Calliostomatidae, Colloniidae, Margaritidae, and Solariellidae (Gastropoda: Trochoidea) collected by the Marion Dufresne (MD55) expedition in southeastern Brazil, with description of a new species of Calliostoma

Abstract

The deep-water mollusks collected during the Marion Dufresne (MD55) expedition to the southeastern Brazilian coast in 1987 have been studied in several recent works. The present paper lists and diagnoses 19 species belonging to the vetigastropod families Calliostomatidae, Colloniidae, Margaritidae, and Solariellidae. A new species, Calliostoma valkuri sp. nov., is described. It is mainly characterized by a very short teleoconch I, with limits marked by conspicuous varices and sculpture consisting of 3 spiral threads; sculpture of the teleoconch II consisting of spiral cords with thorn-like projections; and the presence of two strongly marked cords on the median portion of the whorls. Based on the MD55 samples and complementary specimens from other expeditions, the following species have their geographical and/or bathymetrical range extended: Bathymophila euspira, Calliostoma gemmosum, Calliostoma rude, Callogaza watsoni, Gaza compta, Falsimargarita terespira, Homalopoma boffii, Lamellitrochus cancapae comb. nov., Lamellitrochus pourtalesi, Margarites imperialis, and Solariella quinni.

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ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Accepted by A. Nützel: 19 Mar. 2019; published: 24 May 2019 401
Zootaxa 4609 (3): 401–428
https://www.mapress.com/j/zt/
Copyright © 2019 Magnolia Press Article
https://doi.org/10.11646/zootaxa.4609.3.1
http://zoobank.org/urn:lsid:zoobank.org:pub:F38E6655-0EC4-48E1-A002-CAC599F3556D
Calliostomatidae, Colloniidae, Margaritidae, and Solariellidae
(Gastropoda: Trochoidea) collected by the Marion Dufresne (MD55) expedition
in southeastern Brazil, with description of a new species of Calliostoma
DANIEL C. CAVALLARI1, RODRIGO B. SALVADOR2,3, ANA P. S. DORNELLAS4 & LUIZ R. L. SIMONE5
1Departamento de Biologia, Faculdade de Filosofia, Ciências e Letras de Ribeirão Preto, Universidade de São Paulo. Ribeirão Preto,
SP, Brazil. E-mail: dccavallari@usp.br
2Staatliches Museum für Naturkunde Stuttgart. Stuttgart, Baden-Württemberg, Germany.
3Museum of New Zealand Te Papa Tongarewa, Wellington, New Zealand. E-mail: salvador.rodrigo.b@gmail.com
4Instituto Oceanográfico, Universidade de São Paulo. São Paulo, SP, Brazil. E-mail: dornellas.anapaula@gmail.com
5Museu de Zoologia, Universidade de São Paulo. São Paulo, SP, Brazil. E-mail: lrsimone@usp.br
Abstract
The deep-water mollusks collected during the Marion Dufresne (MD55) expedition to the southeastern Brazilian coast
in 1987 have been studied in several recent works. The present paper lists and diagnoses 19 species belonging to the
vetigastropod families Calliostomatidae, Colloniidae, Margaritidae, and Solariellidae. A new species, Calliostoma valkuri
sp. nov., is described. It is mainly characterized by a very short teleoconch I, with limits marked by conspicuous varices
and sculpture consisting of 3 spiral threads; sculpture of the teleoconch II consisting of spiral cords with thorn-like
projections; and the presence of two strongly marked cords on the median portion of the whorls. Based on the MD55
samples and complementary specimens from other expeditions, the following species have their geographical and/or
bathymetrical range extended: Bathymophila euspira, Calliostoma gemmosum, Calliostoma rude, Callogaza watsoni,
Gaza compta, Falsimargarita terespira, Homalopoma boffii, Lamellitrochus cancapae comb. nov., Lamellitrochus
pourtalesi, Margarites imperialis, and Solariella quinni.
Key words: Calliostoma valkuri new species, deep-water, MD55 expedition, range extension, Vetigastropoda
Introduction
The MD55 expedition of the research vessel Marion Dufresne was born from the combined efforts of the Muséum
national d’Histoire naturelle (Paris, France) and Universidade Santa Úrsula (Rio de Janeiro, Brazil). Aiming at re-
trieving deep-water samples from off southeastern Brazil, the expedition took place in May–June 1987. It revealed a
staggering amount of deep-water mollusk species, several of which have been steadily described during the current
decade (e.g., Pimenta, 2012; Simone & Cunha 2012, 2014; Cavallari et al. 2014; Salvador et al. 2014). The present
work represents an additional effort focused on the gastropod families Calliostomatidae Thiele, 1924, Colloniidae
Cossmann, 1915, Margaritidae Thiele, 1924, and Solariellidae Powell, 1951.
Calliostomatidae is a very diverse family of typically small to medium-sized snails (8–100 mm) with trochiform
shells. It includes over 20 genera and 300 species found living on soft and rocky substrates in all oceans, ranging
from tropical to polar latitudes, from intertidal to bathyal (3,000 m) depths (Marshall 1995; Williams et al. 2010;
Vilvens 2014; Marshall 2016). The genus Calliostoma Swainson, 1840 s.l. within the subfamily Calliostomatinae is
the largest in the family, with over 200 species, 20 of which occur in Brazilian waters (Dornellas 2012; Cavallari &
Simone 2018).
The Colloniidae are characterized by shell features such as the absence of nacre, aperture outline, and a calcified
operculum, though some of these vary considerably (McLean 2012). Because of shell similarities and the calcified
operculum, Colloniidae was previously considered a subfamily of Turbinidae but has recently been treated as a
family (e.g., McLean & Kiel 2007; Williams et al. 2008; Bouchet et al. 2017). The oldest fossil record of the family
dates from the Jurassic (Monari et al. 1996). The family currently includes 19 genera (Bouchet & Gofas, 2011), of
which only Homalopoma Carpenter, 1864 occurs in Brazilian waters.

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Margaritidae was considered a subfamily of Trochidae Rafinesque, 1815, but was recently raised to full family
level by Williams (2012) with support from molecular data. Stoliczka (1868) originally introduced the name, but
his nomenclatural act has since been invalidated (Bouchet et al. 2005, 2017). The authorship is currently attributed
to Thiele (1924), who first reevaluated the group. Margaritids have small to medium-sized (rarely over 20 mm)
turbinoid-trochiform nacreous shells. Three margaritid genera have been recorded in Brazil to date (Rios, 2009),
namely Callogaza Dall, 1881, Gaza Watson, 1879, and Margarites Gray, 1847.
Solariellidae is a diverse, mostly deep-water group, with a circumglobal distribution. Species in this family
are usually small (5–20 mm) and live on fine sediment and unconsolidated substrates in tropical and extra-tropical
seas (Hickman & McLean 1990; Hickman 1998; Williams et al. 2013). Solariellidae was initially introduced as a
subfamily of Trochidae, but the name has been raised to family level by Wareìn & Bouchet (in Bouchet & Rocroi
2005) and this has been corroborated by molecular analyses (Williams et al. 2008). Currently, Solariellidae includes
14 genera, some of which were introduced recently (e.g., Vilvens et al. 2014; Vilvens & Williams 2016). At least
three sollariellid genera are present in Brazilian waters (Rios 2009), namely Microgaza Dall, 1881, Solariella Wood,
1842 and Suavotrochus Dall, 1924.
The present study lists 19 species belonging to the aforementioned Trochoidea families collected by the MD55
expedition, and additional expeditions (e.g., from Canopus Bank, off northeastern Brazil) included herein for com-
parative purposes. We report new range and bathymetric extensions for 11 species and describe a new calliostomatid
species based on shell morphology.
Material and methods
The material studied here mostly comprises empty shells collected by malacologists P. Bouchet, B. Métivier, and
J.H. Leal during the MD55 expedition. These specimens are currently held in the collections of the Muséum nation-
al d’Histoire naturelle (MNHN, Paris, France) and the Museu de Zoologia da Universidade de São Paulo (MZSP,
São Paulo, Brazil). Additional specimens from other institutions listed below, including types, were also examined
for comparative purposes. A list of examined material is given in each species entry. Images and measurements
were obtained with a Zeiss AxioCam MRc 5 and Zeiss AxioVision SE64 Rel 4.8 imaging software. SEM examina-
tion was carried out at the Staatliches Museum für Naturkunde Stuttgart (Stuttgart, Germany). Description of shell
structures of the new species follows the terminology of Marshall (1995).
The following abbreviations are used throughout the text. Specimens: D=greatest shell width (perpendicu-
lar to H); H=shell height (parallel to coiling axis); n=number of specimens; sh=empty shell(s); spm=preserved
specimen(s). Institutional: DMNH=Delaware Museum of Natural History (Wilmington, USA); MCZ=Museum
of Comparative Zoology, Harvard University (Cambridge, USA); MFP=Museo Felipe Poey (Havana, Cuba);
MHNG=Muséum d’Histoire naturelle Genève (Geneva, Switzerland); MORG=Museu Oceanográfico Prof. Eliezer
de Carvalho Rios (Rio Grande, Brazil); RMNH=Naturalis Biodiversity Center (former Rijksmuseum van Natuurli-
jke Historie; Leiden, The Netherlands); USNM=Smithsonian National Museum of Natural History (Washington,
D.C., USA).
Systematics
Family Calliostomatidae Thiele, 1924
Genus Calliostoma Swainson, 1840
Type species. Trochus conulus Linnaeus, 1758, by subsequent designation (Herrmannsen, 1846); Recent, Northeast
Atlantic and Mediterranean.

