THE NAUTILUS
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Volume 130, Number 2
June 15, 2016
ISSN 0028-1344
A quarterly devoted
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THE
CONTENTS
Abraham S.H. Breure
Ad Hovestadt
Angela Fields
David G. Robinson
Vanessa Simao do Amaral
Luiz Ricardo L. Simone
Angel Valdes
Sabrina Medrano
Vishal Bhave
Emilio F. Garcia
N A U T I
L U S
Volume 130, Number 2
June 15, 2016
ISSN 0028-1344
The land Mollusca (Gastropoda) of Saint Kitts and Nevis
(Lesser Antilles), with description of a new species . 27
Comparative anatomy of five species of Saccostrea Dollfus and
Dautzenberg, 1920 (Bivalvia: Ostreidae) from the
Pacific Ocean . 53
A new species of Cuthona Alder and Hancock, 1855
(Gastropoda: Heterobranchia: Nudibranchia: Tergipedidae)
from the Caribbean Sea . 72
The genera Miraclathurella Woodring, 1928 (Gastropoda:
Pseudomelatomidae) and Ddrrylia Garcia, 2008 (Gastropoda:
PHoraiclavidae), with two proposed new combinations
for Darrylia . . .
79
THE NAUTILUS 130(2):27-52, 2016
Page 27
The land Mollusc a (Gastropoda) of Saint Kitts and Nevis
(Lesser Antilles), with description of a new species
Abraham S.H. Breure
Naturalis Biodiversity Center
P.O.Box 9517
NL-2300RA Leiden, THE NETHERLANDS
and
Royal Belgian Institute of Natural Sciences
Vautierstraat 29,
B-1000, Brussels, BELGIUM
[email protected]
Angela Fields
Dept. Biological and Chemical Sciences
University of the West Indies
Cave Hill Campus, BARBADOS
[email protected]
Ad Hovestadt
Dr. Abraham Kuyperlaan 22
NL-3818JC Amersfoort, THE NETHERLANDS
ad .h oves tadt @xs4 all.nl
David G. Robinson
USDA APHIS National Malacology Laboratory
Academy of Natural Sciences of Philadelphia
1900 Benjamin Franklin Parkway
Philadelphia, PA 19103 USA
[email protected]
ABSTRACT
An overview of the land snail fauna of the Lesser Antillean
islands of Saint Kitts and Nevis is given, based on data from
literature and two recent surveys. There are 33 taxa listed, of
which 26 are from Saint Kitts and 22 from Nevis. One taxon is
described as new: Bulimulus ouallensis Breure and Hovestadt.
Furthermore, the following taxa are recorded from these
islands for the first time: Bulimulus diaphanus fraterculus
(Potiez and Michaud, 1835), Obeliscus 'swiftianus (Pfeiffer, 1854),
and Zonit aides arboreus (Say, 1817). Four taxa — Diplosolenodes
sp., Pallifera sp., and two Succinea species — could only be
identified to the genus level. Three taxa, previously thought to
occur on the islands, are now removed from their faunal lists,
due to inaccuracies of provenance of specimens or misidentifi-
cations. Finally, remarks are given on the distribution and con¬
servation status of species collected during the surveys.
Additional Keywords : Taxonomy, distribution, islands, West Indies
INTRODUCTION
The Reverend Smith stands in the time-honored tradition
as a parson-naturalist (Armstrong, 1990). As a clergyman,
he spent five years on Nevis, summarizing his findings in a
1745 book entitled: “A natural history of Nevis and the
rest of the english Leeward Charibee Islands in America
with many other observations on nature and art; particulary
an introduction to the art of decyphering in eleven
letters etc.’ He begins with observations on marine
shells, but there is no reference whatsoever to the non¬
marine malacofauna.
The first overview of the land shells from Saint Kitts
and Nevis was published in 1862 by Thomas Bland. For
Saint Christopher, the oldest English name given to the
island of Saint Kitts, he listed six species in his ‘Catalogue
of the Land Shells of the West Indian Islands’; Nevis was
not mentioned at all. Bland (1862: 361) wrote “[t]he names
of several islands do not appear in the Catalogue, because
I am ignorant as to the species inhabiting them”.
The first mention of an endemic species for either
Saint Kitts or Nevis was by Henry A. Pilsbry, who in 1889
described Helix josephinae var. nevisensis [now Pleurodonte
josephinae nevisensis]; the specimen had been collected by
Frederick Albion Ober. The latter being a very prolific
writer and traveler, he regrettably did not mention his
activities in Nevis in any detail.
William H. Rush visited Saint Kitts in 1891. His methods
are still useful today: “Carriage hire was too expensive and
the time at my disposal too short for any extended trip, so
footing it was resorted to as the only way to reach reason¬
ably near hunting grounds. In this manner the deep gorges
up in the mountains of St. Kitts, at an elevation of one
or two thousand feet were visited (....) Many specimens
of Bulimulus, Helicina and Amphibulima were taken”.
Pieter Wagenaar Hummelinck visited Saint Kitts and
Nevis twice, in 1949 and 1955, but apparently restricted
his collecting to the coastal areas (Figure 4). His surveys
provided the raw data for publications on the Vertiginidae
(Haas, 1960), Subulinidae and Qleacinidae (Haas, 1962),
and Bulimulidae (Breure, 1974).
Page 28
THE NAUTILUS, Vol 130, No. 2
Figure 1. West Indies, showing location of Saint Kitts and Nevis (red circle). Source: Wikimedia.
Saint Kitts and Nevis is a federative, two-island state in
the West Indies (Figures 1-3), and constitutes part of the
British Commonwealth. The total area is 269 km2, of
which Saints Kitts acounts for 176 km2 and Nevis 93 km2.
Figure 2. Saint Kitts. Localities sampled in 2004 (yellow pins),
respectively 2014 (green pins); scale = 2.5 km. Detail of area
around Brimstone Hill within red quadrant; scale = 250 m.
Source: Google™ Earth Pro. Image Landsat. © 2015 Google.
The highest elevation on Saint Kitts is Mount Lianrmiga
(1156 m, also known as Mount Misery), and on Nevis it is
Nevis Peak (985 m). Annual average rainfall is 1625 mm
and 1170 min respectively. On Saint Kitts, the rainfall
is fairly evenly distributed over the island except for
the Southeast Peninsula, which has a very dry climate.
Fi gure 3. Nevis, with localities sampled in 2004 (yellow pins),
respectively 2014 (green pins); scale =1.0 km. Source: Google™
Earth Pro. Image Landsat. © 2015 Google.
A.S.H. Breure et al., 2016
Page 29
From August to November there is a relatively wet season
and a drier season from mid-January to April. On Nevis,
rainfall is lowest on the eastern side and increases widi
altitude. Most rain falls between July and January (all data
from Anonymous, 1991).
Geologically both islands belong to the western chain
of the Lesser Antillean archipelago north of Dominica,
which includes the islands of Basse Terre (Guadeloupe),
Montserrat, Redonda, Nevis, Saint Kitts, Saint Eustatius,
and Saba. These islands consist mostly of volcanic rocks
and remain seismically active, but geological knowledge
about them remains based mainly on older sources. Saint
Kitts has a core of Eocene andesite, and three younger
volcanic centers which were active during the Pleistocene.
Mt. Liamuiga is a stratovolcano with a deep summit crater
of over 300 m diameter and, together with the other
volcanic centers, are aligned along the length of the island
on a NW-SE trend. A small area of Plio-Pleistocene lime¬
stone can be found at Brimstone Hill and at Godwin
Ghaut above 330 m (Trechmann, 1932; Martin-Kaye, 1959;
Westermann and Kiel, 1961; Anonymous, 1991; Toothill
et al., 2007). The oldest rocks of Nevis are of marine
origin, but the island is comprised mainly of dacites, with
andesites in a number of places. Limestone can be found
on the southern slopes of Saddle Hill (Westermann and
Kiel, 1961; Hutton, 1968; Hutton and Nockolds, 1978).
Table 1. Localities mentioned in this paper.
Page 30
THE NAUTILUS, Vol 130, No. 2
Vegetation types occurring on the two islands have been
extensively described by Lindsay and Horwith (1999) based
on a modification of Beard (1949). Helmer et al. (2008) com¬
pared tire data from Beard ( 1949) with recent data and calcu¬
lated the change in land cover data based on satellite imagery.
The aim of this paper is to present the combined
results of two malacological surveys, respectively con¬
ducted in 2004 (by D.G. Robinson and collaborators)
and 2014 (by A. Hovestadt), and to summarize the
malacofauna of these islands.
MATERIALS AND METHODS
Both above-mentioned, recent surveys yielded data on
40 localities in total. Localities are presented with their
coordinates as they were recorded by D.G. Robinson and
collaborators (field work during 2004) and A. Hovestadt
(field work during 2014) respectively (Table 1). Altitudes
for the former have been taken from Google Earth
v.7. 1.2.2041. Coordinates of these localities were plotted
in SimpIeMappr (Shorthouse, 2010) and exported as KM i,
file. For historical comparisons, the localities sampled by P.
Wagenaar Hummelick (1949) have been added in Table 1;
coordinates of these (estimated from Google Earth) are
based on his map (Figure 4; Wagenaar Hummelinck,
1953: 20-21, Figure 19), and are less precise than the
recent ones. The species occurrence data for these locali¬
ties are only partial (Haas 1960, 1962; Breure, 1974).
The above genus-levels taxonomy follows Bouchet et al.
(2005). Under each species, references are made only to
the literature including records from Saint Kitts and Nevis.
The diversity of sampled localities was analyzed to
determine “hotspots” of land snail diversity on the two
Figure 4. Saint Kitts and Nevis, localities sampled in 1949
(Wagenaar Hummelinck, 1953: fig. 16).
islands, following the same method as applied by Robinson
et al. (2009) for Dominica. When more species are present
in a given locality, also rare species will be better repre¬
sented. Therefore, each occurrence was given a “rareness
factor” (R = 1/L; L equals the number of localities where
a species is present). R varies in this study from 0.025 (the
species occurs at all 40 localities where molluscs have
been collected) to 1.000 (the species occurs at a single
locality only). Finally, the diversity per locality is calcu¬
lated, both as total and for endemic species only (Dtot =
E Ktot/S; Dend = Z Rgnj/S; S, species richness as the
number of species per locality). Localities sampled by
Wagenaar Hummelinck, with partial species occurrence
data, have been excluded from these calculations.
Abbreviations for depository collections are: AH, col¬
lection of A. Hovestadt, Amersfoort, the Netherlands;
ANSP, Academy of Natural Sciences of Philadelphia,
Philadelphia, USA; FMNH, Field Museum of Natural
History, Chicago, USA; KMNH, Naturalis Biodiversity
Center (fonnerly Rijksmuseum van Natuurlijke Histone),
Leiden, the Netherlands; SMF, Senckenberg Natur-
Museum, Frankfurt am Main, Germany; UF, Florida
State Museum of Natural History, Gainesville, U.S.A.;
USDA, United States Department of Agriculture, Animal
and Plant Health National Malacology Collection, ANSP,
Philadelphia, USA. Other abbreviations are: leg., for “col¬
lected by,” and “coll.” for “Collection.” An asterisk (*)
indicates an observation only, material not collected.
SYSTEMATICS
Superfamily Helieinoidea sensu Thompson, 1980
Family Helicinidae Ferussac, 1822
Genus Helicina Lamarck, 1799
Helicina fasciata Lamarck, 1822
(Figures 20, 22-23)
Helicina fasciata Lamarck, 1822. — Bland, 1862: 358
(St. Christopher); Clench, 1956: 69 (St. Kitts, Nevis).
Survey Material: St. Kitts: Christchurch, Nichola
Town Parish, trail to Phillips Level, (USDA); St. Thomas,
Middle Island Parish, trail to D’Os Dane Pond (USDA);
Trinity, Palmetto Point Parish, top of Ottley’s Level
(USDA); ibid., near base of Ottley’s Level (USDA); Nevis:
St. George, Gingerland Parish, Golden Rock (AH; USDA);
ibid., Herbert Heights, trail to Nevis Peak (USDA); ibid.,
near Peak Heaven (AH); St. James, Windward Parish,
above Prison Farm, cloud forest (AH; USDA).
Additional Material Examined: St. Kitts: ex Swift
coll (ANSP 14914); W.H. Rush leg., 1891 (ANSP
62064). Nevis: C.A. Barber leg. (ANSP 78304); ibid.,
J. Bond leg., 1929 (ANSP 149430).
Distribution: Lesser Antilles, where it is widespread
as a result of human activities.
A.S.H. Breure et al, 2016
Page 31
Figures 5-19 Helicinidae, Suceineidae, Vertiginidae, and Subulinidae species. 5-7. Lucidella (Poeniella) plicatula christophori
(Pilsbry, 1897), H = 4.2 mm. 8. Succinea species A, H = 9.3 mm. 9-10. Succinea species B, H = 11.3 min. 11. Gastrocopta rupicola
nwrginalha (L. Pfeiffer, 1840), H = 2.2 mm. 12. Gastrocopta servilis (Gould, 1843), H 2.3 mm. 13. Pupoides marginatus nitidulus
(L. Pfeiffer, 1839), H = 3.6 mm. 14. Leptinaria unilamellata (d’Orbigny, 1837), H = 10.6 mm. 15. Allopeas gracile (Hutton, 1834),
H = 7.0 mm. 16. Allopeas micra (d’Orbigny, 1835), H = 6.0 mm. 17. Obeliscus swift ianus (L. Pfeiffer, 1854), H = 7.2 mm.
18. Subulina octona (Bruguiere, 1789), H = 19.7 mm. 19. Beckianum beckianum (L. Pfeiffer, 1846), H = 6.9 mm.
Habitat: Occurs in a wide range of habitats, from rel¬
atively undisturbed (cloud forest) to disturbed (area with
Heliconia plants).
Remarks: This species exhibits a marked preference
for humid conditions in dense vegetation (e.g., humid
scrub forest, cloud forest), being particularly abundant
on the upper slopes of Nevis Peak. It shows a high variety
of colors, even in specimens within the same population.
Genus Lucidella Swainson, 1840
Subgenus Poeniella H.B. Baker, 1923
Lucidella (Poeniella) plicatula christophori
(Pilsbry, 1897)
(Figures 5-7, 21)
Helicina plicatula var. christophori Pilsbry, 1897: 118
(Type locality: St. Kitts).
Page 32
THE NAUTILUS, Vol. 130, No. 2
Figures 20-21. Distribution of Helieinidae. 20. Helicina fasciata Lamarck, 1822 (red star). 21. Lucidella (Poenilla) plicatula
christophori (Pilsbry, 1897) (blue triangle).
Helicina christophori Pilsbry, 1897. — H.B. Baker,
1923: 23.
Lucidella plicatula christophori (Pilsbry, 1897). — Boss
and Jacobson, 1974: 31.
Survey Material: St. Kitts: Trinity, Palmetto Point
Parish, top of Ottley’s Level (USDA); ibid., near base of
Ottley’s Level (USDA); Nevis: St. James, Windward
Parish, above Prison Farm, cloud forest (USDA); ibid.,
Camp’s River (USDA); St. George, Gingerland Parish,
Saddle Hill (AH).
Additional Material Examined: St. Kitts: W.H.
Rush leg. (ANSP 62062, lectotype); ibid. (ANSP 358494,
paralectotypes).
Distribution: Saint Kitts, Nevis.
Habitat: It is believed to live in damp leaf litter and
under rotting logs.
Remarks: This species appears to be relatively rare,
as we found only empty shells.
Superfamily Veronicelloidea Gray, 1840
Family Veronicellidae Gray, 1840
Genus Dvplosolenodes Thome, 1975
Diplosolenode § species
(Figure 27, 31)
Survey Material: Nevis: St. George, Gingerland
Parish, Herbert Heights, trail to Nevis Peak (USDA).
Remarks: There is insufficient material to determine
if this is a more widespread species occurring on other
Antillean islands.
Genus Veronicella de Blainville, 1817
Veronicella cuhensis (L. Pfeiffer, 1840)
(Figures 26, 31)
Survey Material: Saint Kitts: St. Thomas, Middle
Island Parish, fortifications midway up Brimstone Hill
(USDA); Trinity, Palmetto Point Parish, Ottley Plantation
House (AH*); Nevis: St. George, Gingerland Parish,
Golden Rock (USDA).
Distribution: Cuba, Hispaniola, Puerto Rico, Antigua,
Saint Kitts, Nevis, Dominica, Barbados. Introduced to
various Pacific islands (Hawaiian Islands, Guam, northern
Marianas).
Remarks: Robinson et al. (2009) mentioned this spe¬
cies from Saint Kitts and Nevis, without providing fur¬
ther evidence. Although it has been reported as a serious
agricultural pest (especially on Pacific islands), this
species is of no such concern at the moment on these
two islands.
Veronicella aff. floridana (Leidy in Rinney, 1851)
(Figures 25, 31)
Survey Material: Saint Kitts: Trinity, Palmetto Point
Parish, nursery of Mattingly Flower Farm (USDA); ibid.,
top of Ottley’s Level (USDA); Nevis: St. John, Figtree
Parish, Montpellier Estate, botanical garden (USDA).
Distribution: Florida, throughout the West Indies.
Remarks: This invasive slug species is here reported
from Saint Kitts and Nevis for the first time. It has
become an agricultural pest in other parts of the West
Indies, where it feeds on a wide range of species. Dissec¬
tion is required to accurately identify tills and the previous
species, as the two can bear some morphological similarity.
Superfamily Succineoidea Beck, 1837
Family Succineidae Beck, 1837
A.S.H. Breure et al., 2016
Page 33
Figures 22-30. Living snails of the families Helicinidae, Philomycidae, Veronicellidae, Streptaxidae, Subulinidae, and
Pleurodontidae. 22-23. Helicina fasciata Lamarck, 1822. 24. Poll if era species. 25. Veronicella aff. floridana (Leidy in Binney,
1851). 26. Veronicella cubensis (L. Pfeiffer, 1840). 27. Diplosenodes species. 28. Streptartemon glaber (L. Pfeiffer, 1849).
