51 isual e o s a
in e idal (AM), 68 o b ushing (AB), 126 o d edging (AD)
and 204 isual e o s a di ing (AR). No me hod could sam-
ple all 33 ipho id species due o he a i y o many o hem,
bu he isual me hods (AM and AR) we e clea ly poo a he
o al numbe o sampled species and on he equi ed a emp s
o ob ain a single specimen (Fig.22), due o he small size o
ipho ids (Albano e al. 2011). On he o he hand, b ushing
and suc ion we e once again ega ded as e icien me hods
o ob ain ipho ids (Albano e al. 2011), sampling simila
numbe s o species (22 s 23) and p opo ions o species (up
o nine o eigh ) pe s a ion (Fig.22). Despi e ha c.54%
o d edging s a ions did no eco e any ipho id, some
A lan ic side
N
Ca ibbean 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.
T ipho a po o icensis
Ini o is gudeliae
Ma shallo a c . modes a
A
B
C
DEF
T ipho a ellyae
Ini o is pseudo homae
Fig. 24. — A, hype -di e se s a ions o T ipho idae om MADIBENTHOS, con aining mo e han six species; B-F, maps o geog aphic dis ibu ion o selec ed
species. All maps ob ained om Google Ea h.
494 ZOOSYSTEMA • 2024 • 46 (18)
Fe nandes M.R.
hype -di e se d edging s a ions eco e ed up o 13 species,
mainly a dep hs below 50m, in which b ushing and suc ion
did no occu . In addi ion o he es ic ion o dep h, he e
we e mo e d edging s a ions han he sum o b ushing and
suc ion s a ions, and he a ea co e ed by a d edging is o en
la ge han o a AB o AS s a ion, which explain he highe
numbe o sampled species (Fig.22A). Howe e , d edging
usually samples emp y shells, con a y o b ushing and suc-
ion, which in mos cases ob ain li e ipho ids ( a ely shells
wi h he mi c abs; pe s. obs. o b ushing). Un o una ely, he
MADIBENTHOS expedi ion did no aim o he s o age o
li e ipho ids in e hanol, and he amoun o pho og aphed
li e specimens p obably does no e lec he ac ual numbe ,
a oiding u he compa isons.
The i e species cu en ly endemic om Ma inique (“In-
ella”sp., Iso ipho a pa dus n.sp., Iso ipho asp., Me axia discus
n.sp., Me axiasp.) ha e non-plank o ophic de elopmen
(Fig.23A), bu hei ex eme a i y (sampled as single ons o
double ons, wi h he excep ion o I.pa dus n.sp.) and he lack
o in ensi e sampling in o he pa s o he Lesse An illes a oid
such ce ain y. O he species sampled om Ma inique bu
absen om Guadeloupe a e Cosmo ipho a a noldoi, T ipho a
c . scylla and T ipho a po o icensis, al hough he wo o me a e
single on o double on. In con as , 25ou o 29species om
Guadeloupe (86.2%) a e sha ed wi h Ma inique (Fig.23B);
Nanapho a decolla a, Iso ipho a pee e sae Moolenbeek&
Fabe , 1989 (KARUBENTHOS s a ions: GS05 [2], GB11
[1], GS29 [1]) and T ipho a el i ae (KARUBENTHOS s a-
ion: GR10 [1]) we e absen om Ma inique, whe eas he
non-plank o ophic T ipho a guadaloupensis was absen om
MADIBENTHOS and KARUBENTHOS 1. The ipho id
auna om ABC Is. has 14 o he 18 eco ded species (77.8%)
sha ed wi h Ma inique, whe eas Abaco (Bahamas) sha es
wi h Ma inique 14 o 22 species (63.6%) (Fig.23B). The
simila i y d ops when compa ing shallow-wa e ipho ids
om Ma inique wi h hose om Cuba (21 o 40 species
sha ed, i.e., 52.5%), which has mo e species han Ma inique
due o a much la ge coas line and wi h esea che s ac i ely
desc ibing species since 1992; many ipho ids om Cuba
a e non-plank o ophic, which in luences hei endemism.