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Calliostoma caribbechinatum Landau, Van Dingenen & Ceulemans, 2017
(Fig. 1 A–D)
Synonymy see Leal (1991). Complement:
Calliostoma echinatum Dall, 1881: 47; Leal, 1991: 44, pl. 3, fig. A; Benkendorfer & Soares-Gomes, 2009: 147; Massemin et al.,
2009: 84, pl. p83; Rios, 2009: 51, text fig.; Rosenberg et al., 2009: 623; Dornellas, 2012: 103.
Calliostoma caribbechinatum Landau, Van Dingenen & Ceulemans 2017: 123.
Type material. Holotype: USNM 214270.
Type locality. Cuba, off Havana, Blake sta. 62, 146 m (Agassiz & Sigsbee leg., 1877).
Previously known distribution. From Cuba to southern Brazil, live specimens down to 146 m (Benkendorfer
& Soares-Gomes, 2009; Rosenberg et al., 2009).
New occurrence. Abrolhos Slope, 60–295 m (shells only).
Habitat. Coralline and muddy bottoms.
Material examined. Brazil: Espírito Santo: off São Mateus, Abrolhos Slope, 18°59’S, 37°50’W, MD55 sta.
DC75, 295 m, MNHN, 1 sh (27/v/1987); Jaseur Seamount, 20°26’S, 36°17’W, MD55 sta. DC30, 60 m, MNHN,
2 sh (15/v/1987), 20°27’S, 35°54’W, MD55 sta. DC34, 54 m, MNHN, 5 sh (15/v/1987); Dogaressa Seamount,
20°50’S, 33°44’W, MD55 sta. DC43, 63 m, MNHN, 3 sh (17/v/1987). Rio de Janeiro: off Cabo de São Tomé,
21°37’S, 40°18’W, MD55 sta. DC15, 37 m, MNHN, 1 sh (10/v/1987). Additional non-MD55 material: Cuba: ho-
lotype.
Measurements. 6½–7 whorls, H=4.9–6.1 mm, D=4.5–5.5 mm (n=12).
Remarks. The name Calliostoma caribbechinatum was introduced by Landau et al. (2017) as a replacement
for C. echinatum Dall, 1881, a junior secondary homonym of C. echinatum (Millet, 1865), a fossil species from
the Miocene of France. In his study about selected specimens from the MD55 expedition, Leal (1991) reported the
occurrence of C. caribbechinatum (as C. echinatum Dall, 1881) on the islands of Trindade and Martim Vaz as well
as the seamounts Jaseur, Davis, and Dogaressa, but did not mention the Abrolhos Slope (sta. DC75). This location
is newly reported herein and it is also the deepest occurrence of the species known so far, expanding the previous
maximum depth record from 220 m to 295 m. Even though the specimens examined here are exclusively empty
shells, some are very well preserved, which at least means they were collected not too long after death and likely
not far removed from their living area.
Calliostoma gemmosum (Reeve, 1842)
(Fig. 1 E–I)
Synonymy see Clench & Turner (1960) and Leal (1991). Complement:
Trochus gemmosus Reeve, 1842: 165, pl. 218, fig. 9.
Calliostoma gemmosum: Kempf & Matthews, 1968: 91, tab. I; Matthews & Kempf, 1970: 18; Leal, 1991: 45; Gomes et al.,
2006: 185; Benkendorfer & Soares-Gomes, 2009: 147; Rios, 2009: 51, text fig.; Dornellas, 2012: 103, fig. 47.
Type material. Former Stainforth Collection, probably lost (Clench & Turner, 1960).
Type locality. Not given, probably Western Atlantic (Clench & Turner, 1960).
Previously known distribution. Probably endemic to Brazil, ranging from Maranhão state to Espírito Santo
state, live specimens from 10–100 m (Leal, 1991; Rios, 2009).
New occurrence. Vitória-Trindade Chain.
Habitat. Sandy bottoms.
Material examined. Brazil: Espírito Santo: off São Mateus, Abrolhos Slope, 18°49’S, 37°57’W, MD55 sta.
DC83, 60 m, MNHN, 1 sh (28/v/1987); Vitoria Bank, 20°21’S, 36°59’W, MD55 sta. DC26, 97.5 m, MNHN, 1 sh
(14/v/1987); Montague Bank, 20°25’S, 36°42’W, MD55 sta. DC27, 81 m, MNHN, 1 sh (14/v/1987); Jaseur Bank,
20°42’S, 35°22’W, MD55 sta. DC35, 15–82 m, MNHN, 1 sh (16/v/1987). Additional non-MD55 material: Brazil:
Espírito Santo: off Guarapari, under rocks, 20–25 m, NMR 56382, 1 sh (xii/1991, ex coll. J.G.B. Nieuwenhuis).
Measurements. 7 whorls, H=11.0 mm, D=11.0 mm (largest whole specimen).
Remarks. Reeve’s introduction of C. gemmosum in Conchologia Systematica included a single figure depict-
ing a specimen in umbilical view (Reeve, 1842: pl. 218, fig. 9). A complementary description published a year later

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indicated the type locality as unknown (Reeve 1843). No further detail was given and the type material is presum-
ably lost. Nevertheless, subsequent works pointed out the Western Atlantic as the correct provenance of Reeve’s
specimens (Clench & Turner 1960). Although species identity is uncertain due to the absence of type specimens,
there is little doubt that the Brazilian specimens depicted and referred to as C. gemmosum in the works published so
far are part of a single, well-defined entity (Clench & Turner, 1960).
Leal (1991) did not report this species among the MD55 material, but the present specimens compare well with
Brazilian specimens described and illustrated so far (e.g., Rios, 1994, 2009). They also closely resemble specimens
from museum collections examined here in outline, sculpture pattern and ground color. Curiously, the apex and the
typical reddish-brown spiral stripes are much lighter in the MD55 specimens (Fig. 1 E–G). This could be due to
post-mortem discoloration since the color is more vivid in juvenile specimens (Fig. 1 H). Based on the very few
occurrences of C. gemmosum reported so far (Matthews & Kempf 1970; Leal 1991), the present records expand the
range of the species ca. 190 km south from 19°S to 20°42’S.
Calliostoma hassler Clench & Aguayo, 1939
(Fig. 2 A–G)
Synonymy see Dornellas & Simone (2013). Complement:
Calliostoma hassler Clench & Aguayo, 1939: 191, pl. 28, fig. 3; Abbott, 1974: 46; Massemin et al., 2009: 84, pl. p83; Dornellas,
2012: 104, figs 13–15; Dornellas & Simone, 2013: 285, figs 24–25, 44, 51–53, 91–98.
Astele (Leiotrochus) hassler: Macsotay & Campos, 2001: 35, pl. 13, figs 9–10.
Calliostoma sp.: Longo et al., 2014: 2, fig. 4F.
Type material. Holotype: MCZ 104554. Paratypes: MCZ 104555, 1 sh; MFP, 1 sh; all from type locality.
Type locality. Brazil, off Cabo Frio, 75 miles east of Rio de Janeiro, 64 m (20/ii/1872).
Distribution. From Paria, northern Venezuela, to southeastern Brazil, live specimens from 20–100 m (Macso-
tay & Campos 2001; Dornellas & Simone 2013).
Habitat. On bottoms alongside bryozoans and coralline algae (Clench & Turner, 1960).
Material examined. Brazil:Espírito Santo: off São Mateus, Jaseur Bank, 20°27’S, 35°54’W, MD55 sta. DC34,
54 m, MNHN, 1 sh (15/v/1987); Vitoria Bank, 20°32’S, 38°10’W, MD55 sta. DC 22, 52 m, MNHN, 1 sh; Davis
Bank, 20°39’S, 34°41’W, MD55 sta. DC40, 60 m, MNHN, 1 sh; Columbia Bank, 20°41’S, 32°12’W, MD55 sta.
DC47, 94–105 m, MNHN, 2 sh (19/v/1987); Dogaressa Bank, 20°54’S, 34°01’W, MD55 sta. DC42, 60 m, MNHN,
1 sh (17/v/1987); Rio de Janeiro: off Cabo de São Tomé, 21°37’S, 40°18’W, MD55 sta. DC15, 37 m, MNHN, 7 sh
(10/v/1987). Additional non-MD55 material: Brazil: Rio de Janeiro: types.
Measurements. 7¾ whorls, H=25.8 mm, D=25.7 mm (largest whole specimen).
Remarks. The single well-preserved adult individual from the MD55 assemblage (Fig. 2 A–B) is remarkably
similar to the type specimens in outline, sculpture, and coloration. In the original description, Clench & Aguayo
(1939) emphasized the yellowish hue of the types owing to a probable color loss after extended exposure to pre-
servative fluid. However, the material studied herein, as well as the specimens illustrated by Dornellas & Simone
(2013), suggest otherwise: the variation in color from yellowish (Fig. 2 A–B) to strongly red (Fig. 2 D) seems to
be the normal color range of this species. Moreover, even though most specimens have a homogeneous coloration,
except for some faint, darker colored axial stripes and white blotches (Fig. 2 D–F), some have a mixed coloration,
showing a reddish-orange hue on the first whorls that becomes yellowish towards the body whorl (Fig. 2 A). The
protoconch (Fig. 2 G), however, seems to be invariably ivory white. The record from the MD55 sta. CB77 is the
deepest known so far, though it consists of an empty, fragmentary shell.
There is a large conchological overlap between C. hassler and C. jujubinum (Gmelin, 1791), a species known
from Mexico and the Caribbean. Wide-ranging calliostomatid species are not entirely uncommon (Clench & Turner
1960), and a possible synonymy between these two should be further analyzed. Massemin et al. (2009) reported
both species occurring together in French Guyana.