29. Subulina octona (Bruguiere, 1789). 30. Pleurodonte josephinae nevisensis (Pilsbry, 1889).
Genus Succinea Drapamaud, 1801
Succinea species A
(Figures 8, 32)
Survey Material: Saint Kitts: St. George, Basseterre,
road south of Basseterre (USDA).
Remarks: The taxonomy of West Indian succinids is
poorly understood, and a revision is long overdue. Not
many anatomical or molecular studies have been made
on this group, and various names have been used indis¬
criminately by malacological workers over the centuries
for snails on different Antillean islands.
Succinea species B
(Figures 9, 10, 32)
Material Examind: Nevis: St. John, Figtree Parish,
Prospect, greenhouse of Experimental Station (USDA);
St. Thomas Lowland, greenhouse of Taiwanese technical
Mission (USDA).
Distribution: Florida, Puerto Rico, Nevis.
Remarks: This species was likely introduced with hor¬
ticultural imports from southern Florida or from Puerto
Rico, as it closely resembles material found in green¬
houses in those areas.
Superfamily Achatinoidea Swainson, 1840
Family Subulinidae P. Fischer and Crosse, 1877
Genus Allopeas H.B. Baker, 1935
Allopeas gracile (Hutton, 1834)
(Figures 15, 33)
Lamellaxis ( Allopeas ) gracile (Hutton [, 1834]). — Haas,
1962: 56; St. Christopher, Brimstone Hill; Nevis:
Jessops Village.
Page 34
THE NAUTILUS, Vol. 130, No. 2
Figures 31-32. Distribution of Veroniceilidae and Succineidae. 31. Diplosolenodes sp. (green star), Veronicella cubensis
(L. Pfeiffer, 1840) (orange rectangle), and Veronicella aff . floridensis (Leidy in Binney, 1851) (purple circle). 32. Succinea sp. A
(yellow triangle), Succinea sp. B (dark blue polygon).
SAINT KITTS
Survey Material: Saint Kitts: St. George, Basseterre
Parish, Basseterre, La Guerite, Department of Agricul¬
ture (USDA); ibid., road south of Basseterre (USDA);
St. Thomas, Middle Island Parish, base of Brimstone Hill
(USDA); ibid., near barracks (AH); Nevis: St. George,
Gingerland Parish, Saddle Hill (AH); ibid., Montravers
Estate (AH); St. John, Figtree Parish, Montpellier Estate,
botanical garden (USDA).
Distribution: West Indies, southern Mexico, Central
and South America; distributed throughout the subtrop¬
ics worldwide.
A llopeas micro (d’Orbigny, 1835)
(Figures 16, 34)
Lamellaxis (Allopeas) micron (d’Orbigny, [1835]). —
Haas, 1962: 57; St. Christopher, Brimstone Hill;
Morne Hills.
Survey Material: Saint Kitts: St. George, Basseterre
Parish, Basseterre, La Guerite, Department of Agricul¬
ture (USDA); ibid., road to Turtle Bay, south of Salt Pan
(USDA); ibid., road south of Basseterre (USDA);
St. Thomas, Middle Island Parish, lower slope of Brimstone
Hill (USDA); ibid., NW-side Brimstone Hill (AH); Trinity,
Palmetto Point Parish, near base of Ottleys Level (USDA);
Nevis: St. George, Gingerland Parish, Saddle Hill (AH).
Distribution: West Indies, Mexico to Bolivia.
Remarks: The specimens found are up to 6 mm in height.
Figures 33-34. Distribution of Subulinidae. 33. Allopeas gracile (Hutton, 1834) (red star), Beckianum beckianum (L. Pfeiffer, 1846)
(yellow rectangle), Leptinaria unilamellata (d’Orbigny, 1837) (green triangle). 34. Allopeas micro (d’Orbigny, 1835) (light blue star),
Obeliscus swiftianus (L. Pfeiffer, 1853) (orange circle), Subulina octona (Bruguiere, 1789) (purple polygon).
A.S.H. Breure et al., 2016
Page 35
Figures 35-39. Streptaxidae and Sagdidae species. 35. Huttonella bicolor (Hutton, 1834), H = 6.0 min. 36. Streptartemon glaber
(L. Pfeiffer, 1849), H 6.8 mm. 37-39. Hojeda species, D = 4.5 mm.
Genus Beckianum H.B. Baker, 1961
Beckianum beckianum (L. Pfeiffer, 1846)
(Figure 19, 33)
Diaopeas beakianum (L. Pfeiffer [, 1846]). — Haas,
1962: 55. St. Christopher, La Guerite; Nevis, near
Jessops Village.
Survey Material: Saint Kitts: St. Thomas, Middle
Island Parish, lower slope of Brimstone Hill (USDA); ibid,
base of Brimstone Hill (USDA); ibid., near barracks and
on NW side of Brimstone Hill (AH); Nevis: St. George,
Gingerland Parish, Montraves Estate (AH); St. John,
Figtree Parish, Saddle Hill (AH).
Distribution: West Indies, Central America.
Remarks: Material collected was up to 9 mm in height.
This is the first report of this taxon from Nevis.
Genus Leptinaria Beck, 1837
Leptinaria unilamellata (d’Orbigny, 1837)
(Figures 14, 33)
Survey Material: Saint Kitts: Trinity, Palmetto Point
Parish, top of Ottley’s Level (USDA). Nevis: St. George,
Gingerland Parish, Golden Rock (USDA); ibid.,
Frenchman’s Cave (USDA); Herbert Heights, trail to
Nevis Peak (USDA).
Figures 40-41. Distribution of Streptaxidae and Gastrocoptidae. 40. Huttonella bicolor (Hutton, 1834) (grey star), Gastrocopta
rupicola marginalba (L. Pfeiffer, 1840) (yellow circle). 41. Streptartemon glaber (L. Pfeiffer, 1849) (blue rectangle), Gastrocopta
servilis (Gould, 1843) (red triangle).
Page 36
THE NAUTILUS, VoL 130, No. 2
Distribution: West Indies, Central America to Venezuela
and Peru.
Habitat: Generally found in damp leaf litter and under
rotting logs.
Remarks: The specimens collected obtain a maximum
shell height of 15 mm. This species is particularly abun¬
dant in disturbed habitats, especially in agricultural areas.
Genus Obeliscus Reek, 1837
Subgenus Sfenogyra Shuttle worth, 1854
Obeliscus (Sienogyra) swiftianus (L. Pfeiffer, 1853)
(Figures 17, 34)
Opeas octogyrum plicatellum (Guppy [, 1868]). — Haas,
1962: 55; St. Christopher, Brimstone Hill.
Survey Material: Saint Kitts: St. George Basseterre
Parish, road to Turtle Bay, south of Salt Pan (USDA);
St. Thomas Middle Island Parish, lower slope of
Brimstone Hill (USDA).
Distribution: Puerto Rico and northeastern Lesser
Antilles.
Remarks: Haas (1962) misidentified this snail, using
the name of a related species from Trinidad; we believe
that his illustrated species is Allopeas gracile. This is the
first confirmed record for Saint Kitts.
Genus Subulina Reek, 1837
Subulina octona (Rruguiere, 1789)
(Figures 18, 29, 34)
Subulina octona (Bruguiere [, 1789]). — Haas, 1962: 49.
St. Christopher, Winfield River; Brimstone Hill;
Timothy Hill at Frigate Bay; Nevis, Jessops Village,
Nelsons Spring, Jones’ River.
Survey Material: Saint Kitts: St. George, Basseterre
Parish, road to Turtle Bay, south of Salt Pan (USDA);
St. John, Capisterre Parish, Saddler’s Land Settlement,
Lavington Ghut (USDA); St. Paul, Capisterre Parish,
trail head to Mount Liamuiga (AH); St. Thomas, Middle
Island Parish, lower slope of Brimstone Hill (USDA);
ibid., trail to D’Os Dane Pond, off Old Military trail
(USDA); Trinity, Palmetto Point Parish, top of Ottley’s
Level (USDA); ibid., near base of Ottleys Level (USDA);
Nevis: St. George, Gingerland Parish, along Source trail,
N of Golden Rock Inn (AH); St. John, Figtree Parish,
Saddle Hill (AH).
Distribution: Worldwide tropics and subtropics; in
temperate zones in greenhouses.
Remarks: This widely distributed species, probably
introduced centuries ago from Africa, is now one of the
commonest species in the tropics and subtropics. It Is
considered to be an Indicator of disturbed environments.
The species is now recorded for the first time from Nevis.
Superfamily Streptaxoidea Gray, 1860
Family Streptaxidae Gray, 1860
Genus Streptartemon Kohelt, 1905
Sireptartemon glaher (L. Pfeiffer, 1849)
(Figures 28, 36, 41)
Material; Saint Kitts: St. George, Basseterre Parish,
road to Turtle Bay, south of Salt Pan (USDA); St. John,
Capisterre Parish, crater trail up Mount Liamuiga
(USDA); St. Thomas, Middle Island Parish, lower slope
of Brimstone Hill (USDA); ibid., near barracks and NW
side of Brimstone Hill (AH); ibid., trail to D’Os Dane
Pond, off Old Military trail (USDA); Trinity, Palmetto
Point Parish, top of Ottleys Level (USDA); ibid., near
base of Ottleys Level (USDA).
Distribution: Puerto Rico, U.S. Virgin Islands, Saint
Thomas, Saint Croix, Saint Kitts, Dominica, Barbados,
Venezuela, Guyana, Surinam, Brazil.
Remarks: This South American carnivorous species has
been introduced throughout the Lesser Antilles by human
activity. It appears to be very widespread on Saint Kitts;
its effect on the native mollusks remains undocumented
as yet.
Genus Huttonella L, Pfeiffer, 1856
Huttonella bicolor (Hutton, 1834)
(Figures 35, 40)
Ennea (Huttonella) bicolor Hutton, 1834. — Tryon, 1885:
104; introduced to West Indies [no records for Saint
Kitts and Nevis].
Survey Material: Saint Kitts: St. George, Basseterre
Parish, Basseterre, La Guerite, Department of Agricul¬
ture (USDA).
Distribution: Africa; introduced into the tropics and
subtropics worldwide, including USA (Florida), West
Indies, Panama, Brazil.
Remarks: Specimens of this carnivorous species found
on Saint Kitts have a shell height of up to 6.75 mm. The
species might have been imported with potted plants.
This is the first record for this island.
Superfamily Gastrodontoidea Tryon, 1866
Family Gastrodontidae Tryon, 1866
Genus Xonitoides Lehmann, 1862
Z onitoides arboreus (Say, 1817)
Survey Material: Saint Kitts: Saint Thomas, Middle
Island Parish, NW side of Brimstone Hill (AH).
A.S.H. Breure et a!., 2016
Page 37
Figures 42-51. Amphibulimidae and Bulimulidae species. 42-^13. Amphibulima patula christopheri Pilsbry, 1902, H = 17.3 mm.
44-45. Drymaeus (Antidrymaeus) multifasciatus christopheri Pilsbry, 1899, H = 16.4 mm, respectively H = 14.6 mm. 46. Drymaeus
(Antidrymaeus) multifasciatus subspecies, H = 16.2 mm. 47. Bulimulus gittenbergeri Breure, 1974, holotype RMNH 54903,
H = 20.7. 48-49. Bulimulus guadalupensis (Bruguiere, 1789), H 17.6 mm, respectively H 17.4 mm. 50. Bulimulus ouallensis Breure
and Hovestadt new species, holotype RMNH 5003990, H = 19.0. 51. Bulimulus diaphanus fraterculus (Potiez and Michaud, 1835),
H = 16.5 mm.
Page 38
THE NAUTILUS, Vol. 130, No. 2
Table 2. Bulimulus ouallensis Breure and Hovestadt new
species. Measurements of the type material. Abbreviations: D,
diameter; H, shell height; HA, height of aperture; LW, height
of last whorl; P, number of protoconch whorls; W, total number
of whorls; WA, width of aperture (all in mm). See also Breure
(1974; figs 2-5).
Distribution: Widespread in North America, Bahamas,
Puerto Rico.
Remarks: This is the first record for Saint Kitts of this
widespread species, which is likely to have been introduced.
Superfamily Orthalicoidea Albers, 1860
Family Bulimulidae Tryon, 1867
Genus Bulimulus Leaeh, 1814
Remarks: The species of this genus are usually diffi¬
cult to differentiate, most of them being brownish and of
similar shape. The accurate identification of several spe¬
cies from these islands can only be achieved through
careful comparisons.
Bulimulus diaphanus fraterculus (Potiez and
Michaud, 1835)
(Figures 51-52)
Bulimus fraterculus T[erus$ae]| Potiez and Michaud,
1835".— Bland, 1862: 358; St. Kitts.
Figures 52-55. Distribution of Bulimulidae and Amphibulimidae. 52. Bulimulus gittenbergeri Breure, 1974 (purple triangle),
B. diaphanus fraterculus (Potiez and Michaud, 1835) (yellow triangle). 53. Bulimulus guadalupensis (Bruguiere, 1789) (orange
triangle). 54. Bnjmaeus multifasciatus christopheri Pilsbry, 1899 (green circle), D. multifasciatus subspecies (dark blue circle).
55. Bulimulus ouallensis Breure and Hovestadt new species (light blue triangle), Amphibulima patula christopheri Pilsbry, 1902
(dark red polygon).
A.S.H. Breure et a!., 2016
Page 39
Bulimulus fraterculus Potiez and Michaud, 1835. — Pilsbry,
1897 [1897-1898]: 46, pi. 11 figs 24-25 (excluding
Bland’s record).
Bulimulus diaphanus fraterculus Potiez and Michaud,
1835. — Breure, 1974: 32, pi. 3 figs 6-10; pi. 7 fig. 1;
reference to Bland’s record.
Survey Material: Nevis: St. John, Windward Parish,
cloud forest above Prison Farm (AH).
Distribution: Saint Martin, Saint Barts, Saba, Saint
Eustatius, Nevis, Barbuda, Antigua, Guadeloupe, Les
Saintes, Dominica.
Remarks: This taxon has been mentioned from Saint
Kitts by Bland (1862), but no voucher material could be
traced by Breure (1974). Pilsbry (1897 [1897-1898]: 46)
wrote: “The B. fraterculus of American collections,
reported from Porto Rico, St. Kitts, St. Croix, Antigua,
St. John, St. Thomas, Trinidad and Barbados, is not this
species; so the localities cited by Pfr. in Monogr. viii,
p. 189, and by Smith, Ann. Mag. (6), viii, on the authority
of Bland, do not refer to the true fraterculus” . Breure
(1974), after having found type material of Potiez and
Michaud at the Museum National d’Histoire Naturelle,
Paris, confirmed the presence of this taxon only on the
islands of Saint Martin, Saba, Saint Eustatius, Barbuda,
and Guadeloupe. This is the first record for Nevis.
Bulimulus guadalupensis (Bruguiere, 1789)
(Figures 48-49, 53)
Bulimus exilis Gm[elin], 1791. — Bland, 1862: 358;
St. Kitts.
Bulimulus exiles [sic] Gmelin, 1791. — Rush, 1891: 69.
Bulimulus exilis Gmelin, 1791: Pilsbry, 1897 [1897-1898]:
37, pi. 9 figs 61-67; St. Christopher or St. Kitts.
Bulimulus (Bulimulus) guadalupensis (Bruguiere,
1789). — Breure, 1974: 15, figs 6-60, pi. 2.; St. Kitts,
Timothy Hill at Frigate Bay; Frigate Bay beach;
between Jugate Bay and Basseterre; Phillips. Nevis.
Survey Material: Saint Kitts: St. George, Basseterre
Parish, Basseterre, La Guerite, Department of Agricul¬
ture (USDA); St. Thomas, Middle Island Parish, lower
slope of Brimstone Hill (USDA); Trinity, Palmetto Point
Parish, top of Ottleys Level (USDA); ibid., near base
of Ottleys Level (USDA); Nevis: St. James, Windward
Parish, above Prison Farm (USDA); ibid., Camp’s River
(USDA).
Additional Material Examined: Saint Kitts: T.
Bland leg., ex A.D. Brown (ANSP 3519); Dietz leg., ex
Swift coll. (ANSP 25574); C.A. Barber leg., ex T.D.A.
Cockerell (ANSP 78301).
Distribution: Probably originated in the Windward
Islands (Breure, 1974); now distributed throughout the
Caribbean Basin, including Florida.
Bulimulus gittenbergeri Breure, 1974
(Figures 47, 52)
Bulimulus gittenbergeri Breure, 1974. — 27, pi. 5 figs 10-13;
pi. 7 figs 3: St. Kitts, limestone NW Brimstone Hill.
Survey Material: Saint Kitts: St. George, Basseterre
Parish, road south of Basseterre (USDA); St. Thomas,
Middle Island Parish, lower slope of Brimstone Hill
(USDA); ibid.. Brimstone Hill (AH).
Additional Material Examined: Saint Kiits: NW
Brimstone Hill (RMNH 54903, holotype), RMNH
54904, FMNH 174171, SMF 225900 (paratypes); top of
Brimstone Hill (RMNH), Wingfield River (RMNH).
Distribution: Endemic to Saint Kitts.
Habitat: Occurring in arid scrub forests of the coastal
zone, where it was collected among shrubs.
Remarks: The specimens collected have a shell height
up to 20.5 mm.
Bulimulus ouallensis Breure and Hovestadt new species
(Figures 50, 55)
Diagnosis: A species of Bulimulus up to 20.0 mm, with
slightly convex sides and a rather thin shell, sculptured
with spiral striation on the epidermis, protoconch pit-
reticulated in its latter half.
Description: Shell up to 20.0 mm, 2.1 times as long as
wide; narrowly perforated, acute with slightly convex
sides; rather thin. Colour light brown when epidermis
present, apex purplish-brown. Surface opaque, epidermis
usually with very delicate spiral striae; teleoeoneh sculp¬
tured with inerassate growth striae, approximately 1 mm
apart; striae fuse into white tufts, giving the impression
of a narrow white band running just below suture.
Protoconch marked with oblique riblets running at an
angle of 45 degrees with regard to body axis, for first
whorl. Following half- whorl shows also riblets at an
angle of 115 degrees, making the surface pit-reticulated.
