Descriptions of eighteen new species in the genus Granulina Jousseaume, 1888 (Gastropoda: Cystiscidae) from the Caribbean Sea

2010·18.9 MB·English

by  Tony Mccleery

.
T. McCleery NOVAPEX 1 1(2-3): 37-71, 10juin 2010
MCZ
Descriptions ofeighteen new species in the genus Granulina
LIBRARY
Jousseaume, 1888 (Gastropoda: Cystiscidae)
from the Caribbean Sea JL 08
2010
Tony McCleery HARVARD
"Mantaray", Bascombe Road, Churston Ferrers, Brixham, Devon TQ5 OJJ, Englan^JfsjlVERSITY
[email protected]
KEYWORDS. Cystiscidae, Granulina, Caribbean Sea, new species, surface texture, résorption
ABSTRACT. A list ofail described Caribbean species in the genus Granulina is given. Eighteen
new species of Granulina from the southem Caribbean Sea are described for the first time, four
from eastern Panama: G. colonensis, G. darienensis, G. ocella, G. waltergomezi; five from
Colombia: G. cartagenaensis, G. gayracaensis, G. granatensis. G.pinguisa, G. velaensis; six from
Venezuela: G. calla, G. iridisa. G. monjesensis, G. nivalis, G. ovata, G. volcana; one from Aruba:
G. plagula; one from Curaçao: G. producera and one from Trinidad and Tobago: G. tobagoensis
Biodiversity and features of the genus are discussed and previously undescribed features are
presented.
INTRODUCTION Granulina amianta (Dali, 1889), USA, North
Carolina, 26-95 m.
The genus Granulina Jousseaume, 1888, has a World- Granulina antillensis (de Jong and Coomans, 1988),
wide distribution in tropical and subtropical seas. Aruba and Curaçao, 50 m.
Most commonly found in littoral and sub littoral Granulina hadria (Dali, 1889), U.S.A., Florida, Cedar
depths, it has also been recorded from abyssal depths Hill, shallow.
down to 1,285 m, with a single record at 1,700 m Granulina lachrimula (Gould, 1862), USA, Georgia
(Coovert and Coovert, 1995: 74). and Florida, 260-732 m.
Caribbean Sea is used herein to include south Granulina ovuliformis (d'Orbigny, 1842), Gulf of
eastern U.S.A. south of Georgia and Bahamas in the Mexico and Caribbean, moderately deep water.
north, Trinidad and Tobago in the south east. Granulina tinolia (Dali, 1927), USA, north east
In some Cystiscidae gênera Caribbean species do Florida and Georgia. 538 m.
not compare well with the type species - the gênera Since then five more species hâve been described,
Persicula and Gibberula are examples. The genus ail by Espinosa and Ortea, (2000, 2003, 2005),
Granulina appears to be reasonably homogeneous and bringing the présenttotal to twelve:
ail described Granulina species known to the author Granulina aidae Espinosa & Ortea, 2005, Cuba, Pinar
compare well with the type species Granulina isseli del Rio, 25-30 m.
(G. & H. Nevill, 1875), from the Red Sea coast of Granulina guanajatabey Espinosa & Ortea, 2003,
mm
Egypt. Size range forthe genus is given as 0.8-3.2 Cuba, Pinardel Rio, 1-2 m.
(Coovert and Coovert, 1995: 73). Caribbean Granulina lazaroi Espinosa & Ortea, 2005, Cuba,
Granulina collected by the author range from 1.2 - Pinardel Rio, 25-30 m.
3.42 mm. Granulina minae Espinosa & Ortea, 2000, Costa Rica,
A total ofapproximately 75 species of Granulina Manzanilla, 12-15 m.
hâve been described, the majority from north west Granulina molinai Espinosa & Ortea, 2005, Cuba,
Africa and the Mediterranean (approximately 35 Pinar del Rio, 25-30 m.
species). This is foUowed by the Pacific and Indian
Océans (approximately 20 species), and the Caribbean The eighteen new species described herein were
Sea with only twelve species described to date. Thèse collected by the author during the past six years. Five
figures fail to indicate the true diversity ofthis genus; were collected in shallow water down to 10 m on and
more probably they reflect the historical, low level of around dead coral rocks and rubble, eleven by
interest in micro-moUuscs, including families dredging in depths between 18 m and 130 m on sand
Cystiscidae and Marginellidae. This article will deal and mud substrates. The genus was found to be well
with eighteen new species of Granulina from the represented in ail areas sampled at thèse depths.
southem Caribbean Sea, where there appears to be a Undescribed Granulina continued to be found
very large numberof, as yet, undescribed species. regularly indicating that there are probably many more
Preceding the year 2000 only seven recognised species remaining to be discovered. Of the eighteen,
Caribbean Granulina species had been described: fourteen are known only from the type locality.