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FIGURE 1. Calliostoma collected by the MD55 expedition. A–D. Calliostoma caribbechinatum. A. MNHN, sta. DC30, aper-
tural view (H=6.4 mm, D=5.8 mm). B. same, apical view. C. MNHN, from Trindade Island, juvenile specimen, apertural view
(H=1.9 mm, D=1.7 mm). D. MNHN, sta. DC43, detail of apex. E–I. Calliostoma gemmosum. E. MNHN, sta. DC83, apertural
view (H=11.0 mm, D=10.9 mm). F. same, apical view. G. same, umbilical view. H. MNHN, sta. DC27, juvenile specimen,
apertural view (H=3.3 mm, D=3.5 mm). I. MNHN, sta. DC26, detail of apex.

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FIGURE 2. Calliostoma collected by the MD55 expedition. A–G. Calliostoma hassler. A. MNHN, sta. DC47, apertural view
(H=25.8 mm, D=25.7 mm). B. same, umbilical view. C. MNHN, sta. DC15, apertural view (H=10.2 mm, D=11.1 mm). D.
MNHN, sta. DC40, juvenile specimen, apertural view (H=9.6 mm, D=12.2 mm). E. MNHN, sta. DC42, apertural view (H=3.6
mm, D=4.4 mm). F. MNHN, sta. DC22, apertural view (H=1.4 mm, D=1.8 mm). G. MNHN, sta. DC42 detail of apex. H–L.
Calliostoma javanicum. H. MNHN, sta. DC55, juvenile specimen, apertural view (H=5.4 mm, D=5.5 mm). I. MNHN, sta.
CB77, fragmentary specimen (H=~5.0 mm, D=~4.7 mm). J. MNHN, sta. DC55, juvenile specimen, detail of apex. K. same,
apical view. L. same, umbilical view.

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Calliostoma javanicum (Lamarck, 1822)
(Fig. 2 H–L)
Synonymy see Leal (1991). Complement:
Trochus javanicus Lamarck, 1822: 25.
Calliostoma javanicum: Leal, 1991: 45; Hess & Abbott, 1994: 142; Lyons, 1998: 16; Espinosa et al., 2007: 64, fig. 003; Wil-
liams et al., 2008: 487, tab. 2; Rosenberg et al., 2009: 622; Williams et al., 2010: 785, tab. 1.
Calliostoma sp.1: Leal, 1991: 47.
Type material. Holotype: MHNG-MOLL-51496.
Type locality. Java (in error; see Leal 1991).
Previously known distribution. East Florida to Brazil, live specimens from 1–97 m (Leal, 1991; Rosenberg et
al. 2009).
New occurrences. Off southeastern Brazil, Martim Vaz Island and Abrolhos Bank, 780–940 m (shells only).
Habitat. Benthic, on coral, rocky and sandy habitats (Rodríguez-Sevilla et al. 2009).
Material examined. Brazil: Espírito Santo: off São Mateus, Abrolhos Slope, 19°40’S, 37°48’W, MD55 sta.
CB77, 790–940 m, MNHN, 1 sh (27/v/1987); off Vitória, Martim Vaz Island, 20°32’S, 28°52’W, MD55 sta. DC55,
780–795 m, MNHN, 1 sh (20/v/1987). Additional non-MD55 material: holotype.
Measurements. ~6 whorls, H=5.4 mm, D=5.5 mm (juvenile).
Remarks. According to Rosenberg et al. (2009), the bathymetric range of this species is 1–97 m. The well-
preserved juvenile specimen found in the present samples at 780–795 m depth suggests a greater maximum depth,
though this single shell could still have been carried away from its original area. Leal (1991) reported C. javanicum
from Vitoria Bank (MD55 stations DC22, DC26), Montague Bank (sta. DC27), Jasseur Bank (sta. DC34, DC35),
Davis Bank (sta. DC40), Dogaressa Bank (sta. DC42) and Columbia Bank (sta. DC47). The species is newly re-
ported here from Trindade (off Martim Vaz Island) further eastward, and the Abrolhos Bank to the west.
Calliostoma rude Quinn, 1992
(Fig. 3 A–I)
Calliostoma rude Quinn, 1992: 83, figs 7–8; Massemin et al., 2009: 85.
Type material. Holotype: USNM 860251. Paratypes: UMML 305844, 2 sh, FSBC I 44076, 1 sh, from type local-
ity; MCZ 302593, Guyana, off Georgetown, Chain Cruise 35, sta. 35 and 36, 8°10.5–8°10N, 57°48’W, 53–60 m
(28/iv/1963).
Type locality. French Guyana, off Cayenne, Pillsbury sta. P-650, 6°07’N, 52°19’W, 84–91 m.
Previously known distribution. Guyana to French Guyana, live specimens from 53–91 m (Quinn 1992).
New occurrence. Southeastern Brazil, off Espírito Santo, Abrolhos Slope, 120–295 m (shells only).
Habitat. Sandy bottoms, 53–91 m.
Material examined. Brazil: Espírito Santo: off São Mateus, edge of the Abrolhos Slope, 18°59’S, 37°50’W,
MD55 sta. DC75, 295 m, MNHN, 1 sh (27/v/1987); 19°00’S, 37°47’W, MD55 sta. DC81, 120–135 m, MNHN, 1
sh (27/v/1987). Additional non-MD55 material: French Guyana: holotype.
Measurements. 9 whorls, H=13.1 mm, D=9.2 mm (adult specimen).
Remarks. The adult specimen analyzed herein (Fig. 3 A–C) compares exceedingly well with the holotype,
presenting nearly identical proportions, color, and sculpture. Shells of juvenile individuals (Fig. 3 D–I) have a some-
what flatter base, which also lacks the stronger sculpture of spiral ribs (Fig. 3 F) seen in adult individuals (Fig. 3 C).
The new records reported here are the second since the original description of this apparently rare species. The new
occurrence is far away from the previously known distribution in northern South America, extending its range ca.
3,200 km to southeastern Brazil.

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FIGURE 3. Calliostoma collected by the MD55 expedition, Calliostoma rude. A–C. MNHN, sta. DC81 (H=13.1 mm, D=9.2
mm). A. apetural view. B. apical view. C. umbilical view. D–I. MNHN, sta. DC75, juvenile specimen (H=5.0 mm, D=4.5 mm).
D. apertural view. E. apical view. F. umbilical view. G. SEM, apertural view. H. SEM, apical view. I. SEM, detail of apex.

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FIGURE 4. Calliostoma valkuri n. sp. Holotype MNHN-IM-2000-34300, sta. DC79 (H=4.1 mm, D=4.2 mm). A. apertural
view. B. apical view. C. umbilical view. D. SEM, apertural. E. SEM, apical view. F. SEM, umbilical view. G. SEM, detail of
spire. H. SEM, detail of apex showing the protoconch.
Calliostoma valkuri sp. nov.
(Fig. 4)
Type material. Holotype MNHN-IM-2000-34300.
Type locality. Brazil, Espírito Santo, off São Mateus, Abrolhos Slope, 19°01’S, 37°47’W, MD55 sta. CB79,
1500–1575 m (Bouchet, Leal & Métivier leg., 28/v/1987).
Etymology. The name refers to Valkur, a fictional demigod from the Forgotten Realms campaign setting of the
Dungeons and Dragons role-playing game. Valkur is the patron and protector of sailors.