Whorls 6.5, slightly convex; ultimate whorl about 0.66
total height. Suture deepened. Aperture 0.42 shell
height, 1.6 times as long as wide; subovate, whitish
inside; apertural deviation five degrees. Peristome thin;
columellar margin slightly reflexed and dilated above.
Dimensions of holotype: shell height 19.0, diameter
8.9, height of aperture 8.4, width of aperture 4.5, height
of last whorl 13.0; 6.5 whorls.
Type Material: Holotype: RMNH 5003990, A.
Hovestadt leg., 17 November 2014, from type locality;
Paratypes: three shells, St. John Windward Parish,
above Prison Farm.
Type Locality: Nevis, St. George, Gingerland Parish,
along Source trail. North Golden Rock Inn.
Distribution: Endemic to Nevis.
Page 40
THE NAUTILUS, Vol. 130, No. 2
Figures 56-82. Living snails of the family Bulimulidae. 56-60. Drymaeus (Antidmjmaeus) multifasciatus christopheri Pilsbry,
1899; 61-62. Drymaeus (Antidrymaeus) multifasciatus subspecies.
Habitat: Living specimens have been observed
crawling on the ground among fallen leaves and climbing
on tree trunks, In the wet rainforest.
Remarks: Bulimulus ouallensis may be compared to
the neighboring species from Saint Kitts, Bulimulus
gittenhergeri Breure, 1974, from which the new taxon
differs by (1) having the sides slightly convex; (2) the
light brown color. This new species may also be compared
to Bulimulus lehmanni (L. Pfeiffer, 1865), occurring on
Dog Island, Anguilla and Saint Martin-Sint Maarten,
but the latter species has definitely coarser growth striae,
lacks the pit-reticulated surface of the protoconch and
lacks the sutural tufts; the shell is also more solid. The
key to Caribbean Bulimulus species (Breure, 1974: 11-12)
may be adapted as follows:
13a. Sides straight . . . 14
13b. Sides slightly convex ............. ,B. ouallensis
14a. Colour yellowish to greyish white; aperture white
inside. . . B. gittenhergeri
14b. Colour pale russetbrown; inside aperture coloured
like outside . . . B. fuscus
Etymology: The specific epithet derives from Oualla,
the pre-Colombian name for the island, meaning “land
of the beautiful waters”, which presumably refers to
the many water sources and the hot volcanic springs on
the island.
Genus Drymaeus Albers, 1850
Subgenus Antidrymaeus L, Germain, 1907
Remarks: In a forthcoming paper, Breure and Robinson
will show that several Drymaeus species from the Carib¬
bean Basin form a monophyletic group, for which the
name Antidrymaeus L. Germain, 1907 may be used.
The common characteristic in the external morphology
of this group is the bluish color of the body, most notice¬
able in juvenile specimens.
Drymaeus ( Antidrymaeus ) multifasciatus christopheri
Pilsbry, 1899 new combination
(Figures 54, 56-60)
Bulimus multifasciatus Lamjarck], 1822. — Bland, 1862:
358. St. Kitts.
Bulimulus multifasciatus Lamarck, 1822. — Rush, 1891:
69. St. Kitts.
Drymaeus multifasciatus van christopheri Pilsbry,
1899. — 16, pi. 13 figs 98-99. St. Christopher.
A.S.H. Breure et al., 2016
Page 4 1
Drymaeus multifasciatus christopheri Pilsbry, 1899. —
Clench and Turner, 1962: 31.
Drymaeus multifasciatus christopheri Pilsbry, 1899. — H.B.
Baker, 1963: 227 (leetotype designabon).
Survey Material: Saint Kitts: Christchurch, Nichola
Town Parish, trail to Phillips Level (USDA); St. Paul,
Capisterre Parish, crater trail up Mount Liainuiga
(USDA); Trinity, Palmetto Point Parish, trail to Ottley’s
Level (AH).
Additional Material Examined: Saint Kitts: ex Swift
coll. (ANSP 25857, leetotype), (ANSP 325063, paralecto-
tvpes);T. Bland, ex A.D. Brown (ANSP 3417); (UF 176958).
Distribution: Endemic to Saint Kitts.
Habitat: Occurs in rainforest.
Drymaeus ( Antidry maeus) multifasciatus subspecies
(Figures 54, 61-62)
Survey Material: Nevis: St. George, Gingerland Parish,
Herbert Heights, trail up Nevis peak past Rawlins
Entrance, D.G. Robinson leg., 17 March 2004 (USDA);
ibid., near Peak Heaven (AH*); St. James, Windward
Parish, above Prison Farm, D.G. Robinson leg, 12 March
2004 (USDA); ibid. (AH*).
Distribution: Endemic to Nevis.
Habitat: Occurring in cloud forest and rain forest.
Remarks: This subspecies differs from the previous
subspecies by the far less brightly-colored adult shell,
instead showing dull bands of reddish-brown and yellow,
which have a tendency to fade away into the background
color; the patch around the umbilicus is also less colored
than the material from Saint Kitts.
Family Ainphibulimidae P. Fischer, 1873
Genus Amphibulima Lamarck, 1805
Amphibulima patula christopheri Pilsbry, 1902
(Figures 42-43)
Succinea patula Rrug[uiere], 1789. — Bland, 1862: 358.
St. Kitts.
Amphibulima patula Bruguiere, 1789. — Rush, 1891: 69.
Amphibulima patula Bruguiere, 1789. — Pilsbry, 1899:
234, pi. 61 figs 14-19. St. Kitts, Bayford Estate.
Amphibulima patula var. christopheri Pilsbiy, 1902. — Ixviii,
pi. 60 figs 11-12; pi. 62, figs 27-30. St. Kitts.
Amphibulima patula christopheri Pilsbry, 1902. — Clench
and Turner, 1962: 31.
Amphibulima patula christopheri Pilsbry, 1902. — H.B.
Baker, 1963: 227 (leetotype designation).
Amphibidima patula christopheri Pilsbry, 1902. — Breure,
1973: 53.
Survey Material: Saint Kitts: St. John Capisterre
Parish, crater trail up Mount Liamuiga (USDA); St. Thomas
Middle Island Parish, trail to D’Os Dane Pond, off Old
Military trail (USDA); Trinity Palmetto Point Parish, top of
Ottley’s Level (USDA); Nevis: St. James Windward Parish,
above Prison Farm (USDA).
Distribution: Saint Kitts, Nevis.
Habitat: This semi-slug seems to be restricted to rain
and cloud forests.
Remarks: This is the first report for this taxon from
Nevis. Another subspecies, from Dominica, has been
reported to frequent banana and Citrus trees (Robinson
et al., 2009: 638).
Superfamily Pupilloidea Turton, 1831
Family Gastrocoptidae Pilsbry, 1918
Genus Gastrocopta Wollaston, 1878
Gastrocopta harbadensis (L. Pfeiffer, 1854)
Gastrocopta ( Gastrocopta ) harbadensis harbadensis
(L. Pfeiffer [, 1854]).— Haas 1960: 6, pi. 2 figs A-F.
St. Christopher, Brimstone Hill. Nevis, Jessops Village;
Jones’ River.
Survey Material: Not collected during these surveys.
Additional Material Examined: Saint Kitts: Brimstone
Hill (RMNH). Nevis: Jessops Village (RMNH); Jones’
River (RMNH).
Distribution: West Indies.
Gastrocopta rupicola marginalba (L. Pfeiffer, 1840)
(Figures 11, 40)
Gastrocopta (Gastrocopta) rupicola marginalba (L.
Pfeiffer [, 1840]).— Haas 1960: 12, pi. 2, fig. K; pi. 5
figs A-D. St. Christopher, Timothy Hill; Frigate Bay;
Brimstone Hill.
Survey Material: Saint Kitts: St. Thomas, Middle
Island Parish, base of Brimstone Hill (USDA).
Distribution. Saint Thomas, Saint Croix, Dog Island,
Saint Martin, Saint Kitts, southern United States and
northern Mexico.
Gastrocopta servilis (Gould, 1843)
(Figures 12, 41)
Gastrocopta (Gastrocopta) servilis servilis (Gould
[, 1843]). — Haas, 1960: • 1. pi. Digs F-H. St. Christopher,
Morne Hills.
Survey Material: Saint Kitts: St. George Basseterre
Parish, road to Turtle Beach, south of Salt Pan (USDA);
St. Thomas Middle Island Parish, base of Brimstone
Hill (USDA).
Distribution: West Indies, Central America, Venezuela.
Page 42
THE NAUTILUS, Vol. 130, No. 2
Figures 63-66. Distribution Pupillidae, Sagdidae, Philomycidae, Gastrodontidae, Polygyridae and Pleurodontidae species.
63. Pupoides marginatus nitidus (L. Pfeiffer, 1839) (yellow polygon), Hojeda sp. (brown star), Pallifera sp. (purple triangle).
64. Pleurodonte guadeloupensis ssp. (green circle), P. josephinae nevisensis (Pilsbry, 1889) (orange circle). 65. Z achnjsia provisoria
(L. Pfeiffer, 1858) (red rectangle). 66. Polygijra p. plana (Dunker in Philippi, 1843) (light blue triangle), Z onitoides arboreus
(Say, 1817) (dark blue polygon).
Family Pupillidae Turton, 1831
Genus Pupoides L. Pfeiffer, 1854
Pupoides marginatus nitidulus (L. Pfeiffer, 1839)
(Figures 13, 63)
Pupoides ( Pupoides ) marginatus nitidulus (L. Pfeiffer
[, 1839]). — Haas, 1960: 5, pi. 1 Figure D. St. Christopher,
Brimstone Hill; Nevis, Mosquito Bay.
Survey Material; Saint Kitts: St. Thomas, Middle
Island Parish, base of Brimstone Hill (USDA).
Distribution: West Indies.
Superfamily Sagdoidea Pilsbry, 1895
Family Sagdidae Pilsbry, 1895
Genus Hojeda H.B. Baker, 1926
Hojeda species
(Figures 37-39)
Survey Material: Saint Kitts: St. George, Basseterre
Parish, road to Turtle Beach, south of Salt Pan (USDA);
ibid., road south of Basseterre (USDA); St. Thomas, Middle
Island Parish, lower slope of Brimstone Hill (USDA);
Nevis: St. George, Gingerland Parish, Montravers Estate
(AH); St. John, Figtree Parish, Saddle Hill (AH); St. Paul,
Charlestown Parish, Hamilton Estate ruins (AH).
Distribution; Guadeloupe, Saint Kitts, Nevis, Saint
Eiistatius.
Habitat: Found in xerophytie environments on
Saint Kitts.
A.S.H. Breure et al., 2016
Page 43
Figures 67-72. Pleurodontidae species. 67-69. Pleurodonte guadeloupensis subspecies, D 14.1 nun. 70-72. Pleurodonte
josephinae nevisensis (Pilsbry, 1889), lectotype ANSP 32590, D 17.7 mm.
Remarks: This small sagid (up to 4.5 mm in diameter)
closely resembles the Cuban Hojeda boothiana
(L. Pfeiffer, 1839), but differs by its lower spire, which
gives a flatter appearance to the shell. Hojeda vannattai
H.B. Baker, 1924 from Aruba, and Margarita Island (off
the Venezuelan coast) is also very similar but has con¬
sistently a somewhat larger umbilicus. Specimens from
Saint Eustatius, collected during the late 18th century
(ANSP 28312), are virtually identical; similar shells
have also been found on Guadeloupe (Robinson, unpub¬
lished data).
Superfamily Helicoidea Rafinesque, 1815
Family Polygyridae Pilsbry, 1895
Genus Polygyra Say, 1818
Polygyra plana plana (Dunker in Philippi, 1843)
(Figure 66)
Survey Material: Saint Kitts: Gardens of the Sugar
Bay Club, near North Frigate Bay (AH).
Distribution: Bermuda, Saint Kitts.
Habitat: This is an introduced species, occurring in
well-watered gardens.
Remarks: This is the first record for this taxon from
Saint Kitts and Nevis. The nominal subspecies occurs on
Bermuda. Another subspecies, Polygyra plana bahamensis
Vanatta, 1919, can be found in the Bahamas and the
Turks and Caicos Islands.
Superfamily Arionoidea Gray, 1840
Family Philomycidae Gray, 1847
Genus Pallifera Morse, 1864
Pallifera species
(Figure 24)
Survey Material: Nevis: St. George, Gingerland
Parish, Herbert Heights, trail to Nevis Peak (USDA).
Distribution: Nevis; USA.
Remarks: This species lias been reported from
Montserrat (Shoobs and Coote, 2014, as Pallifera dorsalis
(Binney, 1842)) and Martinique and Guadeloupe
(Delannoye et al., 2015, as P. spec.). This is the first record
for Saint Kitts and Nevis. It is most likely introduced
from elsewhere; until its identity can be ascertained, it is
not possible to determine its origin at present.
Page 44
THE NAUTILUS, Vol. 130, No. 2
Dtot
0.20 - 0.39
• 0.40 - 0.69
• >0.70
NEVIS
Dend
# <0.10
# 0.10 • 0.15
• >0.15
NEVIS
Figures 73-76. Diversity of land snails on Saint Kitts and Nevis. 73. Species richness per locality. 74. Frequency of species
richness. 75. Total diversity; calculated using rareness (see methods), only localities scoring 0.20 or higher shown. 76. Diversity of
endemic species (see methods).
Superfamily Helicoidea Rafinesque, 1815
Family Pleurodontidae von Ihering, 1912
Genus Pleurodonte Fischer von Waldheim, 1807
Pleurodonte guadeloupensis subspecies
(Figures 64, 67-69)
Survey Material: Saint Kitts: Trinity Palmetto Point
Parish, top of Ottley’s Level (USDA); ibid., trail to
Ottley’s Level (AH).
Distribution: Saint Kitts.
Remarks: This snail belongs to the Pleurodonte
guadeloupensis species complex of Guadeloupe, Dominica,
Saint Martin, and Martinique. It differs from the other
subspecies in this complex in being higher-spired, with
slightly different labial dentition.
Pleurodonte josephinae nevisensis (Pilsbry, 1889)
(Figures 30, 64, 70-72)
Helix josephinae Fer[ussac], 1832. — Bland 1862: 358;
St. Christopher.
Helix ( Dentellaria ) josephinae Ferussac, 1832. — Pilsbry
1889: 88, pi. 29 figs 31-33; Various islands, including
subfossils from St. Kitts and Nevis.
Helix josephinae var. nevisensis Pilsbry 1889. — 89, pi. 25
figs 54-55; Nevis.
Helix (Dentellaria) josephinae nevisensis Pilsbry, 1899. —
Clench and Tuner 1962: 104 [“no locality given”; sic].
Helix josephinae nevisensis Pilsbry, 1899. — H.B. Baker,
1963: 246.
Survey Material: Nevis: St. George, Gingerland Parish,
Herbert Heights, trail to Nevis Peak (USDA); ibid., near
Peak Heaven (AH); ibid., along Source trail, N Golden
Rock Inn (AH); St. James, Windward Parish, above Prison
Farm (AH, USDA).
A.S.H. Breure et al, 2016
Page 45
□
Drought Deciduous Open Woodland
Drought Deciduous Coastal Shrubland
Drought Semi-Deciduous Forest
Seasonal Evergreen Forest
Evergreen Forest (including Sierra Palm Forest)
Figure 77. Land cover and forest formations on Saint Kitts and Nevis, 1999-2003, with distribution of endemic species. Modified
after I leaner et al. 2006; grid lines refer to the UTM system. Triangles refer to the localities where endemic species were found (light
color, one species; dark colored, two species). Enlarged parts of the islands indicated with corresponding colors. Scale = 4 km.
Page 46
THE NAUTILUS, Vol. 130, No. 2
Additional Material Examined: Saint Kitts: T. Bland
leg., ex A.D. Brown (ANSP 1052, subfossil); ex Swift coll.
(ANSP 32591, subfossil); J.S. Phillips leg. (ANSP 30907;
see remarks); Nevis: T. Bland leg. (ANSP 32590, syntype);
A.D. Brown leg. (ANSP 110428).
Distribution: Saint Kitts?, Nevis.
Habitat: This species exhibits a marked preference for
humid conditions in dense vegetation, being particularly
abundant on the upper slopes of News Peak.
Remarks: The more widespread Pleurodonte josephinae
(Ferussac), occurring on Guadeloupe, and Dominica, has
been reported to occur on both Saint Kitts and Nevis, based
on subfossil specimens. However, Pilsbry (1889) recognized
the population still living on Nevis its a distinct subspecies;
he may have been unaware that the subfossils also belong to
this taxon. Two specimens (ANSP 30907) were collected
early 20th century on Saint Kitts. One, a fully developed
adult, appears to have a higher spire than any of the Nevis-
ian material we have seen. These shells may represent a
surviving Kittitian population that we were unable to locate.
Genus Zachrysia Pilsbry, 1894
Zachrysia provisoria (L. Pfeiffer, 1858)
(Figure 65)
Survey Material: Saint Kitts: St. Thomas, Middleland
Parish, near Romney Manor (AH); Trinity Palmetto Point
Parish, Ottleys Plantation House (AH); Nevis: St. George,
Gingerland Parish, near Peak Heaven (AH); ibid.,
Montravers Estate (AH*); St. John, Figtree Parish,
Prospect, Experimental Station (USDA).
Distribution: Cuba; introduced to Florida, the Bahamas
Islands, Cayman Islands, Jamaica, Saint Thomas, Saint
Croix, Antigua, Anguilla, Saint Martin-Sint Maarten. Saint
Barts, Nevis, Guadeloupe, Barbados, Mustique, Curasao.
Remarks: This potentially serious horticultural pest
appeared to be restricted to a single greenhouse in Nevis
at the time of collection (2004). However, during the past
decade the species has spread and has now been found
at disturbed environments on both islands. It was recently
reported as introduced on Guadeloupe (Massemin and
Pointier, 2010).
Species Doubtfully ok Erroneously Reported
From the Islands
Due to inaccuracies in provenance of snail specimens
(or the total lack of locality data) collected during the
18th and early 19th century, or misidentifications by
later authors, the following species have been reported
from Saint Kitts. As there is no supporting evidence that
these species are actually occurring on one or both
islands, these taxa should be removed from the list of
terrestrial mollusks from Saint Kitts and Nevis.