W
G. agger (Watson, 1886), Indies, offCulebra, 714 Géographie ranges ofthe remaining fourare small.
m.
37
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Map. 1. Caribbean Sea, type localities ofthe new species
Key to map location numbers 9. Venezuela, Monjes del Sur, harbour,
12°21.5'N, 70°54.1'W, 3-lOm.
1. Panama, offColon, to east, 9°31.5'N, 10. Aruba, Boca Grandi, 12°27.3'N,
79°52.0'W, 57 m. 59°52.6'W, 1-2 m.
2. Panama, offIsla Chichime, San Blas, 11. Curaçao, offPiscadera Bay, 12°07.5'N,
9°37.3'N, 78°53.2'W, 75 m. 68°58.5'W, 130 m.
3. Panama, East Holandes Cays, San Blas, 12. Venezuela, Las Aves de Sotavento,
9°35'N, 078°40'W. 3 m. 12°01.66'N, 067°38.05'W, 1 m.
4. Colombia. offCartagena, 10°22.4'N, 13. Venezuela, Cabo Codera, 10°35.2'N,
75°35.8'W, 25-41 m. 66°03.9'W, 18 m.
5. Colombia, Gayraca Bay, Santa Martha, 14. Venezuela, offIsla Cubagua, to north,
iri9.5'N, 74°06.3'W, 7 m. 10°52.4'N, 64°12.4'W, 22 m.
6. Colombia, offSanta Martha, 1 1°18.0'N, 15. Venezuela, Isla Grande, Islas Los Testigos,
74°12.2-W, 90-101 m. 1I°22.8'N, 63°08.rW, 28 m.
7. Colombia, offCabo de Vêla, ! 1°57'N, 16. Venezuela, offIslas Los Testigos, to north,
72°36'W. 58 m. 11°28'N, 63°06'W, 73 m.
8. Colombia, offCabo de Vêla, 12°06.7'N, 17. Trinidad and Tobago, offTobago, to north,
72°19.3'W, 50-59m. 11°16'N, 60°49'W, 86 m.
Terminology. The author has, in gênerai, followed présence or absence ofa notch which is a separate and
terminology established by Coovert and Coovert distinct feature.
(1995). The terms ''siphonal notch" and "posterior
notch" hâve précise meanings and are clearly Shell morphology. Shell shape of Caribbean
described (Coovert and Coovert, 1995: 50). "Siphonal Gnmul'ma ranges between globose, pyriform and
canal" (Coovert and Coovert, 1995: 47), and posterior perfectly oval. AU are colourless but vary slightly in
canal are used by the author. but do not indicate the opacity; are generally, heavily callused; hâve strongly
38
T. McCleery NOVAPEX 11(2-3): 37-71, 10juin 2010
denticulate, curved, strongly curled in lip; four, Coovert and Coovert illustrated internai plications of
generally strong, columellar plications, almost always G. hadria (Dali, 1889), on which species they carried
excavated and intricately formed distally in a wide out a very thorough study. Therefore, it seems
variety of distinct shapes. AH hâve immersed spire, probable that G. hadria differs from many other
médium to very strong extemal varix extending Caribbean Gramilina in this respect. Doubt about the
around posterior and siphonal canals, and most hâve a accuracy of Coovert and Coovert's conclusions was
pariétal calius ridge extending posteriorly from the expressed by Boyer and Rolân (2004: 162), who
columellar plications to meet a lumpy posterior ridge. suggested that Coovert and Coovert had incorrectiy
Ail hâve Type 2 animal (Coovert and Coovert, 1995: interpreted their drawing. They stated that Gramilina
73), and Type 4 radulae (Coovert and Coovert, 1995: hâve fully developed coiling ofthe internai columellar
56). plications, but this is clearly contrary to Coovert and
Surfaces are minutely textured in a wide variety of Coovert's findings. Récent work by the author on a
forms, generally finest on light calius wash extending numberofCaribbean Gramilina sp. revealed that three
over the body whorl, and strongest on the calius plications remain, possibly somewhat reduced, for
around the siphonal and posterior canals (Figs 109- approximately one tum internally, and can frequently
132). Calius on body whoris is not well attached and be detected through the body whorl in fresh dead
very easily removed by light abrasion, confirming that shells (Fig. 55). A number of shells of Gramilina
it is indeed a wash and not micro-sculpture (Figs 139- tohagoensis n. sp. were opened to expose internai
141). When calius wash is removed faint growth Unes whoris and columellar plications (Figs 133-135). It
are often exposed (Fig. 141). In dead collected shells was found that a high degree of résorption had
calius wash on the body whorl was generally found to occurred, that résorption was complète posteriorly,
be absent due to abrasion. and that the plications were much reduced. It can be
it is well known that some Gronii/ina spp. hâve seen in thèse examples that résorption is complète
textured surfaces: Grainilina fenuuidesi Boyer & posteriorly (Figs 133) and in an early stage anteriorly
Rolân 1999, and G. aidae Espinosa & Ortea, 2005, are where the columella remains strong. Onejuvénile was
examples of shells with unusually strong texture for examined (Fig 135) and résorption was noted to be at
the genus. Boyer and Rolân (1999: l-IO) state "This an advanced stage posteriorly. Therefore, it is
feature is however unique and constant in each concluded that résorption commences early in the life
species, and well représentative of the whole genus, of the animal of G. tobagoensis. The sample was
even the type species G. isseli which represents itselfa small and much work remains to be carried out, but
faint "léopard patterned" microsculpture on a smooth thèse preliminaiy findings support those of Coovert
ground". Species which appear to be smooth and and Coovert.