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Diagnosis. Teleoconch I very short (0.1 whorl), sculptured by three very delicate spiral cords, beginning and
transition to teleoconch II marked by very prominent terminal varices. Teleoconch II sculpture consisting of spiral
cords with prominent sharp thorn-like projections; two cords more prominent than remaining cords located on me-
dian portion of whorl.
Description. Shell small (H=4.2 mm, juvenile specimen), trochoid, roughly as tall as wide, with 4 very convex
whorls; color homogeneous cream, with nacreous gleam underneath visible by translucency (Fig. 4 A–C). Spire
tall (height ~1/3 of shell height), prominent; spire angle ~60°. Protoconch (Fig. 4 E, G, H) prominent, of ~1 whorl
(width=0.4 mm), color indistinct from teleoconchs I/II, surface sculptured by intersected thin lines forming a re-
ticulate pattern of roughly hexagonal shapes; transition marked by conspicuous terminal varix. Teleoconch I (Fig.
4 G, H) very short (~0.1 whorl after protoconch), surface sculptured by 3 spiral threads; transition to teleoconch II
marked by conspicuous terminal varix. Teleoconch II of ~3½ convex whorls, mainly sculptured by 3–5 spiral cords
(Fig. 4 A, D), separated by wider interspaces (~3–4x cord width); ontogeny of cords as follows (Fig. 4 B, E): P2
and P3 commencing right after protoconch; P1 commencing at 1.5 teleoconch whorl; S1 commencing at 4.0 whorls;
S2 commencing at 4.1 whorls; cords presenting numerous large, spaced, sharp projections resembling rose thorns;
two cords, on median region of whorl, more prominent than remaining cords; suture very delicate, nearly indistin-
guishable (Fig. 4 G), not channeled; body whorl sculptured by 5 spiral cords, with 3 larger, dominant cords and
two thinner, intercalary cords. Aperture round, prosocline (Fig. 4 A, D), height ~1/2 of shell height, cream-colored,
nacreous; inner lip slightly flared, forming narrow callus (Fig. 4 D). Base convex, sculptured by 8 smooth, well-
marked spiral cords; umbilicus imperforate (Fig. 4 C, F); Operculum unknown.
Distribution. Known only from type locality (empty shell).
Measurements. ~4½ whorls, H=4.1 mm, D=4.2 mm (holotype).
Material examined. Holotype.
Remarks. A single shell among all the Calliostoma material looked clearly different from the remaining speci-
mens. Among the Western Atlantic congeners, the most similar species is C. militare Ihering, 1907 because both
have two strongly marked spiral cords on the median portion of the whorl bearing prominent spikes. The main dif-
ferences between these species are that C. valkuri sp. nov. has a shorter shell with a less rounded profile, and has
conical spikes, as opposed to the rounded ones in C. militare. Other similar species are C. echinatum, C. fascinans
Schwengel & McGinty, 1942, and C. sapidum Dall, 1881, but the similarity is almost exclusive to the juveniles, due
to the spikes on the spiral cords. Nevertheless, the shape of the spikes is much more conical, short and with blunt
rounded tips in these species, and all cords display similar width/length, as opposed to the more prominent median
cords of C. valkuri sp. nov.
Even without more samples, there are enough characters to easily distinguish it as a new entity. Calliostoma
valkuri sp. nov. can be readily told apart from its congeners by two strongly marked spiral cords on the median por-
tion of the whorl bearing prominent spikes resembling rose thorns; subsutural and suprasutural spiral cords similar to
the two stronger ones in structure, but much weaker, with less prominent spikes - probably, an adult specimen would
have more of such cords, as an increase in the number of cords as the shell grows is seen in nearly all congeners.
Genus Falsimargarita Powell, 1951
Type species. Margarites gemma E.A. Smith, 1915, by original designation; Recent, Antarctic Ocean.
Falsimargarita terespira Simone, 2008
(Fig. 5 A–C)
Falsimargarita terespira Simone, 2008: 15, figs 1–7; Dornellas & Simone, 2011: 39.
Type material. Holotype: MZSP 86789. Paratypes: MZSP 86784, 4 sh, Brazil, Santa Catarina, off Itajai, 26º53’S,
48º24’W, 400 m, otter trawl, inside Lophelia cf. pertusa (L.) (local fishermen leg., i/2006); MZSP 86786, 1 sh, from
type locality.
Type locality. Brazil, Santa Catarina state, off Cape Santa Marta, 28º37’S, 48º43’W (stomach content of batfish
Lophius sp.).

MD55 TROCHOIDEANS Zootaxa 4609 (3) © 2019 Magnolia Press · 411
FIGURE 5. Calliostomatidae and Colloniidae collected by the MD55 expedition, and specimens from Northeast Brazil. A–C.
Falsimargarita terespira MNHN, sta. CB103 (H=10.5 mm, D=10.7 mm). A. apertural view. B. apical view. C. umbilical view.
D–J. Homalopoma linnei. D–F. MNHN, sta. DC73 spm #1 (H=5.5 mm, D=4.7 mm). D. apertural view. E. apical view. F. um-
bilical view. G–I. MNHN, sta DC73 spm #2 (H=4.8, D=5.4 mm), under SEM. G. apical view. H. detail of apex. I–J. operculum
of specimen from Canopus Bank, off Northeast Brazil, MZSP 70295. I. inner side (L=1.5 mm), J. outer side. K–N. Homalo-
poma boffii, specimen from Canopus Bank, off Northeast Brazil, MZSP 93535. K–L. operculum (L=1.2 mm). K. inner side. L.
outer side. M. detail of apex under SEM, N. apertural view (H=7.3 mm, D=7.8 mm).

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Previously known distribution. Off Santa Catarina state, Cape Santa Marta and Itajaí, 200–400 m (live speci-
mens).
New occurrence. Southeastern Brazil, Rio de Janeiro, off Cabo Frio.
Habitat. Coralline and sandy bottoms.
Material examined. Brazil: Rio de Janeiro: off Cabo Frio, 23°36’S, 42°01’W, MD55 sta. CB103, 200–217 m
depth, MNHN, 10 sh (02/vi/1987). Additional non-MD55 material: Brazil: Santa Catarina: types.
Measurements. 5¾ whorls, H=10.5 mm, D=10.7 mm (largest specimen).
Remarks. The present material compares extremely well with the types. As remarked by Simone (2008), F.
terespira is the northernmost species of the genus. This distribution is herein extended further to the north (ca. 450
km) in Brazilian waters.
Family Colloniidae Cossmann, 1915
Genus Homalopoma Carpenter, 1864
Type species. Turbo sanguineus Linnaeus, 1758, by monotypy; Recent, northeastern Atlantic and Mediterranean.
Homalopoma boffii Marini, 1975
(Fig. 5 K–N)
Homalopoma boffii Marini, 1975: 31, figs 1–3. Rios, 1985: 26, pl. 11, fig. 109; Rios, 1994: 43, pl. 14, fig. 145; Rios, 2009: 69,
text fig. (switched with preceding species entry in error).
Homalopoma boffi [sic]: Leal, 1991: 58; Benkendorfer & Soares-Gomes, 2009: 151.
Type material. Holotype: MZSP 18772. Paratype: MZSP 18773, 1 spm, from type locality.
Type locality. Brazil, off Rio Grande do Sul state, 31°08’S, 49°31’W, 183–253 m, R/V “W. Besnard” sta. 1858
(6/viii/1972).
Previously known distribution. Off southeastern Brazil, from São Paulo state to Rio Grande do Sul state (Rios,
1994), including Trindade and Martim Vaz Islands, Montague and Columbia Seamounts (Leal, 1991), 60–253 m
(live specimens).
New records. Northeastern Brazil, off Ceará state, down to 260 m (shells only).
Habitat. Coralline and muddy bottoms.
Material examined. Brazil: São Paulo: off Santos, MZSP 36037, 2 spm (local fishermen leg., xii/2002). Ad-
ditional Non-MD55 material: Brazil: Ceará: 120 miles off Fortaleza, Canopus Bank, MZSP 53938, 240–260 m, 2
sh (viii/2005); MZSP 67303, 260 m, 7 sh (xi/2005); MZSP 70298, 260 m, 4 sh (xi/2005); MZSP 93535, 60–200 m,
2 sh (viii/2005); Rio Grande do Sul: types.
Measurements. 6–6½ whorls, H=8.7–10.0 mm, D=9.0–10.4 mm (n=3).
Remarks. This seemingly rare species is usually characterized by its large, reddish shell, with strong rounded
spiral cords and coarse nodulose sculpture starting on the teleoconch, but usually restricted to the first 3–4 whorls
(Marini 1975). The species, in fact, attains a larger size than reported in the original description (H=6.9, D=7.0 mm)
and followed by the literature (e.g., Rios 1994, 2009; Rosenberg et al. 2009), reaching a shell height of 10.9 mm
(MZSP 67303) and a maximum diameter of 10.8 mm (MZSP 36037) in non-MD55 specimens we analyzed (from
Canopus Bank, off northeastern Brazil).
Leal (1991) argued that the figures in the original description by Marini (1975: figs 1–3) were switched with
those of another species, but the present examination of the type material showed that the holotype is the illustrated
specimen: the proportions, sculpture and even the location of damaged areas are identical (MZSP 18772). Both type
specimens are subadult shells, and the slight change in outline with age (Fig. 5 N) may have caused confusion.
With its known distribution restricted to southern-southeastern Brazilian waters in recent works (Leal 1991;
Rios 1994, 2009), Homalopoma boffii is here revealed as more widely distributed than initially assumed, ranging
as far as northeastern Brazil. Part of this newly reported range overlaps with that of Homalopoma linnei (Dall,
1889), which has also been found in the MD55 material (see below). Marini (1975) pointed out the similarity of