Helicina (Analcadia) antillarum G.B. Sowerby II, 1842
Helicina antillarum G.B. Sowerby II, 1842. — Rush, 1891:
67; St. Kitts.
Remarks: This species is distributed on Guadeloupe
(but see, Dominica, and Martinique. The record by Rush
is likely based on a misidentifieation. Delannoye et ah,
2015, however, indicated that the species is most likely
endemic to Martinique). It should be noted that Rush
(1891) also reported an unidentified Helicina species.
We did not encounter any helicinid other than the two
reported herein.
Glyphyalina barbadensis Chase and Robinson, 2001
Hyalina incisa L. Pfeiffer, 1866. — Rush 1891: 68; St. Kitts.
Remarks: This taxon was described from Barbados
as Hyalina incisa by Pfeiffer (1866), and subsequently
renamed Glyphyalina barbadensis by Chase and
Robinson (2001), the original name being unavailable.
We did not encounter this zonitoid species during the
surveys. It may be that Rush misidentified either
immature specimens of Streptaxis glaber (Pfeiffer) or
the Hojeda species mentioned above, whose shells are
superficially similar.
Drymaens ( Mesembrituis ) virgulatus (Ferussac, 1821)
Dn/nmeus ( M esembrinus ) liliaceus Ferussac, 1832. — Pilsbry
' 1889: 11; St. Kitts?
Remarks: Pilsbry reported this taxon from Saint Kitts
without giving a reference. We have been unable to trace
any specimens collected from the island.
DISCUSSION
Our list of land Mollusca from Saint Kitts and Nevis
(Table 3) contains at present 33 taxa, of which six (18%)
are confined to one island (single island endemics; SIEs);
one taxon is endemic to both islands. Two species have
been found which are likely introduced, but at present
their systematic position cannot be ascertained. Of the
list, 22 species are widespread in the West Indies or
beyond (66%). During geological times the areas of these
islands have been dynamic and at the height of the Late
Glacial Maximum (26,500 to 19,000 years before present)
sea levels were ca. 150 m lower (Clark et ah, 2009).
During this time. Saint Kitts and Nevis were united with
Saint Eustatius on the “paleo-island” Saint Kitts-Nevis
Bank. This paleo-island, like others in the Lesser Antilles
at that time, may have well served as separate biogeo¬
graphic areas (Peck, 2011). When we take the land snails
of Saint Eustatius into account (Hovestadt, 1980), eight
taxa (24%) are endemic to this paleo-island; 11 taxa
(33%) may be called Lesser Antillean endemics.
We have furthermore analyzed the fauna with respect
to altitudinal occurrence, and whether they occur on the
A.S.H. Breure et al., 2016
Page 47
Table 3. Summary of species, alphabetically arranged, and their distribution. Abbreviations: E, endemic; I, introduced; x,
collected during surveys mentioned in this paper (Saint Kitts and Nevis) or known from literature (other islands);?, questionable
reports. Species reported for Saints Kitts and Nevis for the first time are shown in bold type.
windward (northeast) or leeward (southwest) side of the
islands. Although the highest elevations have not been
well-sampled (Figures 78-79), some observations may
be made. A number of species occur only at lower altitudes,
viz. Succinea species, Obeliscus swiftianus, Zonitoides
arboreus, Huttonella bicolor, Bulimulus gittenbergeri,
Gastrocopta species, Pupoides marginatus nitidulus.
Poly gym plana plana, and Zachn/sia provisoria. With the
exception of Bulimulus gittenbergeri, these species are
largely introduced species. In contrast, a few species seem
to he restricted to higher elevations, i.e. above 250 m:
Diplosolenodes species, Drymaeus species, Amphibulima
patula christopheri, and Pleurodonte josephinae nevisensis
(Figures 80-81).
600 max =1156
500
max = 985
375
300
250
.cP
78
125
79
Figures 78-79. Altitudinal range of localities sampled. 78. Saint Kitts. 79. Nevis. Dark circles indicate localities at windward side
of the island.
Page 48
THE NAUTILUS, Vol. 130, No. 2
SW
600
450
300
150
Saint Kitts
0
NE
600
x
CD
9.
13
Q.
Q)
CD
<
CD
5. o
(/>
c
o
9.
CD
CL
CD
CD
a
C/D
c
cr
c
Q.
CD
CD
O C/D o
cd =r cd
C/3 CD (/)
T3
CD CD
CD CD
“D
CD
C
s
CD
CD
450
300
1
o - - - — — - - - - - — - — — — — - —
Figure 80. Altitudinal range of species on Saint Kitts, respectively on leeward (upper) and windward (lower) sides of the island.
Colours correspond to those used in distribution maps.
When analyzing the localities related to their diversity
(see Methods), it is clear that most of them have a rather
low species diversity. At four localities no snails were
found; at the remaining localities species diversity
ranged from 1 to 9 (mean 3.12; Figure 73). Taking the
rareness of species into account, the higher elevational
localities tend to score higher (Figures 75-76). There are
six species endemic to both islands (of which five are
SIEs): Bulimulus gittenbergeri, B. ou aliens is, Drymaeus
(. Antidrymaeus ) multifasciatus christopheri, D. (A.) in.
subspecies (Nevis), Amphibulima patula christopheri,
and Pleurodonte josephinae nevisensis. The localitions
where these endemics have been found are given in
Figure 77. The data (adapted from Helmer et ah, 2006)
A.S.H. Breure et ah, 2016
Page 49
sw
Nevis
500
375
250
NE
3
¥
500
375
0 _ _ _ _ _ _ _ _ _ _ _
Figure 81. Altitudinal range of species on Nevis, respectively on leeward (upper) and windward (lower) sides of the island. Colours
correspond to those used in distribution maps.
show that these endemics occur in five vegetation types, of
which three are drought and two are wet types: Drought
Deciduous Open Woodland (. Bulimulus gittenbergeri).
Drought Deciduous Coastal Shrubland {B. gittenbergeri).
Drought Semi-Deciduous Forest (. Drymaeus (. Antidnjmaeus )
multifasciatus christ opheri). Seasonal Evergreen Forest
( D . (A.) multifasciatus christopheri, D. (A.) m. subspecies
(Nevis), Amphibulima patula christopheri, Pleurodonte
josephinae nevisensis), and Evergreen Forest (B. ouallensis,
D. (A.) m. subspecies (Nevis), P. josephinae nevisensis).
Although similar detailed data were not available to us when
writing our study on Dominica (Robinson et al., 2009), we
notice a similar pattern: endemic species are predominantly
found in the wet vegetation types at higher elevations.
Page 50
THE NAUTILUS, Vol. 130, No. 2
In this context it is interesting to take the protection
status of the land snail habitats into account. Helmer
et al. (2008) concluded that during 1949-2000 the land
cover on both islands has changed dramatically, hut a
common aspect is that agricultural land (sugar cane plan¬
tations) has changed into pasture land. Dry and humid
forests increased in area (respectively with 50% for Saint
Kitts and 134% for Nevis respectively) during this
period. Proportional increases in drier formations at
lower elevations were larger than those in evergreen
formations at higher elevations. The degree of (formal)
protection, however, varies for different categories, ranging
from 100% for Evergreen Forest to 0% for lowland
Drought (Semi-)Deciduous Forest (Helmer et al., 2008,
table 5). Gardner (2006) showed that institutional frame¬
works for area protection and protected area programs are
hardly developed on the islands. As pressure for land
development is greatest at lower elevations (Helmer, 2004),
lowland drought shrubland and forest are not well protected.
Hence snail species occurring there may be vulnerable to
habitat fragmentation or destruction. Lugo et id. (2011)
suggested that, due to land cover changes in built-up and
degraded lands, introduced taxa may have a competitive
edge at lowland elevations. From tire data presented on the
lowland species, Bulimulus gittenbergeri especially may be
considered as vulnerable when judged against the IUCN-
eriteria of Critically Endangered species (IUCN, 2012).
In a recent study, Horwith and Kindsay (1999) stated
that data on invertebrates were very limited and informa¬
tion useful to conservation planning was lacking. This void
is filled herein for the land snail fauna of these two islands.
ACKNOWLEDGMENTS
We would like to gratefully acknowledge the support
of the Departments of Agriculture in both Saint Kitts and
Nevis during the 2004 survey. In Saint Kitts, we would
like to thank Director of Agriculture Dr Jerome Thomas,
Mr Kevin Walters, Mr Thomas Jackson, Mr Lawrence
Knight, and Mr Husaini Mawnlawde, all of the Depart¬
ment of Agriculture, Ministry of Agriculture. Of the Nevis
Island Administration, we would like to thank Junior
Minister J. Livingston Herbert, Director of Agriculture
Mr Samuel Powell, Mr Floyd Liburd, and Mr Dwight
Browne, all the Department of Agriculture, Ministry of
Agriculture, Lands, Housing, Co-operatives and Fisheries,
for their support of the USDA project. Finally we thank
Ton de Winter for help with the photograph of the
new species. Laurent Charles and Thomas Watters kindly
provided comments on a previous version of this manu¬
script, which helped to improve it.
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Wagenaar Hummelinck, P. 1953. Description of new localities.
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THE NAUTILUS 130(2):53-71, 2016
Page 53
Comparative anatomy of five species of Saccostrea
Dollfus and Dautzenberg, 1920 (Bivalviai Ostreidae)
from the Pacific Ocean
Vanessa Simao do Amaral
Luiz Ricardo L, Simone
Museu de Zoologia da Universidade de Sao Paulo
Caixa Postal 42494
04299-970 Sao Paulo, BRASIL
[email protected]
[email protected]
ABSTRACT
Ostreids are well known for their high intraspecific variation,
which makes identification problematic. The present paper pro¬
vides a morpho- anatomical analysis of five species of Saccostrea ,
as well as of selected congeneric species from relevant areas in
the Pacific Ocean. Saccostrea cucullata occurs in Africa, Indian
Ocean, Arabian Sea, Red Sea, and part of the Pacific, where it
lives attached to rocks and mangroves roots. Saccostrea glomerate
occurs in the Pacific, New Zealand, Australia and the Philippines,
always associated with rocks. Saccostrea echinafa occurs from
East Africa to japan. S. palmula occurs from Pacific Mexico to
Peru. Saccostrea mordax occurs in tire Red Sea and the Pacific.
The wide distribution and plasticity of these often syinpatric
species led to confusion regarding their identity. Herein we
describe anatomical differences that allow for a more precise
identification, especially when compared to sympatric taxa.
Additional Keywords: oyster, morphology, systematises, pheno¬
typic plasticity
INTRODUCTION
Species of the genus Saccostrea Dollfus and Dautzenberg,
1920 are oysters that live on rocky shores of the Indo-
Pacific region. The taxonomy of this genus is unclear, and
their morphological plasticity makes identification prob¬
lematic (Lam and Morton, 2006). They have small to
medium-sized shells, and a left valve with a prominent
ligament area, with projections in zigzag continuous and
regular. The right valve is flat, bearing projections along
its margin, with corresponding concavities on the left
valve. The genus includes nine species (Huber, 2010):
Saccostrea cucullata (Bom, 1778) - the type species;
Saccostrea cirumsuta (Gould, 1850); Saccostrea echinata
(Quoy and Gaimard, 1835); Saccostrea glomerata (Gould,
1850); Saccostrea kegaki Torigoe and Inaba, 1981;
Saccostrea malubonensis (Faustina, 1932); Saccostrea
palmula (Carpenter, 1857); Saccostrea scyphophilla
(Peron and Lesueur, 1807) and Saccostrea spatulata
(Lamarck, 1819).
Because of shell plasticity, the taxonomy of Saccostrea
has been a troublesome matter; e.g., some “forms” of
S. cucullata have been assigned different names by dif¬
ferent authors, i.e., S. echinata, S. mordax, S. glomerata
(Lam and Morton, 2004). Molecular studies (e.g., Lam
and Morton, 2004; 2006; 2009; Wang and Guo, 2008a;
2008b) have been conducted in attempts to resolve this
taxonomic confusion and better understand the generic
distribution. Reports on the difficulty of proper species
identification, caused by the wide variation in body size
and coloration, are frequent in the literature and deter¬
mine a baffling scenario (Awati and Rai, 1931). This prob¬
lem was further confirmed by Lam and Morton (2006: 1):
“The geographical distribution of lineages of Saccostrea is
somewhat uncertain because of taxonomic confusion”.
Moreover, the widespread, sometimes overlapping, distri¬
butions of these species also make identification problem¬
atic (Huber, 2010).
Molecular systematic studies of Saccostrea are few and
constrained by many taxonomical issues (Lain and Morton,
2006). Furthermore, the genus has been reported to dis¬
play ecomorphological variation (Tack et al, 1992). For
instance, these facts may help explain why all Saccostrea
species from Thailand were previously assumed to be a
single species, S. cucullata (Visoofhivisefh et al, 1998).
However, the results of Klinbunga et al. (2005) indicated
that more than one species of Saccostrea may be present
in Thai waters, which may have implications regarding
its commercial use.
In this paper, we perform a detailed morpho-anatomical
study of five Saccostrea species from the Pacific
Ocean: S. cucullata, from Africa, the Indian and
Arabian Oceans, the Red Sea and part of the Pacific
Ocean; S. glomerata, from the Pacific, New Zealand,
Australia and the Philippines; S', echinata, from East
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THE NAUTILUS, Vol. 130, No. 2
Africa and Japan; S. palmula from Mexico to Peru, in
the Pacific and S. mordax , from the Red Sea and the
Pacific Ocean.
MATERIALS AND METHODS
The study material was preserved in 70-99% ethanol.
Specimens were immersed in the preservative fluid and
dissected by standard techniques under a stereomicro¬
scope (Simone, 1997; 2009; 2011). Details of all systems
(mantle cavity, mantle edge, muscles, digestive, circulatory,
excretory, and nervous systems) and organs were exam¬
ined. Drawings were done with the aid of a camera lucida.
The studied samples are held at several institutional
collections: National Museum of Natural History,
Smithsonian Institution (USNM), Florida Museum of
Natural History (UF), Field Museum of Natural History
(FMNH), and Museu de Zoologia da Universidade de
Sao Paulo (MZSP).
The following abbreviations are used in the figures:
accessory heart (ah), adductor muscle (am), anus (an),
auricles (au), chomata (ch), dorsal hood (dh), esophagus
(oe), external fold (ef), gastric shield (gs), gills (gi), heart
(he), hood (ho), inner fold (if), intestine (in), mantle
edge (me), middle fold (mf), muscle scar (im), pallia]
muscles (pm), palps (pa), Quenstedt muscle (qm), rectum
(rt), sorting area (sa), typhlosole (ty), umbonal cavity (uc),
undulations (pi), ventricle (ve), visceral ganglia (vg), vis¬
ceral mass (vm).
SYSTEM ATICS
Genus Saccostrea Dollfus and Dautzenberg, 1920
Type Species: Ostrea saccellus Dujardin, 1835
(=0. cucullata Born, 1778)
Diagnosis: Shell cup-shaped, with marginal denticles,
promial chamber on the right side.
Distribution: This genus is restricted to tropical and
sub-tropical waters from eastern Atlantic, Indo-West
Pacific, and eastern Pacific, while it is absent in the
western Atlantic; in saline areas (open ocean), but occa¬
sionally occurring in mangrove areas. Most species live
exposed, fixed on rocks or coral (Harry, 1985).
Saccostrea cucullata (Born, 1778)
(Figures 1-15)
Ostrea cuccullata [sic] Born, 1778: 100.
Ostrea cucullata Born, 1780: 114, pi. 6, figs 11, 12;
Dillwyn, 1817: 277; Smith, 1890: 322; Awati and Rai,
1931 1-107.
Ostrea comucopiae Gmelin, 1791: 3336; Kiister, 1868: 77.
Lopha comucopiae: Boding, 1798: 169.
Ostrea gibbosa Lamarck, 1819: 209; Hanley, 1856: 301.
Crassostrea cucullata Ranson, 1960: 20.
Saccostrea cucullata: Stenzel, 1971: N1134-N1135, fig.
J 106; Morris, 1985: 125-128, Pi. 3, figs E-G; Lam and
Morton, 2003: 110-112, pis. 11, 12; Lam and Morton,
2009: 482. fig. 1: 483.
Description: Shell: Elongated, generally cup-shaped,
to 100 mm. Right (upper) valve flat or slightly convex
(Figure 2). Ligament alivineular. Left valve fixed to sub¬
strate, cupped and larger than right valve. Umbonal cavity'
of left valve. Margins of both valves with angles sculpture
in the edge (Figures 1, 3); inner edge of right valve with
small elongated denticles (Figure 2) producing corre¬
sponding depressions on left valve. Adductor muscle
scar reniform, occupying 14 of total shell height, in
posterodorsal region (Figure 2), striate, white or grayish.
Outside of valves variable from white to gray, light or dark
brown, green or purple. Inner surface white, with occa¬
sional puqde spots. Adductor Muscle: Reniform or
slightly oval, located in posterior region, occupying ‘A of
total soft part height (Figures 4, 11, 12). Mantle: Mantle
lobes thick, with well-defined pallia! muscles, most abun¬
dant near adductor muscle (Figures 4, 5). Hood formed by
junction of anterior mantle lobes. Hood robust and well-
defined, usually free or filled by gonad mass (Figures 4,
11). Mantle Edce: Pleated and thick, dark, yellow with
brown spots, bearing three similar-sized folds. Middle and
inner folds with finger-like, uniformly distributed papillae,
single or in pairs, with variable length (Figure 5). Gills:
Gills occupying 50% of total height. Demibranchs mod¬
erately thick, ~l/3 as thick as gill fold (Figures 6, 12).