The (bio)geog aphic dis ance be ween Ma inique and B azil
pa ly explains he low numbe o sha ed species (20 o he
59 species om B azil, i.e., 33.9%); B azil has mo e known
ipho ids owing o a much la ge coas line, comp ising di -
e en ma ine p o inces (Spalding e al. 2007), in addi ion o
se e al species known om he end o he con inen al shel
(Fe nandes& Pimen a 2019a, 2020), a dep h zone poo ly
explo ed in Ma inique. An expec ed bu in iguing absence
om Ma inique is he plank o ophic T ipho a a lan ica
Smi h, 1890, which is widely dis ibu ed in he sou he n
USA and in B azil, bu wi h a single Ca ibbean eco d om
Pue o Rico (Rolán& Fe nández-Ga cés 2008; Fe nandes&
Pimen a 2020).
Ano he sampling bias in MADIBENTHOS was ela ed
o he geog aphic zones wi hin Ma inique, wi h he inc eas-
ing o de (excluding AN s a ions): Baie de Fo -de-F ance
(49s a ions), No d A lan ique (69), No d Ca aibe (95), Sud
A lan ique (131), Sud Ca aibe (147); i.e., he A lan ic side
had less s a ions (200) han he Ca ibbean side (291). E en
hough, he zones wi h mos ipho id species a e, in a dec eas-
ing o de : No d A lan ique (26 species / Chao1 es ima o :
26.1 species), Sud A lan ique (22 / 25.7), No d Ca aibe (20
lso ipho a icingula a
lnella spinosa
Cosmo ipho a a noldoi
“lnella” sp.
Me axia aeniola a
lso ipho a pa dus
Eu ipho a be mudensis
Co iopho a no em
Sageno ipho a osclausu
m
Similipho a in e media
T ipho a ellyae
T ipho a ma ii
T ipho a po o icensis
Ma shallo a mon ei oi
lni o is pseudo homae
T ipho a c . scylla
Me axia sp.
lso ipho a sp.
Similipho a sp.
Me axia discus
lni o is u is homae
Ma shallo a c . modes a
lni o is gudeliae
Nanapho a leei
La i ipho a albida
Chei odon a dupliniana
No o ipho a deco a a
Cosmo ipho a melanu a
Nanapho a e be nei
Monopho us oli aceus
Me axia excelsa
Me axia ugulosa
S obilige a inaudi a
Dep h (m)
0102030405060708090
Fig. 25. — Dep h ange o ipho id species in Ma inique, sampled by MADIBENTHOS.
495
T ipho idae om Ma inique
ZOOSYSTEMA • 2024 • 46 (18)
/ 22.0), Sud Ca aibe (18 / 27.0) and Baie de Fo -de-F ance
(11 / 12.0) (Fig.23C, D). In he nMDS g oupings (Fig.23E-
F), he shallowe (0-30m) s a ions om he Ca ibbean side
(including Baie de Fo -de-F ance) mainly g ouped oge he ,
jus like shallowe s a ions om he A lan ic side, despi e he
skewed ca ego y ‘Sud A lan ique 0-10m’ in he p esence/
absence ma ix (Fig.23F). Deepe (31-85m) s a ions also
g ouped close (Fig.23E-F), excep he ca ego y ‘Sud Ca aibe
61-85m’, which had a highly disc epan ela i e abundance
o Ini o is u is homae (72.7%).
Some hype -di e se s a ions a No d A lan ique de i ed
om deepe d edging (Fig.24A) lye close o he a bi a y
limi wi h No d Ca aibe (Bouche e al. 2019: 217), and
in luenced he highe numbe o ipho ids in he A lan ic
side o Ma inique (Fig.23C, D) despi e he educed num-
be o s a ions, ega ding o he po ions o he island lacked
d edging in deepe le els. The wide ‘shel ’ o he A lan ic
side migh ha e also con ibu ed o he inc eased numbe
o ipho ids owing o a supposed highe a ailabili y o mi-
c ohabi a s (e.g. pe haps mo e sponge hos s). As expec ed
Ini o is u is homae (34.3%)
No o ipho a deco a a (12.0%)
Monopho us oli aceus (11.1%)
Cosmo ipho a melanu a
(10.8%)
Ma shallo a c .