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these species, arguing that the main differences resided in the shell’s ground color (reddish as opposed to pure white
in H. linnei), number of spiral cords (fewer in H. linnei), and overall size (~25% larger than H. linnei). Except for
the discrepancy in size, which is strongly consistent (and even amplified) in the assemblage examined here, the
remaining dissimilarities are not confirmed: some specimens of H. boffii from Canopus are almost pure white, and
the number of spiral cords on the body whorl is variable from 12 to 16 in both species. Even so, there are additional
differences to be noted. In H. boffii, the protoconch is much less inflated, nearly flattened (Fig. 5 M), and the very
first teleoconch whorl already bears coarse, large nodules. The nodules themselves are always rounder and propor-
tionally larger in H. boffii. Moreover, the shell of H. boffii has a more compact outline with a shorter spire, giving
it a more globose profile (Fig. 5 N). The suture is shallower in H. boffii, lacking the prominent, deep subsutural
channel present in H. linnei. The spiral cords in H. boffii are proportionally wider, as are the interspaces, which in
turn are shallower and less prominent than in H. linnei. The operculum in H. boffii is roughly circular (Fig. 5 K-L),
as opposed to squarish in H. linnei (Fig. 5 I-J).
Homalopoma linnei (Dall, 1889)
(Fig. 5 D–J)
Leptothyra linnei Dall, 1889: 353, pl. 33, fig. 8 (9 in error, see below).
Leptothyra linnei var. limata Dall, 1889: 353.
Homalopoma linnei: Poirier, 1954: 26; Abbott, 1974: 60; Rios, 1985: 26, pl. 11, fig. 110; Rios, 1994: 43, pl. 14, fig. 146; Miyaji,
2004: 79; Rios, 2009: 69 [in part]; Rosenberg et al. 2009: 624; Miloslavich et al., 2010: table S6; Welch, 2010: table S1.
Homalopoma sp.1: Leal, 1991: 58.
Type material. Syntypes: MCZ 7538, Cuba, Havana, off Morro Light, Blake sta. 02, 23°14’N, 82°25’W, 1472 m,
1 sh (R/V Blake leg., 1877); MCZ 7539, Cuba, off Havana, 822 m, 1 sh (R/V Blake leg., 1877); MCZ 7540, Cuba,
off Honda Bay, 23°2’30”N, 83°11’0”W, 402 m, 1 sh (R/V Blake leg., i/1878); NMNH 94984, Caribbean Sea, Bar-
bados, 13°17’N, 59°53’W, 183 m, 5 sh (R/V Blake leg.); NMNH 94985, Caribbean Sea, St. Lucia, Blake sta. 220,
13.8375N, 61.0625W, 212 m, 1 sh (R/V Blake leg., 16/ii/1879).
Type locality. Saint Lucia, BLAKE sta. 02, 1472 m depth; BLAKE sta. 20, 402 m depth; sta. 220, 212 m depth.
Cuba, off Havana, 823 m depth. Barbados, 183 m depth.
Distribution. Florida, USA to southern Brazil, 60–260 m (live specimens) down to 1472 m (shells only) (Miy-
aji 2004; Rosenberg et al. 2009).
Habitat. Coralline and muddy bottoms.
Material examined. Brazil: Espírito Santo: off São Mateus, Abrolhos Slope, MD55 sta. DC73, 18°59’S,
37°48’W, 607–620 m, MNHN, 4 sh (27/v/1987); MD55 sta. CB77, 19°40’S, 37°48’W, 790–940 m, MNHN, 1 sh
(27/v/1987). Additional non-MD55 material: Caribbean: types. Brazil: Ceará: 120 miles off Fortaleza, Canopus
Bank, MZSP 53726, 60–260 m, 24 sh (viii/2005); MZSP 55524, 240–260 m, 2 spm (xi/2005); MZSP 67309, 260 m,
5 sh (viii/2005); MZSP 70295, 260 m, 3 sh + 4 spm (xi/2005); MZSP 94229, 260 m, 4 sh (xi/2005).
Measurements. 6 whorls, H=6.26 mm, D=5.07 mm (largest specimen).
Remarks. In the original description, Dall (1889) cited figure 9 on plate 33. That figure, however, corresponds
to another taxon introduced in the same work (Arene bairdii, p. 389). The correct figure is number 8, which perfectly
agrees with the original description as well as the types examined herein. Due likely to a typographical error, the
figure number was switched with figure 8 before the final printing; one can easily notice that by the obvious inter-
ruption in the logical sequence of numbers.
The specimens studied here compare exceedingly well with the original description and the examined syntypes
in size, sculpture elements, and variation. Even so, most shells inspected have spiral cords sculptured by spiny
nodules on nearly all teleoconch whorls, which produce an overall rougher surface and a “spiky” appearance (Fig.
5 D–F). Dall (1889) already mentioned this variation, indicating it as rare, probably based on a syntype specimen
examined herein (UNSM 04984). Nevertheless, based on the current assemblage, this variation seems to be more
common than initially assumed. Leal (1991) described this variation as a probable distinct species based on MD55
material, tentatively naming it “Homalopoma sp.1”.
This species is quite similar to the sympatric congener Homalopoma boffii Marini, 1975, but differs in being small-
er, exclusively cream/white colored, and usually more profusely sculptured. Regarding the sculpture elements, the

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spiral cords referred to as “costae” by Dall in the original (1889) description, are usually narrower (though the num-
ber of spiral cords per whorl is variable and overlaps in both species) and the nodules are more irregular, angular
(spiky), smaller and numerous than in H. boffii. These nodules are usually more closely spaced in H. linnei, and are
often overlain giving the shell a scaly appearance (Fig. 5 G) that is very distinct from the larger and more widely
spaced nodules of H. boffii (Fig. 5 N). Nevertheless, both species display some degree of overlapping variation.
The only unequivocal characters that distinguish H. linnei from H. boffii in the analyzed material is the smoother,
more elevated and inflated protoconch of the former (Fig. 5 H), and the sculpture of the initial whorls, which lacks
the large, sequential nodules found on the initial teleoconch whorls in H. boffii (Fig. 5 M). Furthermore, the deep,
well-marked subsutural channel in H. linnei is not so pronounced in H. boffii. Most importantly, the operculum is
also strikingly distinct, having 9–10 whorls in H. linnei (Fig. 5 I-J), as opposed to 10–12 in H. boffii (Fig. 5 K-L),
and also presenting a prominent widening of the last whorl in H. linnei. This widening produces an almost squarish
outline (Fig. 5 I), which is different from the almost circular operculum of H. boffii (Fig. 5 K).
Family Margaritidae
Genus Callogaza Dall, 1881
Type species. Callogaza watsoni Dall, 1881, by subsequent designation (Clench & Abbott 1943); Recent, Atlantic.
Callogaza watsoni Dall, 1881
(Fig. 6 A–F)
Synonymy see Quinn (1979) and Simone & Cunha (2006). Complement:
Callogaza watsoni Dall, 1881: 50; Simone & Cunha, 2006: 22, figs 41–44, 58–60; Rosenberg et al., 2009: 624.
Gaza (Callogaza) watsoni: Quinn, 1979: 25; Rios, 2009: 55.
Type material. Holotype: MCZ 7544.
Type locality. Cuba, off Havana, 24°34’N, 83°16’W, 324 m (Blake Expedition, 1877).
Previously known distribution. From the Yucatan strait to Northeast Brazil (Simone & Cunha, 2006), live
specimens from 66–1170 m, usually deeper than 250 m (Quinn, 1979).
New occurrence. Southeast Brazil, off Espírito Santo, Abrolhos Slope.
Habitat: Coralline and sandy bottoms.
Material examined. Brazil: Espírito Santo: off São Mateus, Abrolhos Slope, 18°59’S, 37°50’W, MD55 sta.
DC75, 295 m, MNHN, 9 sh (27/v/1987); 18°59’S, 37°48’W, MD55 sta. DC73, 607–620 m, MNHN, 2 sh (27/
v/1987); 19°40’S, 37°48’W, MD55 sta. CB77, 790–940 m, MNHN, 2 lots (1 + 4 sh) (27/v/1987). Additional non-
MD55 material: Cuba: holotype.
Measurements. ~5½ whorls, H=7.2 mm, D=9.2 mm (largest juvenile specimen).
Remarks. The present specimens are young individuals with up to 5½ whorls. Adult specimens have around 7
whorls according to Simone & Cunha (2006). Additionally, juveniles lack a well-developed callus and reflected lip.
The MD55 specimens compare well with the holotype in that they present the strongly marked reticulate sculpture
on the apical side of the whorls (Fig. 6 B), the dominant well-marked spiral sculpture that produces a slightly keeled
shell profile (Fig. 6 A), and a similarly sculptured periumbilical region, with numerous axial furrows (Fig. 6 C). The
youngest specimens (~3¾ whorls, protoconch included) have a distinct rounder profile (Fig. 6 E), which is compat-
ible with the earliest whorls of older individuals. Nevertheless, the first spiral cords appear at 3½ whorls (Fig. 6
F), at which point the shell becomes characteristically keeled. The specimens analyzed here are very similar to the
individual reported by Simone & Cunha (2006: figs 58–60), which was mentioned by Dall (1889: 342) as a record
from Brazil. Dall’s specimen was collected off Fortaleza, Ceará state, R/V Albatross sta. 2756, at 716 m depth. The
present records thus represent a range expansion of over 1,800 km further south to Brazilian waters.

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FIGURE 6. Margaritidae collected by the MD55 expedition. A–F. Callogaza watsoni. A. MNHN, sta. CB77, spm #1, apertural
view (H=6.0 mm, D=8.0 mm). B. MNHN, sta. DC75, under SEM, apical view (D=6.7 mm). C. MNHN, sta. CB77, spm #2,
under SEM, umbilical view (D=6.2 mm). D. MNHN, sta. DC75 under SEM, detail of apex. E–F. MNHN, sta. SY74, juvenile
specimen (H=1.6 mm, D=2.3 mm). E. apertural view. F. apical view. G–L. Gaza compta. G–I. MNHN, sta. CB77, juvenile
specimen (10.2 mm, D=14.1 mm). G. apertural view, H. apical view. I. umbilical view. J–L. MNHN, sta. CB93, juvenile speci-
men #1 (H=1.7 mm, D=2.7 mm). J. apertural view. K. apical view. L. detail of apex under SEM.