Circulatory System: Ventricle large, thick-walled, with
internal beams crossing each other in various directions;
opaque white. Outer part of auricles thin, translucent,
trabecular; external portion of auricles bearing small,
saculiform structures (Figure 7). Pair of kidneys triangular,
tubular, mottled by light brown marks. Each kidney hav¬
ing direct communication with pericardium and gonad on
medial side and communicating laterally by short cham¬
ber. Renal opening located on opposite side of cerebro-
visceral connective. Digestive System: Palps large, length
14 of gill size, spatuliform; internal surface with pleats
extending transversely to edge (Figure 6). Stomach in
globular region of visceral mass, typically occupying 14
of total length. Esophagus short, 14 of palp length,
(Figure 13). Sorting area of stomach (sa) short (1/2 of palp
length) and wide with typhlosole beginning in posterior
region of stomach (Figure 9, 13). Intestine separated from
style sac, returning in opposite direction, passing usually
behind pericardium, immersed in gonads; narrowest por¬
tion of intestine surrounding adductor muscle. Rectum
long and thin, surrounding adductor muscle almost until
canopy-gill junction. Anus sessile, rounded, 14 of height of
adductor muscle, with single fold. Reproductive System:
Gonad composed of two lobes of f ollicular aspect, occupy¬
ing 14 of total animal size. Dorsally and ventrally spread in
left lobe and right lobe; shape indistinct from each other.
Two pairs of separate systems of genital channels, same
as described above (excretory system). Nervous System:
Nervous system similar to other ostreid species. Pair of
V.S. Amaral and L.R.L. Simone, 2016
Page 55
Figures 1-10. Saccostrea cucullata, shell and anatomical features. 1. Shell in right view. 2, Right valve, internal view. 3. Shell in
ventral view. 4. Complete soft parts; (ho) hood; (vm) visceral mass; (am) adductor muscle. 5. Dissected mantle lobe, mantle edge and
accessory heart; (all) accessory heart; (me) mantle edge. 6. Palps (pa), right-slightly ventral view, outer right hemipalp deflected.
7. Heart isolated, showing ventricle (ve) and auricles (au). 8. Anus and rectum. 9. Anterior region, right view, digestive system
exposed; (oe) esophagus; (sa) selection area; (dh) dorsal hood. 10. Visceral ganglia, ventral view, adjacent layer of tissues also shown.
Scale bars = 1 cm.
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THE NAUTILUS, Vol. 130, No. 2
Figures 1 1-15. Saccostrea cucullata, anatomical features. 11. General view of soft parts; (ho) hood; (vm) visceral mass; (pa) palps;
(am) adductor muscle; (gi) gills. 12. General view, right mantle lobe removed; (vm) visceral mass; (pa) palps; (am) adductor muscle;
(gi) gills; (he) heart; (rt) rectum; (an) anus. 13. Stomach, right view, opened longitudinally; (oe) esophagus, (sa) selection area,
(dli) dorsal hood, (gs) gastric shield, (ty) typhlosole, (in) intestine. 14. Visceral ganglia, ventral view. 15. Detail of anus. Scale bars = 1 cm.
V.S. Amaral and L.R.L. Simone, 2016
Page 57
cerebral ganglia in anterior region, near palps; link at pair
of visceral ganglia by connectives; visceral ganglia occupy¬
ing 1/15 of adductor muscle area, with two pairs of
branches in anterior region, three in posterior region and
one laterally (Figures 10,14).
Type Material: Borns holotype is located at the
Naturhistorisches Museum Vienna (Morris, 1985). Ostrea
comucopiae, Syntypes, Geneve Museum n° 1089141,
Lamarck Coll, (examined).
Material Examined: China: Hong Kong, North Pacific
Ocean, USNM 858434, 3 specimens; New Zealand: South
Pacific Ocean, USNM 886486, 5 specimens; Thailand:
Chantahuri, Kung Kraben Bay, MZUSP 55270, (L. R.
Simone coll. 24/vii/2005), 6 specimens.
Distribution: Indo-West Pacific.
Saccostrea glomerata (Gould, 1850)
(Figures 16-27)
Ostrea glomerata Gould, 1850: 346; 1852: 462; Sowerby,
1871: sp. 64.
Ostrea attenuata Sowerby, 1871: pi. 21.
Ostrea vitrefacta Sowerby, 1871: sp. 80; Lamy, 1929: 166;
ile-Rodriguez, 1938: 289.
Ostrea mordax. — Hutton, 1873b: 84 (non Gould, 1850).
Lopha glomerata. — Finlay, 1928: 268.
Ostrea forskali var. Glomerata. — Lamy, 1929: 158.
Ostrea (Lopha) glomerata. — Hiro, 1936: 36; Wada, 1942: 70.
Crassostrea cucullata. — Ranson, 1967: 188; Nishimura
et ah, 1998: 85 ( non Born, 1778).
Saccostrea glomerata. — Torigoe, 1981: 310, 332; Dinamani,
1991a: 335; Xu, 1997: 94.
Saccostrea cucullata. — Oliver, 1992: 88; Nishimura et al.
1998: 86, 88 ( non Born, 1778).
Description: Shell: Rounded, to 50 mm; left valve
concave, slightly larger than right valve, lacking well-
developed umbonal cavity (Figure 16). Right valve slightly
convex; ligament short, with mild bulging in central region
(Figure 17); margins of both valves with well-developed
angles (Figures 16, 17); color internally dark brown at
edges, fading toward median region of shell. Right valve
with single row of rounded denticles spread along entire
edge (Figure 17), with corresponding depressions on left
valve. Adductor muscle scar reniform, occupying 1/5 of
total inner shell area, located in median region, with ante¬
rior portion normally pigmented gray/brown and posterior
portion with pigmented lines; surface slightly pearlescent,
white-colored. Quenstedt muscle scar rounded, occupy¬
ing 1/20 of total area of adductor muscle scar, located in
anterior-ventral region of valves (Figure 17). Muscles:
Adductor muscle reniform, occupying 1/3 of total height,
with a convexity in posterior region (Figures 18, 24).
Mantle: Mantle lobes slightly thick, with well-defined
pallial muscles more abundant near adductor muscle
(Figures 18, 19). Mantle edge thick, reddish; middle fold
with .3-4 short to long, elongated papillae; interned fold with
short to medium papillae, presenting 2 — 3 medium papil¬
lae for each short one (Figure 19). Gills: Gills occupying
M> total body size. Alimentary channel narrow, ~ 1/5 as thick
as gill fold (Figure 18). Heart Heart well-developed. Ven¬
tricle relatively large, with internal bundles crossing each
other in different directions, light beige. Auricles thin, with
more abundant bundles, same texture, and coloration,
but more elongated than ventricle (Figure 21). Digestive
System: Palps occupying ~l/5 of visceral mass, spatuliform,
rounded; internal surface with plicae extending trans¬
versely to edge (Figures 20, 23, 25). Esophagus long and
narrow; length ~ 1/2 of palp length (Figures 22, 26).
Selection area (sa) long and thin, with typhlosoles begin¬
ning in posterior region of stomach (Figures 22, 26).
Intestine passing behind pericardium, with narrowest por¬
tion surrounding adductor muscle. Rectum slightly long
and thin, contouring adductor muscle at dorsalmedial
region. Anus with single fold (Figure 22). Nervous System:
Two pairs of branches in the anterior region and three in
posterior region (Figure 27).
Type Material: Holotype, USNM 5960 (examined);
Paratypes USMN 612314, MCZC 178590, 178591.
Material Examined: Fiji: Nanuya-Lailai Island, Yasawa
Group T. Moala. MZUSP 71454; 2 specimens.
Distribution: India to Australia and New Zealand.
Saccostrea echinata (Quoy and Gaimard, 1835)
(Figures 28-42)
Ostrea mytiloides Lamarck, 1819: 207; Deshayes, 1836:
227; Hanley, 1856: 300; Hidalgo, 1905: 387; Lamy, 1924:
155, 1929: 138 ( non Gmelin, 1791).
Ostrea echinata Quoy and Gaimard, 1835: 455; Hanley,
1856: 302; Kiister, 1868: 76; Hidalgo, 1905: 387; Chen
et al. 1980: 174.
Ostrea arakanensis Sowerby, 1871: sp. 83.
Ostrea nigromarginata Sowerby, 1871: sp. 85; Lamy,
1928: 140.
Ostrea cucullata. — Pilsbry, 1895: 146 ( non Born, 1778).
Crassostrea echinata. — Thompson, 1954: 152: Carreon,
1969: 113.
Saxostrea mytiloides [sic]. — Habe and Kosuge, 1966:145
(non Gmelin).
Saccostrea echinata. — Stenzel, 1971: N962; Torigoe and
Inaba, 1981: 126; Torigoe, 1981: 308, 330; Li and Qi,
1994: 171; Hayami, 2000: 927.
Saccostrea cucullata. — Morris, 1985: 125; Harry, 1985:
150; Oliver, 1992: 88, 91; Xu, 1997: 95 (non Born, 1778).
Description: Shell: Elongated, ~ 60 mm, with slight
ripples in margin of valves extending toward central
region (Figures 28-30); left valve slightly larger than
right valve, with deep umbonal cavity, lacking any major
projections (Figure 29); internal color white/cream to
gray. Adductor muscle scar elongate-reniform, occupy¬
ing ~14 of total height of shell, non-pigmented (Figure 30).
Denticles present across edge of right valve (Figure 30),
Page 58
THE NAUTILUS, Vol. 130, No. 2
Figures 16-23. Saccostrea glomerata, shell and anatomical features. 16. General right view. 17. Right valve, internal view;
(ch) chomata, (im) muscle impression. 18. General view of soft parts right view, right mantle lobe removed; (vm) visceral mass,
(am) adductor muscle, (pa) palps, (gi) gills. 19. Mantle lobes and edge; (if) inner fold; (mf) middle fold;(pm) pallial muscles.
20. Heart, ventral view; (ve) ventricle; (au) auricle. 21. Anterior region, (qm) Quenstedt muscle. 22. Anterior region, right view, with
Stomach opened longitudinally; (oe) esophagus, (sa) selection area, (dh) dorsal hood. 23. Palps (pa), right-slightly ventral view. Scale
bars = 1 cm.
V.S. Amaral and L.R.L. Simone, 2016
Page 59
oe
Page 60
THE NAUTILUS, Vol. 130, No. 2
Figures 28-36. Saccostrea echinata, shell and anatomical features. 28. External view. 29. Internal view, left valve; (im) muscle
impression; (eh) chomata. 30. Internal view right valve. 31. General view of soft part;(vm) visceral mass, (am)adductor muscle,
(me) mantle edge. 32. Anterior region; (qin) Quenstedt muscle, (pa) palps, (gi) gills. 33. Mantle edge (me), detail of pallia! muscles
(pm) and accessory heart (ah). 34. Visceral ganglia. 35. Heart, (ve) ventricle, (au) auricle and rectum (rt). 36. Anus with expansion
and rectum. Scale bars = 1 cm.
VS. Amaral and L.R.L. Simone, 2016
Page 61
Figures 37-42. Saccostrea echinata, anatomical features. 37. General view of soft parts; (qm) Quenstedt muscle, (vm) visceral
mass, (he) herart), (am) adductor muscle, (rt) rectum, (an) anus, (pm) pallial muscles, (all) accessory heart, (me) mantle edge,
(gi) gills, (pa) palps. 38. Palps, detail of median fusion of external hemipalps. 39. Anus (an) and rectum (rt), detail of expansion in
anus. 40. Stomach, right view; (oe) esophagus, (sa) selection area, (gs) gastric shield, (dli) dorsal hood, (ty) typhlosole. 41. Heart in
pericardium;(ve) ventricle, (au) auricle. 42. Visceral ganglia. Scale bars = 1 cm.
Page 62
THE NAUTILUS, Vol. 130, No. 2
with corresponding depressions on left valve (Figure 29),
distributed in various rows along posterior region, being
elongated, rounded or both. Muscles: Reniform, occupy¬
ing ~ !4 of total height, with a convexity in posterior
region (Figures 30, 37). Mantle: Mantle edges delicate,
with well-defined pallial muscles in all mantle surfaces.
Hood filled with gonads (Figures 31, 33, 37). Mantle
Edge: Mantle edge thick, with yellow and/or brown pig¬
mentation. Papillae of middle fold alternately long and
short; usually with a long papilla for each three short ones.
Accessory heart with only one branch on both lobes, thin
and translucent; lobes not reaching adductor muscle
(Figure 33). Gills: Gills occupying Vi of total height.
Alimentary chamber thick, as thick as gill fold
(Figures 32, 37). Circulatory System: Ventricle thin, usu¬
ally translucent, auricles elongated, less cross-linked, non-
pigmented (Figures 35, 41). Digestive System: Palps small,
~ ’A of adductor muscle size, curved, sickle-shaped, with
folds along anterior margin. External palps fused in
middle region and overlying internal palps (Figures 32, 39).
Esophagus long (~l/3 of adductor muscle) and thin.
Selection area of stomach (sa) short (~ Vi of esophagus
size) and wide, with typhlosole (ty) beginning in posterior
region of stomach (Figure 38). Intestine passing behind
pericardium; narrowest portion circling adductor muscle.
Rectum long and thin, girdling adductor muscle. Anus
papillae shaped similarly to shell (Figure 35, 36, 40).
Nervous System: With three branches in anterior region
and only one in posterior region (Figures 34, 42).
Type Material: Holotype in MNHN (examined).
Materia] Examined: Mariana Islands: Guam, UF 284793;
3 specimens.
Distribution: E. Africa to Japan.
Saccostrea palmula (Carpenter, 1857)
(Figures 43 -53)
Ostrea palmula Carpenter, 1857b: 163, 550; 1864: 538;
Dali, 1914: 2; Lamy, 1929: 150; Keen, 1971: 84; Abbott,
1974: 456.
Ostrea plumula [sic]. — Carpenter, 1857b: 351, 353.
Ostrea amara Carpenter 1864a: 363; 1864: 541, 552, 621,
666; Lamy, 1930: 242.
Ostrea from. — Carpenter, 1864: 520 ( non Linnaeus, 1758).
Ostrea mexicana Sowerby, 1871: sp. 35.
Ostrea cumingiana var. Mexicana. — Dali, 1914: 2.
Ostrea conchaphila (pars). — Abbott, 1974: 456 (non
Carpenter, 1857).
Saccostrea palmula. — Harry, 1985: 138.
Crassostrea palmula. — Rodrigues and Garcia-Cubas,
1986: 269.
Description: Shell: Shell rounded, to ~60 mm. Left
valve larger than right valve, with small umbonal cavity
and slight projection in ligament region; projections and
evident ripples present along margin (Figures 43, 44).
Right valve smaller than left valve, opercular, following
pattern of expansion of left valve (Figure 44). Color
white/cream with puqule spots on outer surface; inner
surface white, slightly nacreous, with occasional punple
spots and lines. Adductor muscle scar well-defined,
showing purple concentric lines in anteroventral portion
of valves, reniform (Figures 44, 45), occupying 1/5 of
total height. Denticles rounded in anterior region, aligned,
uniformly distributed; corresponding depressions on right
valve shallow, almost inconspicuous (Figure 44). Muscles:
Same morphology as other species, reniform (Figures 46,
47, 50), occupying ~l/5 of total height. Mantle: Mantle
lobes thin, with well-defined pallial muscles in posterior
region (Figures 46, 50); hood absent. Mantle edge thinner,
beige to cream; middle fold bearing two to five finger-like,
medium papillae for each two long ones (Figure 47).
Accessory heart with three well-defined branches,
reaching and circling adductor muscle in ventral region
(Figure 47). Gills: Gills occupying ~ 1/3 of total body
size. Alimentary chamber narrow (Figures 46, 48).
Heart Ventricle elongated, slightly thick, reticulated
and whitish. Auricles thin, small and non-pigmented
(Figures 46, 50). Digestive System: Palps large, occupy¬
ing half as large as adductor muscle, thin, with folds
on upper margin (Figures 48, 51).
Nervous System: Visceral ganglia with two anterior
expansions, two lateral and tree posterior ones (Fig¬
ures 49, 53).
Type Material: Syntypes, RMNH 1857.6. 4.735, 1857.6.
4.736, 1857.6. 4.737 (examined).
Materia] Examined: Panama: Miraflores Lock, USNM
734199, 2 specimens; Arraijan, FMNH 27226, 2 specimens.
Distribution: Mexico to Peru.
Saccostrea mordax Gould, 1850
(Figures 54-64)
Ostrea mordax Gould, 1850: 346; 1852: 464; 1856: pi. 43;
Sowerby, 1871: sp. 31; Saville-Kent, 1891: 2; 1892: 65,
245; Hutton, 1937b: 84; Hirase and Taki, 1951: pi. 7.
Ostrea comucopiae Saville-Kent, 1891: 3 ( non Gmelin,
1791).
Ostrea mordax var. comucopiaeformis Saville-Kent,
1893: 248.
Ostrea cucullata. — Iwakawa, 1915: 15; 1919: 254; Hatai,
1941: 58; Wells and Biyce, 1988: 162.
Ostrea forskali var. sueli Lamy, 1925: 192.
Ostrea (Lopha) cucullata. — Kuroda, 1928: sp. 36; Hirase,
1930: 25; 1934: 5; Takatsuki, 1949: 5 ( non Bom, 1778).
Ostrea forskali var. mordax. — Lamy, 1929: 158.
Ostrea (Lopha) mordax.— Kuroda, 1930: 51; Tchang and
Lo, 1956: 74.
Saxostrea amasa Iredale, 1939: 399; Allan, 1959: 273.
Ostrea glomerata. — Blanco et al., 1951: 52 ( non Gould, 1850).
Crassostrea amasa. — Thompson, 1954: 154; Carreon,
1969: 111.
V.S. Amaral and L.R.L. Simone, 2016
Page 63
Figures 43-49. Saccostrea palmula, anatomical features. 43. External view. 44. Internal view of left valve; (im) muscle impression.
45. Internal view of right valve. 48. General view of soft parts; (qm) Quenstedt muscle, (vm) visceral mass, (he) heart, (rt) rectum,
(an) anus, (am) adductor muscle, (pm) pallial muscles, (me) mantle edge, (gi) gills, (pa) palps. 47. Posterior region, detail for a gills
(gi), mantle edge (me), accessory heart (ah) and adductor muscle (am). 48. Detail of palps. 49, Visceral ganglia. Scale bars = 1 cm.