modes a (5.6%)
Sageno ipho a
osclausum (4.4%)
Ini o is gudeliae
(3.5%)
Nanapho a leei
(3.2%)
Similipho a in e media
(2.6%)
Me axia excelsa (2.6%)
T ipho a ellyae (2.3%)
La i ipho a albida (2.0%)
Nanapho a e be nei (2.0%) O he s
(3.2%)
Ini o is u is homae
(51.0%)
Cosmo ipho a melanu a
(10.8%)
Monopho us
oli aceus (6.6%)
No o ipho a deco a a
(6.4%)
Ini o is gudeliae (4.9%)
Sageno ipho a
osclausum (4.5%)
Co iopho a no em (3.6%)
Me axia excelsa (3.1%)
Ini o is pseudo homae (1.4%)
)
Similipho a in e media (1.2%)
Nanapho a leei (1.1%)
)
T ipho a po o icensis (1.1%)
O he s (4.1%)
Ini o is u is homae (48.5%)
Cosmo ipho a
melanu a (12.4%)
Monopho us
oli aceus (9.4%)
Ini o is
pseudo homae
(6.4%)
Me axia
excelsa (4.5%)
Sageno ipho a
osclausum (4.5%)
Co iopho a no em (4.2%)
No o ipho a deco a a (3.0%)
T ipho a po o icensis (1.5%)
Similipho a in e media (1.2%)
O he s (3.6%)
Ini o is u is homae
(12.0%)
Cosmo ipho a melanu a
(24.1%)
Ma shallo a
mon ei oi (14.4%)
Monopho us
oli aceus (7.2%)
Sageno ipho a
osclausum (7.2%)
Co iopho a
no em (6.0%)
Me axia
excelsa (4.8%)
Me axia
ugulosa (3.6%)
No o ipho a deco a a
(3.6%)
T ipho a ellyae (3.6%)
O he s (13.2%)
Cosmo ipho a melanu a (29.5%)
Ini o is u is homae (24.5%)
Co iopho a no em (8.0%)
Me axia excelsa
(8.0%)
No o ipho a deco a a
(5.3%)
Me axia ugulosa
(4.5%)
Ini o is pseudo homae
(3.1%)
Sageno ipho a osclausum (3.1%)
Inella spinosa (1.8%)
Monopho us oli aceus (1.8%)
T ipho a ellyae (1.8%)
O he s (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. — P opo ion o ipho id species sampled by MADIBENTHOS, acco ding o dep h zones and o al numbe o shells/specimens (n).
502 ZOOSYSTEMA • 2024 • 46 (18)
Fe nandes M.R.
S a ion Locali y La i ude (N) Longi ude (W) Ini ial dep h (m) Final dep h (m) Sampling me hod
GS04 G and Cul-de-sac ma in 16°21.97’ 61°37.98’ – 11 Suc ion
GB05 Poin e à Léza d 16°08.43’ 61°46.92’ – 12 B ushing
GS05 G and Cul-de-sac ma in 16°21.72’ 61°36.35’ – 22 Suc ion
GB06 Tê e à l’Anglais 16°22.9’ 61°45.94’ – 23 B ushing
GM06 Île Fo une 16°09’ 61°33.67’ – 1 Visual a in e idal
GS06 G and Cul-de-sac ma in 16°21.75’ 61°36.07’ – 23 Suc ion
GB07 Poin e du Quesy 16°06.07’ 61°46.37’ – 6 B ushing
GS07 Poin e à Léza d 16°08.43’ 61°46.92’ – 12 Suc ion
GB08 Poin e à Léza d 16°08.43’ 61°46.92’ – 12 B ushing
GM08 Poin e de l’E mi age 16°07.57’ 61°46.45’ – 1 Visual a in e idal
GS08 Tê e à l’Anglais 16°22.9’ 61°45.94’ – 23 Suc ion
GB09 Baie de Bouillan e 16°08.07’ 61°46.71’ – 6 B ushing
GS09 Poin e de l’E mi age 16°07.61’ 61°46.53’ – 11 Suc ion
GB10 Poin e de Malendu e 16°05.95’ 61°47.5’ – 8 B ushing
GD10 N Baie de Bouillan e 16°08.48’ 61°47.03’ – 54 D edging
GR10 Poin e à Léza d 16°08.43’ 61°46.92’ – 29 Visual a di ing
GB11 Cô e Fe y 16°17.69’ 61°48.23’ – 10 B ushing
GM11 Cab i 16°11.97’ 61°34.28’ – 1 Visual a in e idal