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Genus Gaza Watson, 1879
Type species. Gaza daedala Watson, 1879, by original designation; Recent, Pacific.
Gaza compta Simone & Cunha, 2006
(Fig. 6 G–I)
Gaza compta Simone & Cunha, 2006: 4, figs 1–10, 49–52, 67–77, 92.
Type material. Holotype: MZSP 40324. Paratypes: MZSP 40325, 5 spm; ANSP 413312, 1 sh; MNRJ 10531, 1 sh;
ZSM 20060173, 1 sh, all from type locality.
Type locality. Brazil, Rio de Janeiro, off Cabo Frio, 22º53’S 42º01’W, 700–800 m depth (otter trawl, i/2004.
C.M. Cunha leg.).
Previously known distribution. Known only from type locality, 800–900 m (live specimens).
New occurrence. Off Espírito Santo, Abrolhos Slope, down to 940 m depth (shells only).
Material examined. Additional material: Brazil: Espírito Santo: off São Mateus, Abrolhos Slope, 18°59’S,
37°48’W, MD55, sta. DC73, 607–620 m, MNHN, 1 sh (27/v/1987); 19°36’S, 38°53’W, MD55 sta. CB93, 640 m,
MNHN, 1 sh (30/v/1987); 19°40’S, 37°48’W, MD55 sta. CB77, 790–940 m, MNHN, 1 sh (27/v/1987). Additional
non-MD55 material: Brazil: Rio de Janeiro: holotype.
Measurements. ~5¾ whorls, H=10.2 mm, D=14.1 mm (largest juvenile specimen).
Remarks. The MD55 specimens are all juvenile and fragmentary but retain enough conchological features to
allow identification, such as the faintly purple-colored initial teleoconch whorls that also show well-marked axial
undulations, the overall shell profile, and the compatible height/width ratio (~1.45 according to Simone & Cunha
2006). The new records from the MD55 expedition presented herein are the first published since the original de-
scription and expand the range of the species ca. 470 km to the north. The bathymetric range is also expanded from
the previously known maximum depth of 800 m to 940 m.
Genus Margarites Gray, 1847
Type species. Margarites diaphana Gray, 1847, by monotypy; Recent, North Atlantic Ocean [as Margarites helicinus (Phipps,
1774)].
Margarites imperialis Simone & Birman, 2006
(Fig. 7 A–F)
Margarites imperialis Simone & Birman, 2006: 14, figs 6–10; Rios, 2009: 104, text fig. (in part); Dornellas & Simone, 2011:
19.
Type material. Holotype: MZSP 46025. Paratypes: MZSP 46026, 12 sh, ANSP 412951, 1 sh, BMNH 20050261, 1
sh, from type locality; Rio Grande do Sul, off Porto Alegre, 30º43’S 44º05’W, MZSP 40193, ~90 m, 3 sh (Colella
leg., 1978).
Type locality. Brazil, Espírito Santo state, off Itapemirim, 21º37’S, 40º03’W, 900 m, IOUSP sta. E321 (09/
ii/1969).
Previously known distribution. Off South-Southeastern Brazil, 21.62°S to 26.75°S, 46.91°W to 40.05°W, live
specimens from 90–900 m.
New occurrence. As far northeast as the Abrolhos Slope, 19°36’S–18°58’S, 37°48’W–38°53’W, down to 1200
m (shells only).
Material examined. Brazil: Espírito Santo: off São Mateus, Abrolhos Slope, 18°58’S, 37°48’W, MD55 sta.
CB78, 1200 m, MNHN, 2 sh (27/v/1987); 19°36’S, 38°53’W, MD55 sta. CB93, 640 m, MNHN, 2 sh (30/v/1987).
Rio de Janeiro: off Cabo Frio, 23°46’S, 42°10’W, MD55 sta. CB105, 610 m, MNHN, 2 sh (06/v/1987). Additional
non-MD55 material: Brazil: types.

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FIGURE 7. Margaritidae and Solariellidae collected by the MD55 expedition. A–F. Margarites imperialis A–C. MNHN, sta.
CB105, spm #1 (H=8.8 mm, D=11.7 mm). A. apertural view. B. apical view. C. umbilical view. D. MNHN, sta. CB105, spm #2
(H=8.6 mm, D=11.2 mm), apertural view. E. MNHN, sta. CB78, juvenile specimen (H=5.5 mm, D=6.6 mm), apertural view.
F. MNHN, sta. CB93, juvenile specimen (H=4.3 mm, D=5.2 mm), apertural view. G–L. Bathymophila euspira. G–H. MNHN,
sta. DC72 (H=4.0 mm, D=4.1 mm). G. apertural view. H. apical view. I. MNHN, sta. CB77 (D=3.2 mm), umbilical view. J–L.
MNHN, sta. DC72 under SEM. J. apertural view. K. apical view. L. detail of apex.

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Measurements. 5 whorls, H=8.8 mm, D=11.7 mm (largest specimen).
Remarks. The present material compares extremely well with the types, showing a shell sculptured by uniform
cords that become more closely spaced at the base, three more widely spaced spiral cords from the periphery to
the adjacent suture on each whorl (Fig. 7 A, D), a subsutural cord with round nodules (Fig. 7 B), and an umbilicus
covered by a smooth callus (Fig. 7 C). A single exception is a poorly preserved specimen that exhibits both the sub-
sutural spiral cord and the following cord ornamented by triangular spikes (Fig. 7 E). A considerable conchological
variation in sculpture strength is known in many trochoid species (e.g., Quinn 1979; Marshall 1999). Thus, this case
is here interpreted as intraspecific conchological variation. The present record extends the range of the species ca.
320 km northwest to the Abrolhos Slope and the maximum depth from 900 m to 1200 m.
Family Solariellidae
Genus Bathymophila Dall, 1881
Type species. Margarita euspira Dall, 1881, by monotypy; Recent, Portugal, Gulf of Mexico, Caribbean Sea.
Bathymophila euspira (Dall, 1881)
(Fig. 7 G–L)
Synonymy see Quinn (1979). Complement:
Margarita euspira Dall, 1881: 44.
Trochus euspira: Jeffreys, 1883: 98, pl. 20, figs 6, 6a, 6b.
Margarites (Bathymophila) euspira: Quinn, 1979: 5, figs 1–2.
Bathymophila euspira: Marshall, 1999: 20.
Margarites euspira: Rosenberg et al., 2009: 624; Welch, 2010: table S1.
Margarites euspirus: Miloslavich et al., 2010: table S6.
Type material. Unknown (Quinn 1979).
Type locality. Cuba, off Havana, Blake sta. 2, 23º14’N, 82º25’W, 1472 m.
Previously known distribution. Portugal, Gulf of Mexico, Caribbean Sea, live specimens from 713–1472 m
(Rosenberg et al., 2009).
New occurrence. Southwestern Atlantic, Brazil, off Espírito Santo, Abrolhos Slope.
Habitat. Coralline and muddy bottoms.
Material examined. Brazil: Espírito Santo: off São Mateus, Abrolhos Slope, 19°00’S, 37°48’W, 950–1050 m,
MNHN, 2 sh; 19°40’S, 37°48’W, MD55 sta. CB77, 790–940 m, MNHN, 1 sh (27/v/1987).
Measurements. 4½ whorls, H=4.0 mm, D=4.1 mm (largest specimen).
Remarks. The diagnostic spiral sculpture pattern of the early whorls, as well as the overall shell shape and
proportions and the large circular aperture, is consistent with Bathymophila euspira. The holotype of this species
could not be located. However, topotypes and the syntypes of the variety (and synonym) B. euspira nitens Dall,
1881 (USNM 214275, from off Portugal), alongside the redescription of Quinn (1979), support the identifica-
tion of the present specimens. Bathymophila euspira is known for its considerable conchological variation, from
completely smooth specimens to a variable number of fine spiral striae and having a subsutural spiny spiral thread
(Quinn 1979). The present specimens lack the subsutural spiral thread and are almost completely smooth (Fig. 7 G,
J), counting with circa 8 very fine spiral striae on the base of the whorl, circling the umbilicus (Fig. 7 I, J). More-
over, the present specimens have narrower shells than the typical B. euspira. The present record greatly extends the
species range ca. 6,900 km to the southeast, which is relatively common for the MD55 material (e.g., Calliostoma
echinatum, above).
Genus Lamellitrochus Quinn, 1991
Type species. Margarita lamellosa Verrill & Smith [in Verrill], 1880; Recent, New Jersey to Gulf of Mexico.