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THE NAUTILUS, Vol. 130, No. 2
Figures 50-53. Saccostrea palmula, shell and anatomical features. 50. General view of soft parts; (qm) Quenstedt muscle,
(vm) visceral mass, (he) heart, (rt) rectum, (an) anus, (all) accessory heart, (pm) pallia! muscles, (gi) gills, (pa) palps. 51. Detail of
palps. 52. Stomach, right view; (oe) esophagus, (sa) selection area, (gs) gastric shield, (dh) dorsal hood, (ty) typhlosole, (pa) palps,
(he) heart, (am) adductor muscle, (rt) rectum, (in) intestine. 53. Visceral ganglia. Scale bars = 1 cm.
V.S. Amaral and L.R.L. Simone, 2016
Page 65
Figures 54-81. Saccostrea inordax, shell and anatomical features. 54. External view. 55. Internal ’view of right valve; (ch) chomata,
(im) muscle impression. 58. Internal view of left valve, detail of undulations (pi) of the edge valves and umbonal cavity (uc). 57. General
view of soft part; (ho) hood, (an) anus. 58. Mantle edge, detail of accessory heart (ah) and pallia! muscles (pm); (ef) external fold,
(m.Q middle fold, (if) inner fold, (me) mantle edge. 59. Detail of the palps (pa) and gills (gi). 60. Heart;(ve) ventricle, (au) auricle.
61. Visceral ganglia. Scale bars = 1 cm.
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THE NAUTILUS, Vol. 130, No. 2
Figures 62-64. Saccostrea mordax, anatomical features. 62. General view of soft parts; (pa) palps, (gi) gills, (vm) visceral mass,
(he) heart, (rt) rectum, (an) anus, (am) adductor muscle, (pm) pallia] muscles, (me) mantle edge. 63. Stomach, right view; (pa) palps,
(oe) esophagus, (sa) selection area, (gs) gastric shield, (dh) dorsal hood, (ty) typhlosole, (in) intestine. 64. Visceral ganglia. Scale
bars = 1 cm.
V.S. Amaral and L.R.L. Simone, 2016
Page 67
Saxostrea mordax.- — Habe, 1951: 94; Kira, 1959: 127;
1962: 144; Tald, 1960: 192; Habe and Kosuge, 1966:
145; 1967: 137.
Saccostrea mordax. — Habe and Okutani, 1975: 194;
Habe, 1977: 109; 1981: 83; Torigoe, 1981: 306, 328;
Okutani and Soyama, 1987: 152; Habe and Matsuda,
1990: 98; Xu, 1997: 95; Hayami, 2000: 925.
Saccostrea cucullata — Morris, 1985: 125; Li and Qi, 1994:
170; Lamprell and Healy, 1998: 134 ( non Bom, 1778).
Description: Shell: Shell elongated, to ~40 mm, with
undulations on entire edge of both valves up to
ligamentary region (Figure 54). Left valve concave,
umbonal cavity well-developed, with extensive anterior
projection (Figure 56). Right valve flat or slightly convex,
not opercular (Figure 55). Color beige/cream, with edges
of valves dark purple. Denticles present along edges;
rounded, relatively large and distributed in single row
(Figure 55). Adductor muscle scar oval, grayish or non-
pigmented (Figure 55). Mantle: Mantle thick, with pallial
muscles more evident in adductor muscle region. Hood
present, robust, filled by gonads and palps (Figure 57).
Mantle edge thin, dark brown and yellow; middle fold
with small and medium finger-like papillae; 2^4 small
papillae for each larger one. Internal fold with small,
uniformly distributed finger-like papillae. Accessory heart
inverse-T shaped, well-marked, and surrounding ventral
region of adductor muscle (Figures 58, 62). Gills: Gills
occupying of total area. Alimentary chamber narrow
(Figures 59, 62). Circulatory System: Ventricle large, ~2/3
as large as auricles, with slight bundles more concentrated
in posterior region; whitish. Auricles elongated, translu¬
cent (Figure 60). Digestive System: Palps large and thick,
of adductor muscle size, superior edge with folds;
external palps covers ~l/3 of internal palps (Figures 59,
62). Esophagus long, twice as long as palps. Selection area
of stomach (sa) long and narrow, Vz as high as esophagus;
typhlosole in terminal region of globular portion (Figure 63).
Nervous System: Visceral ganglia with tree nerves in ante¬
rior and posterior region and one laterally (Figures 61, 64).
Type Material: Syntype, NMNH 5958 (examined).
Material Examined: China: Taiwan, Taipei Co.,
Wanli, Yehliu, UF 303012, 3 specimens.
Distribution: Indo-West Pacific.
DISCUSSION
It is an accepted fact that that the taxonomy of Saccostrea
is problematic because of shell plasticity (Lam and
Morton, 2006). Still, except for the shell, the anatomy of
the species is mostly unknown (e.g., Lam and Morton,
2004, 2006, 2009). Among the species studied here, the
only one with a described anatomy was S. cucullata, by
Awati and Rai (1931).
The shell of Saccostrea differs from that of Crassostrea
basically by the presence of chomata and the ornamen¬
tation of the edge in some species. Both genera have
medium to large-sized shells and occur in similar habi¬
tats, but Saccostrea species they were found only in the
Pacific and Indian Oceans. An invasive species of the
genus Saccostrea, found in Sao Sebastian (Sao Paulo,
Brazil), is identified as S. cucullata (see Amaral and
Simone, in press). As we demonstrate herein, among the
Saccostrea species, taxonomy is problematic only when
based solely on shell characters (even so, differences in
shell morphology may be found when the characters are
considered in sufficient detail). Anatomical characters are
useful and sufficient for species-level identification.
Methods in molecular systematics have been advocated
as the only tools for identification of the various lineages
of Saccostrea (Lam and Morton, 2006: 7). Observations of
S. mordax suggest that it occurs only on exposed, wholly
marine rocky shores, whereas other Saccostrea lineages
occupy a wide range of habitats from brackish mangroves
to somewhat less exposed marine shores (Lam and
Morton, 2006).
Shell: The shells of Saccostrea cucullata are elongated,
normally cup-shaped, with a deeply concave left valve
showing an expansion in the ligament area. The right
valve is fiat or slightly convex, like a lid. The shell of
S. glomerate is rounded, with a concave left valve slightly
larger than the right valve, and a slightly convex right
valve. In S. echinata, the shell is more elongated, with
slight undulations, along the edge of the valves that stretch
toward the central area. 8. palmula has a rounded shell,
with both valves laterally expanded; the left valve is larger
than the right, and the right valve is small and opercular.
Saccostrea mordax has a usually elongated shell, with
undulations going from the margins to the ligament
region; the left valve is concave, the right valve is almost
flat or slightly convex, but it is not opercular as in the
remaining species, and both valves have a similar width.
The shell edge undulations are characteristic of Saccostrea,
with this character usually separating its species from
those of Crassostrea. When present in Crassostrea, this
undulation is only observed in the posterior region and in
follicular layers.
The umbonal cavity is well-defined in Saccostrea
cucullata , S. echinata, and S. mordax. In S. glomerate
and S. palmula it is small or absent. According to Stenzel
(1971: N995), the deepest umbonal cavities and the
largest number of chambers are found in Saccostrea,
particularly in the high-conical, rudist-like ecomorphs of
the “living complex superspecies S. cucullata” from the
tropical Indo-Pacific.
The shell edge in S. cucullata has small tubercles
(elongated chomata). They are present along the inner
edge of the right valve, with corresponding depressions
on the left valve. In S. glomerata, the margins of both
valves have well-developed angles; the right valve shows
rounded chomata spread along its entire edge, arranged
in single row, with corresponding depressions on the left
valve. In S. echinata, the chomata are present along the
right valve’s edge, with corresponding depressions on the
left valve; these chomata are distributed in multiple rows
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THE NAUTILUS, Vol. 130, No. 2
in the posterior region, and may have an elongated or
rounded contour. Saccostrea palmula has evident projec¬
tions and undulations along the shell’s edge; following
the pattern of expansions of the left valve, both valves
show more rounded chomata on the edge than those of
S. cucullata, which is restricted to the anterior region,
aligned and uniformly distributed. The corresponding
depressions on the right valve are somewhat shallow.
Saccostrea mordax has chomata along the edges on both
valves; they are rounded, relatively large and distributed
in a single row.
Some oysters have chomata distributed around the
peripheries of their valves, including the valve margins
directly opposed to the hinge. True teeth are never
found in this area of bivalve shells. Chomata are not
analogous to teeth or sockets. The genus Saccostrea has
strongly developed chomata, tall and strong tubercles on
the right, and deep pits on the left valves along the whole
periphery of the valves (Stenzel, 1971).
In Saccostrea cucullata , the external surface of the
valves is variable in color, from pale white, light-gray to
dark-brown, greenish, or purple. The internal surface
is white, and may present purple spots. These charac¬
teristics support those presented by Awati and Rai
(1931). Saccostrea glomerata is internally colored dark
brown at the edges, fading toward the median region
of the shell. Comparatively, S. echinata is colored
white/cream to grayish. Saccostrea palmula is colored
white/cream with purple spots externally, and the inter¬
nal surface is white and slightly nacreous; there may be
purple spots in the region corresponding to the mantle
edge on both valves. In Saccostrea mordax , color is
cream in the middle, with dark purple colored in the
internal margins.
The adductor muscle scar in Saccostrea cucullata is
reniform, striated, white or grayish. In S. glomerata, the
adductor muscle scar is reniform, occupying 1/5 of total
area, located in the median region, with anterior portion
normally pigmented gray/brown and a posterior portion
with pigmented lines. Its surface is slightlv opalescent
and white. In S. echinata , the adductor muscle scar is
also reniform but more elongated and unpigmented; in
S. palmula , it differs from the above-mentioned species:
the adductor muscle scar is defined by concentric purple
lines in the anteroventral portion of the valves, but it is
also reniform. S. mordax has an oval adductor muscle
scar, grayish, with concentric lines that may be pigmented.
These characteristics fit with those described by Lam and
Morton (2009).
All species of Saccostrea exhibit a slight Quenstedt
muscle scar, a rounded mark located in the anteroventral
region of the valves. These imprints have been noticed
on many oyster species, fossil and living (Dali, 1880;
Hedman and Boyce, 1890, Stenzel, 1971, Harry, 1985).
The adductor muscle (and scar) of Crassostreinae in
general do not differ among genera but show variations
among species. These variations have been pointed out
in the literature (Nascimento, 1991 for C. rhizophorae-
Galtsoff, 1964 and Kennedy et ah, 1996 for C. virginica),
and are possibly associated with the high level of
shell plasticity.
Adductor Muscle: The adductor muscle is short and
stout and directly connects the two valves. It is differen¬
tiated by its whole length into two coalescent subdivisions:
the catch muscle (with opalescent and opaque color in
live oysters, comprising the ventral or distal part of the
muscle), and the quick muscle (flesh, colored and trans¬
lucent, comprising the dorsal or proximal part of the mus¬
cle). The relative sizes of the two subdivisions differ
among genera (Stenzel, 1971). Ostrea has a larger catch
muscle then Crassostrea and Saccostrea ; Hyotissa has the
smallest one.
Saccostrea cucullata shows a normally reniform adduc¬
tor muscle, which is sometimes oval while S. palmula has
a strictly reniform muscle. Saccostrea glomerata has a
reniform adductor muscle, with a convexity in the poste¬
rior region. Saccostrea. echinata also has a reniform
adductor muscle, but with a convexity toward the poste¬
rior region (that of S. cucullata is anterior). Saccostrea
mordax has a strictly oval adductor muscle.
The Quenstedt muscles are a pair of tiny muscles,
each attached at one extremity to a valve, and having the
opposite extremity ending among the oral part of the
gills. The pair of Quenstedt muscles has no differentia¬
tion regarding the origin on the shell and the insertion in
the base of the palps in the five species. Galtsoff (1964),
Stenzel (1971), and Mesquita (1993), observed that the
Quenstedt muscles are difficult to find, but, according to
Amaral and Simone (2014), the pair of Quenstedt mus¬
cles are visible when opening the valves in all species of
Crassostreinae. From their origin on the shell, the mus¬
cles run through the anterior visceral mass up to the
insertion in the base of the palps. This peculiar morpho¬
logical arrangement shows no probable homology with
the anterior adductor muscle or any foot muscles
(Galtsoff, 1964; Stenzel, 1971).
Mantle: The mantle lobes of an adult oyster are thick
and somewhat solid, having many muscles strands, some
arranged in concentric and others in radial patterns
(Stenzel, 1971). In Saccostrea cucullata, the lobes are
thick, with well-defined pallial muscles, mainly around
the adductor muscle; the hood is robust, well-defined,
normally free or filled by gonads. In S. glomerata, the
mantle is slightly thick, with pallial muscles more evident
in the adductor muscle region, and the hood is absent. In
S. echinata, the mantle lobes are thin, with well-defined
pallial muscles along the mantle’s extension. It has a
smaller hood than S. cucullata. Saccostrea palmula has a
thick mantle with pallial muscles more evident around the
adductor muscle. The hood is not present. In S. mordax,
the mantle lobes are thick with pallial muscles more
evident in the adductor muscle region. It also has a
well-defined hood, filled by gonads and palps. The pallial
muscles show some differences among species, in quan¬
tity, girth, or location.
V.S. Amaral and L.R.L. Simone, 2016
Page 69
Saccostrea cucullata lias a thick mantle edge, colored
dark vellow with brown spots. Its three folds have similar
size; the middle and inner folds also have finger-like
papillae, one or two short for each long one. Saccostrea
glomerata has a thick mantle edge with reddish color; the
middle fold has both short and long papillae, at a ratio of
three to four short for each long one; the inner fold has
short and medium papillae, usually two or three medium
for each short one. In S. echinata, the mantle edge is
thick and colored yellow and/or brown; the papillae of
the middle fold are alternately long and short, with three
short for each long one. Saccostrea palmula has a thin
edge, colored heige/cream; the finger-like papillae of the
middle fold are small, elongated, alternately short and
medium/long. Usually, two to five small papillae are pres¬
ent for each one or two long/medium ones. S. mordax has
a thin edge, colored dark brown and/or yellow; the middle
fold bears medium and short finger-like papillae, at a ratio
of two to four short papillae for each medium one; the
internal fold bears small, uniformly distributed finger-like
papillae. This variation (size and distribution) was also
Found in Amaral and Simone (2014) for Crassostrea species.
The accessory heart in Saccostrea cucullata has three
well-defined branches in the right lobe, forming a short
y-shaped structure while the left lobe shows only a single
large branch. In S. echinata, the accessory heart is slim,
translucent, with thin walls, not reaching the adductor
muscle in both lobes. In S. palmula , it has tree branches
that contour the adductor muscle. Saccostrea mordax
shows a different arrangement, forming a well-defined
inverted T-shaped structure, and girdling the adductor
muscle in the ventral region. Saccostrea glomerata has
three branches in both lobes, forming a thick y-shaped
structure, and one branch, about 1/3 thinner, which bor¬
ders the adductor muscle.
Differendy from die species described here, Crassostrea
had tree branches in all studied species in Amaral and
Simone (2014). This characteristic can be used as a distinc¬
tion between Saccostrea and Crassostrea.
The promial passage was first reported by Kellogg
(1892: 396-397). Nelson (1938) was the first to recognize
its significance and to elucidate its function. The promial
passage is present in all Saccostrea. This is a characteristic
of the nonincubatory ostreid genera (e.g., Crassostrea,
Saccostrea, Striostrea, Hijotissa, Neopci/nodonte; Stenzel,
1971). The promial passage varies in size: in species of
Crassostrea it appears on both sides as a long chamber,
from the margin of the visceral mass to the base of the
gills; in Saccostrea, the passage is open only in the left
lobe and open to half of length of the visceral mass.
Gills: The number of gill folds and filaments is very
variable, apparently increasing proportionally to the
animal’s size. However, these numbers can vary among
individuals of similar size. Ridewood (1903) mentioned
9-12 filaments per fold in Ostrea edulis, while Atkins
(1937a) mentioned 8-17. Atkins (1937a) mentioned 8-17
in Crassostrea virginica; Nelson (1938) mentioned 8-15.
Galtsoff (1964) mentioned 10-16 filaments per fold in
C. angulata, while Atkins (1937a) referred to 15-16 and
Nelson (1938) 8-14 filaments per fold. Amaral and
Simone (2014) observed that the number of filaments
per fold in C. rhizophorae, C. mangle, C. hrasiliana, and
C. virginica varies from 12 to 13, reaching 15 in C. gigas.
Based on the similarity of these numbers, this parameter
seems to be of little use in distinguishing the examined
species. In all species of Saccostrea studied here, the
variation is of 12-14 filaments per fold. The gills of all
species in this study occupy about 14 of the pallia! cavity
area. Saccostrea cucullata has a medium-sized alimen¬
tary chamber, approximately 1/3 as thick as the gill fold;
S. echinata has a thick chamber, about 14 as thick as the
fold; S. palmula, S. mordax, and S. glomerata have narrow
alimentary chambers, about 1/5 as thick as the gill fold.
Visceral Mass: An oyster is composed of two major
interconnected subdivisions: 1) visceral mass, containing
all the organ systems (digestive, excretory, reproductive),
most of the muscles, and much of the nervous and circu¬
latory systems; 2) gills and mantle/shell, which is com¬
posed of a hard portion (shell) that protects all other
organs and soft parts, the mantle, which carries sensitive
organs, as well as lesser parts of musculature, nervous and
circulatory systems (Stenzel, 1971).
The visceral mass extends from the ventral region of
the ligament to the anterior surface of the adductor muscle.
It is formed by normally yellowish-colored gonads, with
abundant digestive diverticula and renopericardial struc¬
tures, and occupies nearly 14 of the total body size in all
species. The gonads of all species are massive, apparently
regardless of the gonadal maturation period, and sur¬
round the whole visceral mass, in some cases surround¬
ing the region near the pyloric process. They are usually
colored beige or yellowish. The genital opening has been
observed in S. cucidlata in the kidney and distinguishes
itself from the surrounding area by a brownish spot.