GS11 S Roc oy 16°02.38’ 61°45.71’ – 17 Suc ion
GB12 Po -Louis 16°25.61’ 61°32.57’ – 14 B ushing
GR12 Tê e à l’Anglais 16°22.9’ 61°45.94’ – 21 Visual a di ing
GS12 Baie de Bouillan e 16°08.07’ 61°46.71’ – 6 Suc ion
GB13 Po -Louis 16°23.26’ 61°31.79’ – 10 B ushing
GD13 Pe i e Anse 16°05.72’ 61°46.26’ – 5 D edging
GR13 Poin e de l’E mi age 16°07.61’ 61°46.53’ – 11 Visual a di ing
GS13 Île Pigeon 16°02.4’ 61°45.6’ – 50 Suc ion
GB14 Nea Po -Louis 16°23.74’ 61°32.07’ – 49 B ushing
GR14 Vieux Habi an 16°03.25’ 61°46.17’ – 27 Visual a di ing
GS14 Poin e de Malendu e 16°05.95’ 61°47.5’ – 8 Suc ion
GB15 S Po -Louis, W Pe i Canal 16°22.57’ 61°31.74’ – 8 B ushing
GD15 Anse à la Ba que 16°05.39’ 61°46.48’ – 50 D edging
GR15 Poin e du Quesy 16°06.07’ 61°46.37’ – 11 Visual a di ing
GS15 Sec Fe y 16°17.51’ 61°48.96’ – 27 Suc ion
GB16 Po -Louis 16°27.34’ 61°32.07’ – 19 B ushing
GD16 Anse Ca aïbe 16°12.37’ 61°47.2’ – 10 D edging
GS16 Po -Louis 16°25.61’ 61°32.57’ – 25 Suc ion
GB17 Poin e G ig i 16°23.26’ 61°31.79’ – 13 B ushing
GR17 Baie de Bouillan e 16°08.07’ 61°46.71’ – 13 Visual a di ing
GS17 Po -Louis 16°23.26’ 61°31.79’ – 3 Suc ion
GB18 Po -Louis 16°25.99’ 61°32.92’ – 45 B ushing
GS18 Nea Po -Louis 16°23.74’ 61°32.07’ – 49 Suc ion
GB19 Po -Louis 16°23.26’ 61°31.79’ – 11 B ushing
GR19 Île Pigeon 16°02.4’ 61°45.6’ – 15 Visual a di ing
GS19 S Po -Louis 16°22.57’ 61°31.74’ – 8 Suc ion
GB20 Po -Louis 16°26.78’ 61°32.41’ – 16 B ushing
GD20 Nea Baie de Baille-A gen 16°15.54’ 61°48.71’ – 35 D edging
GR20 Poin e de Malendu e 16°05.95’ 61°47.5’ – 8 Visual a di ing
GS20 Po -Louis 16°27.34’ 61°32.07’ – 19 Suc ion
GB21 T ou à l’o age 16°22.88’ 61°31.43’ – 8 B ushing
GD21 Nea Baie de Baille-A gen 16°15.55’ 61°48.8’ – 40 D edging
GR21 Sec Fe y 16°17.51’ 61°48.96’ – 27 Visual a di ing
GS21 Poin e G ig i 16°23.26’ 61°31.79’ – 14 Suc ion
GB22 Poin e Mon agnie 16°30.57’ 61°28.45’ – 12 B ushing
GS22 Poin e d’An igues 16°25.99’ 61°32.92’ – 45 Suc ion
GB23 G o e Amédie 16°30.04’ 61°28.79’ – 16 B ushing
GS23 Po -Louis 16°23.26’ 61°31.79’ – 7 Suc ion
GB24 Sec Pâ é 15°54’ 61°39.3’ – 25 B ushing
GD24 Po -Louis 16°25’ 61°33’ – 150 D edging
GS24 Po -Louis 16°26.78’ 61°32.41’ – 16 Suc ion
GB25 Ma ina de Ri iè e Sens 15°59’ 61°43.08’ – 25 B ushing
GD25 Po -Louis 16°25’ 61°33’ – 160 D edging
GS25 Poin e Mon agnie 16°30.57’ 61°28.45’ – 12 Suc ion
GB26 Nea Ca e 16°22.28’ 61°38.14’ – 29 B ushing
GS26 G o e Amédie 16°30.04’ 61°28.79’ – 16 Suc ion
GB27 Îlo Ca e 16°21.26’ 61°37.79’ – 2 B ushing
GN27 Po -Louis 16°23’ 61°33’ – 120 T ap lines
GR28 Po -Louis 16°27.34’ 61°32.07’ – 19 Visual a di ing
GM29 G and Cul-de-sac ma in 16°17.42’ 61°33.19’ – 1 Visual a in e idal
GS29 Nea Ca e 16°22.28’ 61°38.14’ – 29 Suc ion
GB30 Caye Pla e 16°10.97’ 61°32.41’ – 16 B ushing
Appendix 1. — Con inua ion.