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Lamellitrochus cancapae (Vilvens & Swinnen, 2007) comb. nov.
(Fig. 8 A–C)
Solariella cancapae Vilvens & Swinnen, 2007: 124, figs 1–2.
Type material. Holotype: RNMH.MOL.109036.
Type locality. Portugal, Azores, east of Flores, CANCAP-V sta. 5171, 39º20’N, 30º53’W, 1874–1887 m.
Previously known distribution. Known only from type locality, 1874–1887 m (empty shells).
New occurrence. Western Atlantic, Brazil, off Espírito Santo, Abrolhos slope, 607–620 m (shells only).
Material examined. Additional material: Brazil: Espírito Santo: off São Mateus, continental slope of Abrol-
hos, 18°59’S, 37°48’W, MD55 sta. DC73, 607–620 m, MNHN, 2 sh (27/v/1987). Additional non-MD55 material:
Portugal: Azores: holotype.
Measurements. 6¼ whorls, H=9.1 mm, D=9.0 mm (largest specimen).
Remarks. The species was originally classified in the genus Solariella S. Wood, 1842, but we propose here a
new combination within the Atlantic genus Lamellitrochus. This is justified by the following characters (as defined
by Quinn 1991): (1) The angular shell profile formed by a subsutural cord, a peripheral carina and a basal carina,
with nearly flat regions between them. The two carinae are fainter in the rounded body whorl of L. cancapae, al-
though they can be easily seen in earlier whorls. (2) A tuberculate subsutural cord. (3) Early whorls sculptured by
strong spiral cords and well-marked axial riblets.
The present material compares exceedingly well with the holotype (and only known specimen) of L. cancapae
(RMNH.MOL.109036) in overall dimensions, outline with a rounded body whorl (Fig. 8 A–B), and main spiral
sculpture elements (see Vilvens & Swinnen 2007 for a full description). However, the present specimens only show
spiral striae on the whorl base (Fig. 8 C) and do not have axial sculpture, thus they are lacking the type’s reticulate
sculpture pattern; they also lack interumbilical spiral striae. Nevertheless, as seen in other solariellids, the strength
of the sculpture can be very variable intraspecifically (Quinn 1979; Marshall 1999) and thus, the present differences
are here interpreted as morphological variation. The present record greatly extends the species range to the Western
Atlantic and ca. 6,500 km to the south.
Lamellitrochus carinatus Quinn, 1991
Lamellitrochus carinatus Quinn, 1991: 84, figs 7–12, 34–35; Barros et al., 2008: 25, figs 50–56; Benkendorfer & Soares-Gomes,
2009: 147; Rios, 2009: 57, text fig.; Rosenberg et al., 2009: 623; Barros, 2010: 69, figs 50–56; Welch, 2010: table S1.
Solariella sp. 1: Leal, 1991: 47, pl. 3, figs B–D.
Type material. Holotype: DMNH 179393. Paratypes: Brazil, Espírito Santo, off Trindade Island, 20°29’02’’S,
29°18’02’’W, MD55 sta. DC61, 63 m, MORG 26529 (22/v/1987); 20°30’04’’S, 29°18’05’’W, MD55 sta. DC59,
52–60 m, MNHN-IM-2012-37681 (22/v/1987).
Type locality. USA, Florida, SW of Egmont Key, 366–421 m depth (J. Moore leg.).
Distribution. North Carolina, USA and Trindade Island, Brazil, 52–421 m depth (Quinn 1991).
Material examined. Brazil: Espírito Santo: paratypes.
Measurements. ~4¼ whorls, H=2.5–2.7 mm, D=2.7–2.9 mm (Quinn 1991).
Remarks. The paratypes, stemming from the MD55 expedition, were already studied and reported by Quinn
(1991). No further specimens of L. carinatus were found in the remaining MD55 material.
Lamellitrochus pourtalesi (Clench & Aguayo, 1939)
(Fig. 8 D–F)
Synonymy see Quinn (1979). Complement:
Solariella (Solariella) pourtalesi: Quinn, 1979: 42, figs 63–64.
Lamellitrochus pourtalesi: Quinn, 1991: 83, figs 4–6; Rosenberg et al., 2009: 623; Barros, 2010: 67, figs 43–49.

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FIGURE 8. Solariellidae collected by the MD55 expedition. A–C. Lamellitrochus cancapae MNHN, sta. CB73 (H=9.1 mm,
D=9.0 mm). A. apertural view. B. apical view. C. umbilical view. D–F. Lamellitrochus pourtalesi. D. MNHN, sta. CB76 (H=5.2
mm, D=5.2 mm), apertural view. E–F. MNHN, sta. CB79, juvenile specimen (H=3.5 mm, D=3.4 mm) under SEM. E. apertural
view. F. umbilical view.
Type material. Holotype: MCZ 135018. Paratypes: MCZ 7579, Cuba, NW of Havana, 23°42‘N, 83°13‘W, 1572 m
(24/ii/1878); MCZ 7585, Mexico, Yucatan Strait, 1170 m; MCZ 135019, Cuba, 23°24’N, 80°44’W, 1060 m (W.C.
Schroeder leg., 1938); MCZ 135020, Cuba, 23°10’N, 81°29’W, 420 m (W.C. Schroeder leg., 1938).
Type locality Cuba, Atlantis sta. 2993, 23°24’N, 80°44’W, 1061 m.
Previously known distribution. Florida, USA, to Alagoas state, Brazil, 275–2350 m (Quinn, 1979; Barros,
2010).
New occurrence. Brazil, off Espírito Santo state, Abrolhos Slope.
Habitat. Quinn (1979) states that the species is more common in depths greater than 1000 m.
Material examined. Brazil: Espírito Santo: off São Mateus, Abrolhos Slope, 18°58’S, 37°49’W, MD55 sta.

MD55 TROCHOIDEANS Zootaxa 4609 (3) © 2019 Magnolia Press · 421
CB76, 637 m, MNHN, 2 sh (27/v/1987); 19°01’S, 37°47’W, MD55 sta. CB79, 1500–1575 m, MNHN, 3 sh (28/
v/1987). Additional non-MD55 material: Mexico/Cuba: types.
Measurements. 5 whorls, H=5.2 mm, D=5.2 mm (largest specimen).
Remarks. The present material compares well to the types of L. pourtalesi. The species is readily diagnosable
by its sculpture pattern (see Quinn 1979 for a full description). The present record greatly expands the species range,
ca. 1,000 km to the south.
Genus Solariella S. Wood, 1842
Type species. Solariella maculata S. Wood, 1842, by monotypy; Pliocene, England.
Solariella carvalhoi Lopes & Cardoso, 1958
(Fig. 9 A–C)
Solariella carvalhoi Lopes & Cardoso, 1958: 59, figs 1–3; Rios, 1970: 23; Abbott, 1974: 41; Rios, 1975: 21, pl. 4, fig. 45;
Rios, 1985: 23, pl.10, fig. 95; Calvo, 1987: 59, fig. 24; Rios, 1994: 35, pl. 10, fig. 105; Miyaji, 2004: 78; Barros, Santos &
Francisco, 2008: 11, figs 8–14; Rios, 2009: 55, text fig.; Barros, 2010: 58, figs 8–14; Daccaret & Bossio, 2011: 54, fig. 92;
Dornellas & Simone, 2011: 10, figs 41–43; Cavallari et al., 2016: 1, figs 1–23.
Solariella sp. 2: Leal, 1991: 47, pl. 3, figs E–G.
Solariella quadricincta Quinn, 1992: 50, figs 1–4; Barros, Santos & Francisco, 2008: 9, figs 1–7; Barros, 2010: 56, figs 1–7;
Miloslavich et al., 2010: table S6.
Solariella staminea Quinn, 1992: 53, figs 11–12; Benkendorfer & Soares-Gomes, 2009: 147.
Type material. Holotype: MZSP 18446. Paratype: Paulo de Sá Cardoso Collection, same data as Holotype (prob-
ably lost; J. Heise, pers. comm.).
Type locality. Brazil, off São Paulo state, 31°35’08”S, 50°50’00”W, 54 m (Moreira leg., 11/xi/1956).
Distribution. From Venezuela to southern Brazil, live specimens from 0–350 m (Cavallari et al. 2016).
Habitat. Coralline and muddy bottoms.
Material examined. Brazil: Espírito Santo: Davis Seamount, 20°40’S, 34°40’W, 60 m, MNHN-IM-2012-
37682, 3 paratypes of S. staminea, MORG 26530, holotype of S. staminea MD55 sta. DC40 (17/v/1987). Rio de
Janeiro: off Cabo Frio, 22°58’S, 42°06’W, MD55 sta. CB101, 50 m, MNHN, 4 sh (01/vi/1987). Additional non-
MD55 material: Brazil: São Paulo: holotype.
Measurements. 5–6 whorls, H=5.0–7.0 mm, D=5.5–8.0 mm (n=4).
Remarks. The present material compares well with S. carvalhoi. This species can be readily told apart from
the local congener Solariella quinni Barros & Pereira in Barros et al., 2008 (Fig. 9 D–G - see below) mainly by
its larger and proportionally wider shell, its much wider umbilicus (Fig. 9 C), which was correctly emphasized by
Lopes & Cardoso (1958), its more delicate axial and much stronger spiral sculpture (Fig. 9 A–B). The two remaining
local congeners, S. quadricincta Quinn, 1992 and S. staminea Quinn, 1992, were considered junior synonyms of S.
carvalhoi by Cavallari et al. (2016). Their study was based on a large sample, which revealed that both morphotypes
(S. quadricincta and S. staminea) fall within a continuum of conchological variation.
Solariella quinni Barros & Pereira in Barros et al., 2008
(Fig. 9 D–G)
Solariella quinni Barros & Pereira in Barros et al., 2008: 13, figs 15–21; Barros, 2010: 60, figs 15–21; Cavallari et al., 2016:
44.
Type material. Holotype: MZSP 77083. Paratypes: MZSP 77082, 08°11’S, 34°34’W, 66–71 m (18/xii/2004); MZSP
77084, 08°11’S, 34°36’W, 69–71 m (19/xii/2004); MZSP 77085, 08°11’S, 34°36’W, 69–71 m (19/xii/2004).
Type locality. Brazil, off Pernambuco, 08°09’S, 34°34’W, gravel bottom, 51–71 m (19/xii/2004).