Pericardium/Heart: The heart is located in the peri¬
cardium, a thin-walled chamber between the visceral mass
and the adductor muscle. On the right side, the promial
chamber extends down over the heart region and the
mantle separates the pericardium wall from the shell
(Galtsoff, 1964). In most Lamellibranehia, the rectum per¬
forates the heart and passes through it, but in Ostrea, the
heart and rectum remain apart, die heart lying beneadi die
rectum (Awati and Rai, 1931).
Saccostrea cucullata has a slightly enlarged ventricle
with thick walls, of opaque- white color; the auricles are
diin-walled and translucent and bear external sac-like
structures. Saccostrea glomerata has a robust heart, with
a relatively large ventricle, colored light cream; the auricles
are elongated, robust and have the same color as the ven¬
tricle. Saccostrea echinata has a thin ventricle, normally
translucent, and two non-pigmented auricles, both more
elongated than in S. cucidlata. The ventricle is elongated
in S. palmula, slightly thick and whitish; the auricles are
small and non-pigmented. Saccostrea mordax has a large
ventricle, about 2/3 the size of the auricles, with whitish
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THE NAUTILUS, Vol. 130, No. 2
coloration; the auricles are translucent and elongated. The
heart size and coloration are variable in Crassostreinae.
Palps: The outline of the palps is hatchet-shaped and
slightly different in each genus (Stenzel, 1971). The
labial palps are large triangular flaps King in front of the
gills and attached to the visceral mass by the broad base
(Awati and Rai, 1931). The palps of Saccostrea cucullata
are large, spatuliform, have a folded anterior edge and a
slightly arched ventral region. In S. glomerata , the palps
occupy ~l/5 of the visceral mass area and are also
spatuliform; their internal surface bear plicae that extend
transversely to the edge. The palps of S. echinata are
small, arched, sickle-shaped, with folds along the ante¬
rior margin. The external palps are fused in the middle,
overlying the inner palps. In S. palmula , the palps are
larger than those of the remaining species, thin, with a
superior margin bearing folds. Saccostrea nio relax has
large and thick palps, with folds in the superior margin,
as is the case in S. echinata, and the external palps overly
the inner palps for about 1/3 of their length.
Esophagus: The mouth leads into a long esophagus;
the esophagus lumen is dorsoventrally compressed and
appears as a narrow slit in transverse sections. The lumen
becomes narrower as it passes backward and upward to
enter the dorsal region of the stomach (Awati and Rai,
1931). The esophagus is dorsoventrally compressed in all
the study species; in Saccostrea cucullata, it is short and
large, Vz of palps in size; in S. glomerata, it is long and
narrow, and also ~!4 of palps size. In S. echinata , it is long
and thin, and ~l/3 as long as the palps; in S. palmula, it is
elongated and thin, and ~2/3 of palps size; in S. mordax,
it is large and long, of same size as the palps.
Stomach: The stomach is a large sac occupying a central
position in the visceral mass. This is true also for O. edulis
(Yonge, 1926; Graham, 1949; Purchon, 1957; Reid, 1965),
C. parasitica (Shaw and Battle, 1957) and O. chilensis
(Purchon, 1957). Other details of the morphology and
physiology of the digestive tract in Ostreidae, such as
ciliary mechanisms and stomach structure and operation,
can be found in the literature (Shaw and Battle, 1957;
Nelson, I960; Reid, 1965; Yonge, 1966). Internally, the
study species differ in the selection area 1 (sal), typhlosole,
gastric shield, and dorsal hood. The sal in Saccostrea
cucullata is short and large, with the typhlosole beginning
in the posterior region of the stomach; S. echinata has a
longer sal, and the typhlosole begins more posteriorly
than in S. cucullata. In S. palmula, the sal is short and
narrow, with the typhlosole beginning laterally. The sal in
S. mordax is long and narrow and the typhlosole is located
more posteriorly than in S. cucullata.
The stomach has been described as bag-shaped, inter¬
nally divided into a smaller anterior chamber and other
larger posterior chamber, by means of a thick fold; such a
fold is projected to the organ s lumen and seems to serve as
a means to direct particles to the posterior region through
a narrow channel (Shaw and Battle, 1957). The dorsal
hood length is nearly Vz of the length of the sal in
S. cucullata. In S. mordax, the dorsal hood has about the
same length as the sal. In S. echinata and S. palmula, the
dorsal hood is short and narrow, about Vz the size of those
of other species.
Intestine, Rectum, Anus: The mid-gut, or intestine,
originates from the stomach on the right side of the style-
sac, turns around immediately to its left, and runs parallel
to it all along its length; on reaching the distal end of the
style-sac, the intestine turns anteriorly and runs completely
back on this course (Awati and Rai, 1931). The intestine,
before the style sac, passes behind the pericardium,
immersed in the gonads in the visceral mass. In Saccostrea
cucullata, S. echinata, S. mordax, and S. glomerata, the
intestine passes behind the pericardium; in S. palmula,
behind the esophagus. The rectum of S. cucullata is long
and thin, surrounding the adductor muscle in 2/3 of its
concave region, with the anus bearing a single fold.
Saccostrea echinata has a larger elongated rectum, with a
shell-like anal expansion; S. palmula it is short and large,
with the anus ending in a bell -shaped fold. Saccostrea
mordax has a short and large rectum, and the anus bears
a single fold.
In all living oysters, except for the Pycnodontinae, the
rectum skirts the dorsal flank of the pericardium. In the
Pycnodontinae, the rectum passes through the pericardium
and the ventricle of the heart, and the anal papilla is more
projected. These relative positions of the rectum and
ventricle in oysters are of great taxonomic importance.
Central Nervous System: The nervous system is rel¬
atively simple, showing a pair of cerebral ganglia and a
pair of visceral ganglia connected by commissures. The
cerebral ganglia are somewhat inconspicuous because of
their very small size and location in the visceral mass.
The visceral ganglia can be easily distinguished in the
anterolateral region of the adductor muscle. Saccostrea
cucullata and S. glomerata have two pairs of nerves in
the anterior region and three in the posterior region;
S. echinata presents two pairs in the anterior and poste¬
rior regions; S. mordax and S. palmula have three pairs
in the anterior and posterior regions.
LITERATURE CITED
Abbott R.T. 1974. American Seas hells. 2nd edition. Van Nostrand
Reinhold, New York, 663 pp.
Awati, P.R. and H.S. Rai. 1931. Ostrea cucullata (the Bombay
oyster). Indian Zoological Memoirs 3: 1-107.
Born, 1., 1778. Index Rerum Naturalium Musei Caesarei
Vindobonensis, pars prima, Testacea. Ex Officina Krausiana,
Vindobonae, 442 pp.
Born, I., 1780. Testacea Musei Caesarei Vindobonensis.
Sumptibus Joannis Pauli Kraus, Vindobonae, 458 pp.
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naturelle des animaux sans vertebres. Deuxieme edition,
revue et auginente des notes par G.P. Deshayes et H.
Milne Edwards. Paris, 580 pp.
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Dillwyn L.W. 1817. A descriptive catalogue of recent shells.
John & Arthur Arch, Comhill, pp. 1-2, xii, 1-580, iv, 581-1121.
Gmelin, J.F. 1791. Caroli a Linne Systema naturae per regna
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Pp. 3021-3910.
Gould, A. A., 1850. Shells of the United States Exploring Expe¬
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THE NAUTILUS 130(2):72-78, 2016
Page 72
A new species of Cuthona Alder and Hancock, 1855 (Gastropoda;
Heterobranchia: Nndibranchia: Tergipedidae)
from the Caribbean Sea
Angel Valdes
Sabrina Medrano
Department of Biological Sciences
California State Polytechnic University
3801 West Temple Avenue
Pomona, CA 91768 USA
[email protected]
Vishal Shave
Bombay Natural History Society
S.B. Singh Road, Mumbai
Maharashtra 400001, INDIA
ABSTRACT
Tergipedid nudibranch specimens from the Caribbean Sea
previously identified as Cuthona caerulea are here described
as a new species. Sequence data for the mitochondrial COl
and 16S genes as well as the nuclear H3 are provided. A pre¬
liminary molecular phylogeny including other Cuthona species
available in GenBank produced inconclusive results, but the
new species is morphologically distinct from European specimens
of Cuthona caerulea. Differences include radular teeth and
reproductive morphology, as well as the external coloration.
INTRODUCTION
Cuthona Alder and Hancock, 1855 is a group of tergipedid
nudibranchs characterized by having crowded rows of
cerata, a broad foot, and arch-shaped radular teeth
(Miller, 1977). Species of Cuthona feed on hydroids,
and are most diverse in tropical and subtropical regions
(Gosliner, 1981). Cuthona is taxonomically complex, and
its taxonomic placement in relation to other genera in
the Tergipedidae is controversial (Williams and Gosliner,
1979; Gosliner and Griffiths 1981; Miller; 1977; Brown,
1980; Miller, 2004).
Cuthona caerulea is a northeastern Atlantic species char¬
acterized by having a white body with numerous cerata
with blue (or green) and yellow (or orange) pigment. The
coloration of this species is extremely variable, but speci¬
mens with distinct color patterns are morphologically
similar and regarded as members of the same species
(Thompson and Brown, 1984).
Thompson and Brown (1984) reported this species for
the first time from the western Atlantic, based on speci¬
mens collected from Florida, as well as records from
Sao Paulo, Brazil (based on a personal communication by
Ev. Marcus). Later, another specimen from Florida was
illustrated by Valdes et al. (2006). The western Atlantic
animals display some differences in color pattern in com¬
parison to the European ones, and some authors consid¬
ered the former to belong to an undescribed species
(Picton and Morrow, 1994; Calado, 2002).
In this paper we examined additional specimens recently
collected in Bocas del Toro, Panama, which are externally
similar to the animals illustrated from Florida by Valdes
et al. (2006). These animals were found to be distinct from
C. caerulea and are herein described as a new species.
MATERIALS AND METHODS
Specimen Collection: Four specimens were collected
on unidentified hydroids at 1 m depth in Crawl Cay,
Bocas del Toro, Panama, on July 30, 2015. Two specimens
were preserved in ethanol 95% and two in RNAlater. The
type material is deposited at the Museo de Malacologia,
Universidad de Panama (MUMAUP) and the Natural
History Museum of Los Angeles County (LACM).
Morphological Examination: One specimen (paratype)
was dissected. The buccal mass was extracted and placed
in a small glass container with NaOH 10% water solution
for 60 min until the tissue was soft. The jaws were then
removed and placed in ultrapure water for 5 min. The
radula and remaining tissue was left for another 24 hrs.
After tills period, the radula was also removed from the
NaOH solution and placed in ultrapure water for 5 min.
The radula and jaws were mounted on a stub for scanning
electron microscope (SEM) examination. The stub with
the samples was coated with an Emitech K550x sputter
coate > at the Natural History Museum of Los Angeles
County. The samples were examined under a Jeol
JSM-601G variable pressure SEM at the California State
Polytechnic U niversity.
A. Valdes et al, 2016
Page 73
The reproductive system was dissected from the
paratype (LACM 3335), examined under a dissecting
microscope (Nikon SMZ-100), and drawn with the aid
of a camera lucida attachment. The penis was removed
from the rest of the reproductive system and placed In
1 mL of hexamethyidisiiazane until all the liquid evapo¬
rated. The dry and hardened penis was then mounted on
a stub and sputter coated for SEM examination.
DMA Amplification and Sequencing: DNA from the
paratype was sequenced for this study. DNA extractions
were performed using approximately 1-3 mg of tissue
taken from the foot of the animal, followed by a hot
Chelex® extraction protocol with minor modifications.
The tissue sample was placed into a 1.7-mL tube contain¬
ing 1.0 mLTE buffer (10 mM Tris, 1 mM EDTA, pH 7.8)
and incubated overnight at room temperature in a rotator.
The sample was centrifuged for 3 min at 21,130 g. Subse¬
quently, 975 pL of the original 1 mL of TE buffer was
removed without disturbing the pellet of tissue. Then,
175 p.L of Chelex® solution was added and heated in a
56°C water bath for 20 min and placed in a 100°C heating
block for 8 min. The supernatant was the final product
used for the polymerase chain reaction (PCR).
PCK was used to amplify portions of the mitochondrial
cytochrome c oxidase 1 (COl) and 16S ribosomal RNA
(16S) genes, as well as the nuclear histone 3 (H3) gene.
The following universal primers were used to amplify the
fragments of interest: COl (LCQ149Q 5'-GGTCAAC
AAATCATAAAGATATTGG-3', HC02198 5'TAAACTT
CAGGGTGACCAAAAAATCA-3' developed by Folmer
et al, 1994), 16S rRNA (16S ar-L 5'-CGCCTGTTTAT
CAAAAACAT-3', 16S br-H S'-CCGGTCTGAACTCAG
ATCACGT-3' developed by Palumbi, 1996) and H3 (H3
AF S'-ATGGCTCGTACCAAGCAGACGGC-S', 113 AR
S'-ATATCCTTGGGCATGATGGTGAC-S' developed by
Colgan et al., 1998). Confirmation of amplification was
carried out using agarose gel electrophoresis with ethidium
bromide to detect the presence of DNA. PCR prod¬
ucts were sent to Source RioScience (Santa Fe Springs,
California, USA) for sequencing. Sequences were assem¬
bled and edited using Geneious Pro R8 (http://www
.geneious.com, Kearse et al. 2012). The sequences obtained
were deposited in Gen Bank, under the accession num¬
bers presented in Table 1.
Phylogenetic Analyses: Phylogenetic analyses were run
with the new sequences obtained and a data set of other
species of Cuthona compiled from GenRank (Table 1).
Phylogenetic analyses were conducted for all genes
concatenated and each gene individually. The best-fit
models of evolution (GTR + G for COl, HKY + G for 16S,
HKY for H3, and GTR + I for the entire concatenated
data set) were determined using the Akaike information
criterion (Akaike, 1974) implemented in jModelTest
(Darriba et ah, 2012). A Bayesian analysis was conducted
with MrBayes 3.2 (Ronquist et ah, 2012), partitioned by
gene (unlinked). The Markov chain Monte Carlo analysis
was run with two runs of six chains for 10 million genera¬
tions, with sampling every 100 generations. The default
25% bum-in was applied before constructing the majority-
rule consensus tree. Convergence was confirmed by eye
THE NAUTILUS, Vol. 130, No. 2
Page 74
using the “Trace” function in Tracer 1.5 (Rambaut and
Drummond, 2007). Maximum likelihood analyses were
conducted for the entire concatenated alignment with
raxmlGUI 1.0 (Silvestro and Michalak, 2012) using the
bootstrap + consensus option (10,000 replicates) and the
GTR + I model.
RESULTS
The Bayesian consensus tree was relatively well-resolved( but
most nodes were not supported in tire maximum likelihood
tree (Figure 1). Only two clades, one including Cuthona
fulgens (MacFarland, 1966) from California and two uniden¬
tified species from Washington and tire Pliilippines, and
another including Cuthona divae (Er. Marcus, 1961) and
Cuthona concinna (Alder and Hancock, 1843), are well
supported. Additionally, the phylogenentie position of tire
specimens from Panama, sequenced here in relation to a spec¬
imen of Cuthona caendea from Europe, was not resolved.
Anatomical data revealed consistent differences between
Panamanian and European specimens. Therefore, the
taxon from Panama is below described as a new species.
The morphological differences are described in the Dis¬
cussion section.
SYSTEMATICS
Tergipedidae Bergh, 1889
Cuthona luciae new species
(Figures 2-11)
Cuthona caendea ( non Montagu, 1804). — Thompson
and Brown, 1984: 121; Valdes et al. 2006: 264, 265
Externa] Morphology: Live animals up to 12 mm
length. Body narrow, elongated (Figure 2). Cerata elon¬
gated, cylindrical, dorso-lateral, arranged in 13-14 verti¬
cal rows, with 4 — 5 cerata in each row. Oral tentacles
smooth. Rhinophores smooth, similar in length to oral
tentacles. Reproductive opening located on right side
of body, between first and second groups of cerata.
Anus acleiproctic, dorso-lateral, posterior to pericardium.
Body background color gray with irregular yellow spots.
Posterior of dorsum dark blue. Dense yellow spotting on
pericardium, behind second row of cerata. Rhinophores
100
0 98
0.86
0.81
100
0.58
0.91
0 99
Tergipes tergipes CASIZ 182699
Cuthona sp. 2 Antartica
0.03
0.94
Cuthona sp. 9 Antartica
0 66
0.86
Cuthona sp. 35 CASIZ 186436 Philippines
Cuthona Columbiana Canada
- Cuthona luciae Panama
Cuthona lagunae California
— — Cuthona sibogae
- - — - Cuthona caerulea North Sea
Cuthona sp. 7 Washington
0.51
Cuthona sp. A CASIZ 186435 Philippines
— Cuthona fulgens CASIZ 174484 California
- Cuthona sp 8 Washington
. Cuthona abronia CASIZ 174485 California
100
Cuthona divae CASIZ 174495 California
Cuthona concinna Washington
Cuthona ocellata Portugal
0.7 . .
0 93
Cuthona sp. 4 Antartica
Cuthona sp. 3 Antartica
- Cuthona sp. 1 Antartica
Cuthona sp 5 Antartica
- Cuthona sp. PW-2014 French Polynesia
Cuthona sp. 6 Antartica
Cuthona cocoachroma Washington
Figure 1. Cuthona. Bayesian consensus tree of the concatenated analysis including posterior probabilities and bootstrap values
from the maximum-likelihood analysis.
A. Valdes et al., 2018
Page 75
Figures 2—4. Cuthona luciae new species. Photographs of live holotype (MUMAUP MOL-GAS-O01). 2. Dorsal view of animal on
its hydroid prey, with egg mass visible. 3. Lateral view of head. 4. Dorsal view on black background.
opaque gray with while mid-region and orange apices.
Yellow pigment on head, surrounding base of rhinophores.
Cerata opaque yellow with blue band toward distal
end before reverting to yellow. Oral tentacles opaque
yellow with dark orange tips. Blue transverse band
connecting bases of oral tentacles. Anterior end of head
dark orange.