503
T ipho idae om Ma inique
ZOOSYSTEMA • 2024 • 46 (18)
S a ion Locali y La i ude (N) Longi ude (W) Ini ial dep h (m) Final dep h (m) Sampling me hod
GS30 Îlo Ca e 16°21.26’ 61°37.79’ – 2 Suc ion
GB31 Pe i e Te e 16°09.71’ 61°07.73’ – 15 B ushing
GD31 Po -Louis 16°24.97’ 61°32.8’ – 85 D edging
GD32 Po -Louis 16°23.88’ 61°32.47’ – 80 D edging
GS32 W Fajou 16°21.35’ 61°35.79’ – 2 Suc ion
GD33 Po -Louis 16°24’ 61°33’ – 130 D edging
GM33 Anse Ta a e 16°15.37’ 61°11.92’ – 6 Visual a di ing
GS33 Caye Pla e 16°10.97’ 61°32.41’ – 14 Suc ion
GB34 Pe i e Te e 16°10.45’ 61°08.16’ – 10 B ushing
GR34 Poin e de la Fon aine 16°27.74’ 61°31.84’ – 9 Visual a di ing
GS34 Pe i e Te e 16°09.71’ 61°07.73’ – 15 Suc ion
GD35 Po -Louis 16°22.77’ 61°34.19’ – 66 D edging
GB36 W Pe i e Te e 16°09.45’ 61°10.5’ – 16 B ushing
GD36 Vieux-Bou g 16°22.87’ 61°33.05’ – 65 D edging
GR36 Poin e Mon agnie 16°30.57’ 61°28.45’ – 12 Visual a di ing
GS36 Pe i e Te e 16°07.87’ 61°12.52’ – 50 Suc ion
GR37 G o e Amédie 16°30.04’ 61°28.79’ – 16 Visual a di ing
GS37 Pe i e Te e 16°10.45’ 61°08.16’ – 10 Suc ion
GD39 G and Cul-de-sac ma in 16°17.34’ 61°33.32’ – 1 D edging
GS39 W Pe i e Te e 16°09.45’ 61°10.5’ – 16 Suc ion
GD42 G and Cul-de-sac ma in 16°21.12’ 61°32.33’ – 3 D edging
GD49 Pe i Cul-de-sac ma in 16°13.62’ 61°32.38’ – 3 D edging
GD50 Île Gosie 16°11.31’ 61°29.59’ – 22 D edging
GR50 Lagon de Pe i e Te e 16°10.55’ 61°06.67’ – 4 Visual a di ing
GD51 Île Gosie 16°11.44’ 61°29.34’ – 15 D edging
GD52 Île Gosie 16°12.09’ 61°29.71’ – 6 D edging
GD54 Nea Île Fajou 16°22.2’ 61°35.01’ – 60 D edging
GD55 Nea Île Fajou 16°22.48’ 61°35.46’ – 85 D edging
GD58 Nea Île Fajou 16°22.68’ 61°34.95’ – 95 D edging
GD59 Nea Île Fajou 16°22.55’ 61°35.38’ – 88 D edging
GD61 E Pe i e Te e 16°11.97’ 61°03.96’ – 80 D edging
GD65 Banc des Vaisseaux 16°08.14’ 61°16.96’ – 20 D edging
GD68 Banc des Vaisseaux 16°09.03’ 61°17.31’ – 33 D edging
GD69 Anse Ta a e 16°15.97’ 61°10.18’ – 60 D edging
KARUBENTHOS 2
DW4545 N G ande-Te e 16°29.7’ 61°31.4’ 60 82 D edging
DW4546 N G ande-Te e 16°31’ 61°32’ 268 306 D edging
DW4550 N G ande-Te e 16°37’ 61°31’ 432 482 D edging
DW4551 N G ande-Te e 16°31.3’ 61°24.5’ 59 74 D edging
DW4552 E La Dési ade 16°23.9’ 60°56.2’ 48 60 D edging
DW4555 E La Dési ade 16°24’ 60°51’ 100 258 D edging
DW4574 E La Dési ade 16°22’ 60°54’ 140 340 D edging
DW4583 N Ma ie-Galan e 16°09’ 61°19’ 233 396 D edging
DW4586 W Ma ie-Galan e 16°00’ 61°23’ 204 251 D edging
DW4587 W Ma ie-Galan e 16°01’ 61°24’ 281 406 D edging
DW4599 W Ma ie-Galan e 15°53’ 61°25’ 262 266 D edging
Appendix 1. — Con inua ion.