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FIGURE 9. Solariellidae collected by the MD55 expedition. A–C. Solariella carvalhoi MNHN, sta. CB101 (H=6.9 mm, D=7.8
mm). A. apertural view. B. apical view. C. umbilical view. D–G. Solariella quinni MNHN, sta. DC73 (H=4.2 mm, D=4.2 mm).
D. apertual view. E. apical view. F. umbilical view. G. SEM, protoconch detail. H–L. Suavotrochus lubricus. H–J. MNHN, sta.
DC75, apertural view (H=4.8 mm, D=3.5 mm). I. apical view. J. umbilical view. K. MNHN, sta. DC75, detail of protoconch
under SEM. L. MNHN, sta. CB93, juvenile specimen with ribbed sculpture (H=2.2 mm, D=2.1 mm).
Previously known distribution. Northeast Brazil, off Pernambuco state, live specimens from 51–71 m (Barros
et al. 2008).

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New occurrence. Espírito Santo state, off Southeast Brazil, Abrolhos Slope, 290–620 m (shells only).
Habitat. Gravel and muddy bottoms (Barros et al. 2008; present study).
Material examined. Brazil: Espírito Santo: off São Mateus, Abrolhos Slope, 18°59’S, 37°48’W, MD55 sta.
DC73, 607–620 m, MNHN, 1 sh (27/v/1987); 18°59’S, 37°49’W, MD55 sta. CB80, 290–315 m, MNHN, 1 sh (28/
v/1987). Additional non-MD55 material: Brazil: Pernambuco: types.
Measurements. 5 whorls, H=4.2 mm, D=4.2 mm (largest specimen).
Remarks. The present specimens compare well with the type material. The species can be readily diagnosed by
the sculpture pattern on the whorl shoulder (Fig. 9 E) and base, as well as by the narrow, internally smooth umbilicus
(Fig. 9 F). This species is similar to S. lacunella (Dall, 1881) from the Caribbean, but can be distinguished from it
by: (1) being much smaller (H=~4.2 mm, as opposed to 8.5 mm in S. lacunella); (2) presenting an overall stronger
axial sculpture with conspicuous axial ribs on the whorl shoulder, and weaker spiral sculpture with narrower cords,
especially on the base; (3) having a more convex base; and (4) presenting a much narrower umbilicus. The records
presented here greatly expand the species range ~1,200 km southward to Espírito Santo state. The maximum depth
for the species is also updated herein from 71 to 620 m.
Solariella quinni was originally described by Barros et al. (2008). The subsequent paper by the senior author
(Barros 2010) is an almost entirely verbatim repetition of the previous version and is herein considered a re-publica-
tion, even though it does not contain a reference to the former.
Genus Suavotrochus Dall, 1924
Type species. Margarita lubrica Dall, 1881, by monotypy and original designation; Recent, Florida Straits to Brazil.
Suavotrochus lubricus (Dall, 1881)
(Fig. 9 H–L)
Synonymy see Quinn (1979). Complement:
Margarita lubrica Dall, 1881: 44.
Solariella (Suavotrochus) lubrica: Quinn, 1979: 42, figs 68–74.
Solariella lubrica: Springer & Bullis, 1956: 25; Lopes & Cardoso, 1958: 62; Rios, 1975: 21, pl. 4, fig. 46; Rios, 1985: 23, pl. 10,
fig. 96; Rios, 1994: 36, fig. 109; Gracia et al., 2004: 50, fig. 17; Miyaji, 2004: 78; Benkendorfer & Soares-Gomes, 2009:
150; Rios, 2009: 56, text fig.; Rosenberg et al., 2009: 623; Barros, 2010: 62, figs 22–28; Miloslavich et al., 2010: table S6;
Welch, 2010: table S1; Daccaret & Bossio, 2011: 54, fig. 91.
Suavotrochus lubricus: Bouchet, 2015.
Type material. Lectotype: USNM 95061.
Type locality. Cuba, North of Havana, Blake sta. 2, 23º14’N, 82º25’W, 1472 m (24/v/1893).
Distribution. From Florida Straits to southernmost Brazil, 73–1472 m (Rosenberg et al. 2009). Quinn (1979),
however, remarked that the species might actually inhabit only depths up to 500 m, being carried post-mortem to
greater depths. Barros (2010) gave a bathymetric range of up to 5633 m of depth based on empty shells.
Habitat. Muddy bottoms (Barros et al., 2008)
Material examined. Brazil: Espírito Santo: off São Mateus, continental slope of Abrolhos, 18°59’S, 37°50’W,
MD55 sta. DC75, 295 m, MNHN, 100 sh (27/v/1987). Rio de Janeiro: East of Cabo de São Tomé, 21°31’S,
40°08’W, MD55 sta. CB96, 295–300 m, MNHN, 1 sh (31/v/1987); S of Cabo Frio, 23°41’S, 42°06’W, MD55 sta.
CB104, 430–450 m, MNHN, 50 sh (02/vi/1987). Additional non-MD55 material: Cuba: lectotype.
Measurements. 5½–6 whorls, H=5.0–5.5 mm, D=4.0–4.5 mm (n=20).
Remarks. This species is by far the most abundant trochoid found in the MD55 material. The present speci-
mens compare very well with the lectotype, even though they show an overall less-pronounced sculpture; this is
in line with the known degree of morphological variation of this species (Quinn 1979). The variety (subspecies)
Margarita lubrica iridae was described by Dall (1889) as a nearly smooth form (lectotype USNM 95063, paralec-
totype USNM 887343), but was later considered a synonym of S. lubricus by Quinn (1979), who observed several
intermediate stages between these two forms.
Interestingly, some specimens from one locality (sta. CB96) have the teleoconch sculptured by regularly spaced

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424 · Zootaxa 4609 (3) © 2019 Magnolia Press
well-marked prosocline ribs. Moreover, S. lubricus has some axial corrugations restricted to the periumbilical area
(Fig. 9 J), but in these specimens the corrugations seem to be expanded, forming complete ribs (Fig. 9 L). To the best
of our knowledge, this feature has never been reported for this species. In the present specimens, the axial sculpture
is of variable strength and there are even completely smooth specimens.
Discussion
A total of 19 species were found in the present sample, including Calliostoma valkuri sp. nov. This study extends
the geographic and/or bathymetric ranges for 11 species, some to a great extent. The following species have their
geographical range extended within Brazilian waters: Calliostoma gemmosum (190 km southward), Falsimargarita
terespira (450 km to the north), Gaza compta (470 km to the north), Callogaza watsoni (1,800 km to the south),
Margarites imperialis (320 km northwest), Lamellitrochus pourtalesi (1,000 km to the south), and Solariella quinni
(1,200 km southward). Species from other regions that are here newly reported from Brazil are: Calliostoma rude
from the Guianas, Bathymophila euspira from the Caribbean, and Lamellitrochus cancapae from the Azores.
A few species (e.g., Suavotrochus lubricus) are extremely abundant in the sample, while others (e.g., Calliosto-
ma javanicum, Solariella quinni and Calliostoma valkuri sp. nov.) appear to be much rarer, being reported here from
just a few or even a single specimen. Large range extensions such as the present ones were also reported in other
studies of MD55 material, including other vetigastropods (e.g., Salvador et al. 2014). Overall, the amount of new
information brought in by the present study is on par with previous works focused on gastropods collected during
the MD55 expedition (e.g., Simone & Cunha 2014; Salvador et al. 2014). Sampling efforts such as this expedition
have added a wealth of information to the knowledge on Brazilian marine snails, especially to that of deep-water
species, of which we still know surprisingly little about. Fortunately, studies about deep-sea species in Brazil have
received increased attention in recent years. Knowledge of this fauna is imperative for further environmental studies
and proper legal protection. This is especially true when it comes to potential petroleum extraction areas, which can
greatly affect the environment and the local benthic biota (OSPAR 2009). The new species described herein, which
appears to be extremely rare, illustrates this situation very well. Since it is only known from the type locality, caus-
ing disturbances in the area could greatly affect it, or even ultimately bring it to extinction.
Acknowledgements
We are grateful to Philippe Bouchet (MNHN) for the opportunity to study the MD55 material; to Karin Wolf-
Schwenninger and Christina G. Martin (SMNS) for the SEM images; to Carlo M. Cunha (Unimes), Claude Vil-
vens (MNHN) and Bruce Marshall (NMNZ) for several helpful suggestions and corrections; and to the following
people for providing photos and information of type specimens: Bárbara L. Valentas-Romera, Leonardo Souza
(MNRJ), Emmanuel Tardy (MHNG), Adam Baldinger and Jennifer Lenihan (MCZ), Ellen Strong and Thomas
Jorstad (USNM). RBS acknowledges the bequest of Bruce Fraser Hazelwood and the NMNZ.
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