Anatomy: Radular formula 68 x 0.1.0 in 12 mm pre¬
served length paratype (LACM 3335). Radular teeth
with 9-10 large, sharp denticles, which decrease in size
toward lateral sides of teeth and again toward center
(Figure 5). Denticles separated by gaps, which become
wider towards center of teeth. Gaps filled with tiny, sharp
denticles, which vary in number depending on width of
gap and are absent from most lateral gaps. Cusp about
same length, or shorter, than central denticles, and only
distinguishable from denticles because it emerges from
slightly higher plane. Jaws elongate (Figure 8) with smooth
masticatory borders (Figures 7-8).
Reproductive system (Figure 10) with an elongate
ampulla connecting directly into female gland com¬
plex. Prostate emerges from female gland complex,
near insertion point of ampulla. Prostate long and con¬
voluted, narrowing abruptly at distal end, to expand
again into deferent duct. Distal portion of deferent duct
containing large penis with apical stylet (Figures 9, 11).
Vagina slightly curved, connecting directly into rounded
bursa copulatix.
Type Materials HOLOTYPE: MUMAUP MGL-GAS-
001, July 30, 2015; PARATYPE: LACM 3335, July 30,
2015; all from type locality.
Type Locality: Crawl Cay, Bocas del Toro, Panama.
Geographic Range: Florida (Thompson and Brown
1984, Valdes et al. 2006) to Panama (present paper) and
possibly Brazil (Thompson and Brown 1984).
Page 76
THE NAUTILUS, Vol. 130, No. 2
Figures 5-9. Cuthona luciae new species. Scanning electron micrographs of radular teeth, jaws, and penis of paratype
(LACM 3335). 5. Radular teeth. 6. Jaw. 7. Dorsal view of the masticatory border. 8. Ventral view of the masticatory border. 9. Penis.
Etymology: Named after Lucia Valdes, daughter of the
senior author.
DISCUSSION
The phylogenetic analyses resulted in poorly supported
trees. Although the Bayesian consensus tree contains
well-supported nodes, many of those are not supported
in the maximum likelihood consensus tree. This study
has not produced a reliable phylogeny for the species of
Cuthona sequenced to date. The results of this phyloge¬
netic analysis are also inconclusive as to the position of
the species here described relative to a European specimen
of Cuthona caerulea. However, a Bast-n search in Gen Bank
revealed that the COI sequence of C. caerulea from the
North Sea in GenBank (AF249807) and the sequence
from C. luciae are only 82% identical, which is consistent
with species-level differences. In addition, the morpho¬
logical examinations revealed the presence of several
unique characteristics that support that the Caribbean
animals constitute a distinct species.
The radular teeth of Cuthona luciae are very different
from those of C. caerulea described from European spec¬
imens. Schmekel and Portman (1982) illustrated three
radular teeth in lateral view of a specimen collected in
Naples, Italy. These teeth had 6 lateral denticles of similar
size and a huger central cusp. Thompson and Brown
(1984) illustrated one radular tooth of a specimen from
Lundy, England, which had 5 lateral denticles, but was
odierwise similar to tire Mediterranean radula illustrated
by Schmekel and Portman (1982). These radulae are very
different from the Caribbean specimens here examined,
in which the teeth contain denticles separated by gaps
filled with tiny, sharp denticles, varying in number
depending on the width of the gap. Additionally, the jaws
of European specimens have a distinct masticatory
border with denticles (Thompson and Brown, 1984), which
is absent in the Caribbean animals, although Schmekel
and Portman (1982) reported that it can be absent in
Mediterranean specimens as well.
Schmekel and Portman (1982) illustrated the reproduc¬
tive system of a specimen from Naples, Italy. Although the
reproductive system of the specimen examined from the
Caribbean is similar, there are two fundamental differ¬
ences, the European specimens have a well-formed penial
gland, absent in the Caribbean animal; in addition, the
Caribbean animal has a penial stylet, which is not reported
in the European specimen.
A similar species to Cuthona luciae is Cuthona herrerai
Ortea, Moro, and Caballer, 2001, originally described
from Cape Verde, Eastern Atlantic. The radular teeth of
A. Valdes et al., 2016
Page 77
of the two species bear similar colors, they are much
brighter in C. luciae.
Also, Cuthona iris Edmunds and Just, 1983, originally
described from Barbados, has a similar color pattern with
yellow cerata, each with a blue band (see Valdes et al.
[2006] for a color illustration). But the body of this species
is predominantly yellow, including the rhinophores and
oral tentacles, it has a light blue dorsal band, absent in
C. luciae , and lacks the characteristic head pigmentation
of C. luciae.
Although morphological evidence confirmed that
Cuthona luciae is distinct from C. caeurela and other
similar species such as C. herrerai and C. iris, further
research is necessary to determine the phylogenetic posi¬
tion of C. luciae and to resolve the evolutionary relation¬
ships within Tergipedidae. The sequence data provided
here should facilitate future work toward these goals.
0.5 mm
Figures 10, 11. Cuthona luciae new species. Reproductive
system of paratype (LACM 3335). 10. Dorsal view of the repro¬
ductive system. 11. Detail of the penis. Abbreviations: am,
ampulla; be, bursa copulatrix; dd, deferent duct; fgc, female
gland complex; pe, penis; pr, prostate; s, penial stylet.
C. herrerai are very similar to those of C. luciae in
having large denticles separated by gaps containing
tiny denticles (Ortea et al. 2001). However, many other
characteristics differentiate these two species, for
example the jaws of C. herrerai contain denticles on the
masticatory border, absent in C. luciae-, C. herrerai
has less rows of cerata and less cerata per row than
C. luciae-, more importantly, C. herrerai lacks orange
pigment on the oral tentacles and the characteristic bright
blue band on the head of C. luciae. Although the cerata
ACKNOWLEDGMENTS
The SEM work was conducted at the California State
Polytechnic University SEM laboratory supported by
the US National Science Foundation (NSF) grant DMR-
1429674. The fieldwork in Panama was conducted during
the course “Taxonomy and Biology of Sea slugs” funded
by awards NSF DEB-1355190 and DEB-1355177, and
organized by Rachel Collin of the Smithsonian Tropical
Research Institute in Bocas del Toro. The travel expenses
of AV and SB were also supported by the NSF, and
those of VB by the British Ecological Society (travel
grant 5585-6629), and the Bombay Natural History
Society. Several staff members of the Smithsonian Tropi¬
cal Research Institute in Bocas del Toro facilitated the
fieldwork activities: Plinio Gondola, Urania Gonzalez,
Tanyusha Grenald, Arcadio Castillo, Deyvis Gonzalez,
and Nerea Nieto. Collecting of specimens was possible
with permits ARAP 27 (2015) and ANAM SE/A-64-15
issued by the Panamanian Government.
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Valdes, A., J. Hamann, D.W. Behrens, and A. DuPont. 2006.
Caribbean Sea Slugs: A Field Guide to the Opisthohranch
Mollusks from the Tropical Northwestern Atlantic. Sea
Challengers Natural History Books, etc, Gig Harbor,
Washington, 289 pp.
Williams, G.C. and T.M. Gosliner. 1979. Two new species of
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America, with a revision of the family Cuthonidae. Zoo¬
logical Journal of the Linnean Society 67: 203-223.
THE NAUTILUS 130(2):79-81, 2016
Page 79
The genera Miraclathurella Woodring, 1928 (Gastropoda:
Pseudomelatomidae) and Darrylia Garcia, 2008 (Gastropoda:
PHoraiclavidae), with two proposed new combinations for Darrylia
Emilio F. Garcia
1 15 Oak Crest Dr.
Lafayette, LA 70503
Efg21 [email protected]
ABSTRACT
The genera Miraclathurella Woodring, 1923 and Darrylia
Garcia, 2003 are compared. Miraclathurella clendenini and
M. peggijwilliamsae are reassigned to Darrylia.
INTRODUCTION
Miraclathurella Woodring, 1928 and Darrylia Garcia,
2008 are two genera that are similar in conchological
characters. However, Bouchet et al. (2011) have placed
Miraclathurella in the family Pseudomelatomidae and
tentatively placed Darrylia in the recently erected family
Horaiclavidae. According to Bouchet et al. (2011: 293),
the family Horaclaividae “shares many characters with
Pseudomelatomidae, conchologically differing by a small
stout shell with short siphonai canal and usually poorly
developed spiral sculpture”. Puillandre et al. (2001) further
define Horaiclavidae, stating that it “mostly includes genera
previously placed in the Drilliidae and Pseudome¬
latomidae.” Molecular data clearly show that the included
genera are distinct in these families and correspond to a
highly supported clade, currently arranged as a sister
clade to the Clavatulidae (Puillandre, 2011: 269). Unfor¬
tunately, molecular data has not been available for either
Miraclathurella or Darrylia.
The genus Miraclathurella was proposed by Woodring
(1928: 191) to accommodate turrid-like mollusks with,
among other characters, a stout, broad-tipped nucleus
“consisting between two and a half and three whorls,
the end of the last whorl bearing a few coarse curved,
protractive axial riblets.” The aperture is “very long and
narrow”, and the anterior canal, “relatively long.” Woodring
named two species in Miraclathurella : M. vittata (Figure 1),
with a protoconch of “about three whorls, about the last
half whorl bearing an anterior keel, behind which lie axial
riblets,” and M. eniemna (Figure 2), with a protoconch
“consisting of about two and a half whorls, the last quarter
whorl bearing an obscure anterior keel, behind which lie a
few obscure axial riblets.” Although Woodring considered
the Bowden beds to be of Miocene origin, studies of the
foraminifers in the area have proved them to be younger,
from the Pliocene Epoch (Donovan, 1998).
Use of the genus Miraclathurella remained restricted to
fossil species from Bowden described by Woodring until
1971, when Shasky described Miraclathurella mendozana
from the recent fauna of the Panamic Province. In his
Discussion (Shasky, 1971: 68), that author compared this
species to Clathrodrillia woodringi Pilsbry and Olsson,
1941, a species described as a Pliocene fossil from the
Canoa Formation of Ecuador, which Shasky also assigned
to Miraclathurella. Also in 1971, Keen transferred
Pleurotorm bicanalifera Sowerby, 1934 to Miraclathurella
(Keen, 1971: 728, fig. 1719).
The genus Miraclathurella did not appear in the lit¬
erature of the recent fauna of the western Atlantic until
1988, when Jong and Coomans (1988) transferred Drillia
kleinrosa Nowell-Usticke, 1969, from the shallow waters
around Aruba, Bonaire and Curasao, to Miraclathurella. And
two decades later a second species, M. clendenini Garcia,
2008, from Bahia de Campeche, was tentatively placed in
that genus. A third species, M. peggy williamsae Fallon,
2010, from St. Vincent and the Grenadines soon followed.
Although Jong and Coomans placed kleinrosa in
Miraclathurella, the species did not seem to fit Woodring’s
description of that genus, lacking any vestige of a keel on
the protoconch and possessing a short, not long, siphonai
canal so when a second species with the same differ¬
entiating characters was discovered in Roatan Island,
Honduras, there seemed to he a need to create a taxon that
would include those two species, and the genus Darrylia
was proposed.
DISCUSSION
Darrylia differs from Miraclathurella in having more
solid, more stout shells; a paucispiral nucleus of less than
two whorls that are almost completely, or partially, axially
ribbed; a short aperture which occupies no more than
Page 80
THE NAUTILUS, Voi 130, No. 2
Figures 1-10. Species of Miraclathurella and Darrylia. 1. Miraclathurella vittata Woodring, 1928, Holotype, USNM 135376,
length 12 mm, width 4 mm, Bowden Formation, Jamaica. 2. Miraclathurella entemna Woodring, 1928, Holotype, USNM 135374,
length 15.5 mm, width 4.7 mm, Bowden Formation, Jamaica. 3, 4. Darrylia harryleei Garcia, 2008, Holotype, ANSP 416409, length
5.9 mm, width 2.1 mm, Oakridge, Roatan I., Honduras, 0.2 m depth. 5, 6. Darrylia kleinrosa (Nowell-Usticke, 1969), EFG 13924,
length 6.5 mm, off Hotel Bonaire, Bonaire 1., Netherlands Antilles, 2. 5-3.0 m depth. 7, 8. Darrylia clendenini (Garcia, 2008),
Holotype ANSF 416411, length 8.2 mm, width 3.2 mm, Bahia de Campeche, southwestern Gulf of Mexico, 20°51.49' N, 92°21.44' W,
63-65 m depth. 9, 10. Darrylia peggijwilliamsae (Fallon, 2010), Holotype USNM 1139716, length 10.3 nun, width 3.6 mm, Baliceaux
Island, St. Vincent and the Grenadines, Caribbean Sea.
E.F. Garcia, 2016
Page 81
one third the length of the shell; and a truncated shell
with a shorter anterior canal. When the genus Darrylia
was proposed, only D. harryleei (Figures 3, 4) and
D. kleinrosa (Figures 5, 6) were considered. Despite its
obvious similarities with Darrylia, the species clendenini
(Figures 7-8) was tentatively placed in Miraclathurella
mainly because of the presence of (1) a subsutural cord
and (2) a somewhat longer anterior canal. However,
the discovery of “ Miraclathurella ” peggywilliamsae
(Figures 9-10), which also has a subsutural cord, and
which otherwise fits well in Darrylia, has led me to
conclude that some members of the genus Darrylia do
possess a subsutural cord. One may point out that the
almost completely ribbed protoconch of D. clendenini
(Figure 4) does compare well with that of D. harryleei
(Figure 8), the type species of Darrylia-, and that the first
smooth, then ribbed protoconch of D. peggywillianisae
(Figure 10) reflects the characters of the protoconch
of D. kleinrosa (Figure 6).
Comparing the plate images of Miraclathurella spp.
(Figures 1-2) and Darrylia spp. (Figures 3, 5, 7, 9) they
illustrate the differences between the two genera.
Darrylia clendenini still seems to show a somewhat more
elongated anterior canal, but the specimen figured, as
well as all other specimens from the type material, were
sub-adult, and the outer lip had not yet thickened, which
gives it that seemingly uncharacteristic appearance.
Drawing from the considerations above, I propose
that Miraclathurella clendenini Garcia, 2008 and
M. peggywilliamsae Fallon, 2010 be assigned to Darrylia.
Two recent species from the Panamic Province have
been placed in Miraclathurella, However, Keen’s new
combination of M. bicanalifera is believed to be in error,
as the species characters are more similar to those of
Glyphostoma. Glyphostoma epicasta Bartsch, 1934 is
its western Atlantic cognate. Compare Keen (1971:726,
fig. 11) with Bartsch (1934, pi. 4, figures 4, 7, 9), and with
the figure at http://www.jaxshells.org/glypepicj.htm.
The multispiral protoconchs in Mirachlaturella species
indicate planhtotrophie larval development and resulting
broader distributions for these species. This could explain
the presence of a broadly distributed ancestor to both
the new species and the recent eastern Pacific species
M.mendozana. Although Shasky described the protoconch
as “smooth” (Shasky, 1971: 68), without the axial riblets
described by Woodring, the shell does have all the other
characters of Miraclathurella.
In the western Atlantic, the genus Miraclathurella
seems to be restricted to the Pliocene beds at Bowden,
Jamaica. The genus Darrylia, characterized by a
paucispiral protoconch, has so far produced four distinct
populations: Darrylia peggywilliamsae in the St. Vincent
Grenadines area in the southeastern Caribbean,
D. kleinrosa around the ABC islands, Netherlands Antil¬
les, in the southwestern Caribbean, D. harryleei in the
Bay Islands, Honduras, and D. clendenini, in the south¬
western Gulf of Mexico.
ACKNOWLEDGMENTS
My thanks to Phillip Fallon for allowing me to reproduce die
images of the holotype of Miraclathurella peggywilliamsae;
he has also critically read the ms, providing information
that improved the quality of the paper. The figures of
Miraclathurella vittata and M. entemna are reproduced
from Woodring (1928: pi. 8, figures 2 and 5).
LITERATURE CITED
Bartsch, P. 1934. Reports on the collections obtained by the
first Johnson-Smithsonian Deep-sea Expedition to the
Puerto Rican Deep. New mollusks of the family Turritidae.
Smithsonian Miscellaneous Collections 91(2): 1-29, 8 pis.
Bouchet, R, Yu. I. Kantor, A. Sysoev, and N. Puillandre. 2011.
A new operational classification of the Conoidea. Journal
of Molluscan Studies 77: 273-308.
Donovan, S.K. 1998. An introduction to the Bowden Shell Bed,
Southeast Jamaica. Contributions to Tertiary and Quater¬
nary Geology 35: 3-8, 4 figs.
Fallon P.J. 2010 Descriptions and illustrations of some new and
poorly known turrids of the tropical northwestern Atlantic.
Part 1. Genera Buchema Corea, 1934 and Miraclathurella
Woodring, 1928 (Gastropoda: Turridae: Crassispirinae).
The Nautilus 124: 166-174.
Garcia, E.F. 2008. Eight new molluscan species (Gastropoda:
Turridae) from the western Atlantic, with the description
of two new genera. Novapex 9: 1-15.
Jong, K.M. de and H.E. Coomans. 1988. Marine gastropods
from Curagao, Aruba and Bonaire. Studies on the Fauna
of Curagao and other Caribbean Islands 69: 1-261, 47 pis.
Pilsbry, H.A. and A. A. Olsson. 1941. A Pliocene fauna from
western Ecuador. Proceedings of the Academy of Natural
Sciences of Philadelphia 93: 1-79, pis. 1-19.
Puillandre, N., Yu. I. Kantor, A. Sysoev, A. Couloux, C. Meyer,
T. Rawlings, J.A. Todd and P. Bouchet. 2001. The dragon
tamed? A molecular phylogeny of the Conoidea Journal of
Molluscan Studies 77: 259-272.
Shasky, D.R. 1971. Ten new species of tropical eastern Pacific
Turridae. Veliger 14: 67-72.
Nowell-Ustieke, G.W. 1969. A Supplementary Listing of New
Shells, to be added to the Check List of the Marine Shells
of St. Croix. Author edition, St. Croix, 32 pp., 6 pis.
Woodring, W.P 1928. Miocene Mollusks from Bowden,
Jamaica. Part II. Gastropods and discussion of results.
Carnegie Institute of Washington Publication 385, vii +
564 pp., 40 pis.
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