shiny to the human eye were previously believed to be The shell morphology within the genus Gramilina
without texture, but récent S.E.M. work by the author is very variable and it is no surprise to find variations
confirms that probably ail Graniilina exhibit some in the précise way internai résorption takes place. The
degree of texture. It was observed that the fonn of important point in this matter is that significant
texture is variable over the surface. For example, in résorption and réduction of columellar plications do
Graniilina colonensis n. sp., surface of the posterior take place in Gramilina species. Partial résorption of
canal is evenly covered by scales, but calius on the internai whoris with reduced columellar plications is
body whorl, adjacent to the lumpy ridge bordering the the key feature to be considered in family assignment
posterior canal is, at first, comprised ofdistinct round, of ail marginelliform gênera (Coovert and Coovert,
evenly sized granules, changing abruptiy into less 1995:43).
regularly shaped and very variably sized granules
(Figs 110-112). The body whorl of Gramilina nivalis Animal chromatism. In Gramilina extemal animal
n. sp. was found to be lightly textured internally (Figs morphology and chromatism exhibit more features
136-138). It is not known ifthis feature is widespread than are seen in other Cystiscidae. Some of thèse
as only this species has been examined so far. It seems features hâve not previously been documented. For
probable that surface texture will be usefui in species example, the extrême distal elongation of the
identification but frst, more research is necessary in metapodium (Figs 12-13), and the posteriorpapillae in
order to establish the degree of both intra-population some species (Figs 5, 12, 35). On occasions, when
and inter-population variation which exists. photographing live animais an apparently rounded
Gramilina hâve partially resorbed internai whoris metapodium was observed to gradually extend and
which Coovert and Coovert (1995: 73) termed become long, thin and very finely elongate distally
"moditled cystiscid internai whoris". A drawing (Figs 12-13). Occasionally the distal portion is
presented by Coovert and Coovert (1995: 49, tlg. 4), transparent and unmarked and can be very diftlcult to
and their description: "columella with 2 internally observe (Fig. 14). Mantles are often only partially
reduced columellar plications, plus one or two pseudo- visible, seldom being seen fully extended, and are
continuous plications" (Coovert and Coovert,1995: very variable: surfaces can be smooth, pustulose, and
73) is somewhat confusing and is at variance with the occasionally bear long, plume-like posterior papillae
author's tmdings. An explanation for this may be that which are located on a thick, cap-like area, posterior
39
T. MtClkfry Eightccn now species o\'Gnuiuliiui
medially (Figs 16-17). Thèse papillac were nbseived and ollen becoming very strong in slieils of okl
to extcnd and relraet rapidi\: approxiinaleK une animais. For example, in Granidiiui, the callus around
seeond to fully cxlend lo a lengtli whieli oqualed with the posterior canal, and to a lesser extent around the
shell length, and a similar lime to fully retract. Il sipiional canal, appears to continue to grow
seems likeK that éléments whicli inllate papillae are througluHil the life of the animal, whereas growlii of
locaied in the eap-like area. Tlie nuniber ot" papillae labial denticles and columellar plications appears to
varies trom one to t'ive (Figs 12, 21, 35). Pustules also slow down or stabilise. It was observed that old shells
e.xtend and retraet, but were ne\erobser\ed to do so as hâve a iower W:L ratio than young adult shells. The
rapidly as the papillae (See "Remarks" under reason is that callus growth at both ends of maturing
description oïGninulina cartagenaensis n. sp.). When shells increases shell length more than width.
magnitled. the mantle chromatism was observed to be It appears that the intricate and complicated
comprised of extremely small spots amongst more émergent parts ofcolumellar plications are associated
prominent features (Figs 30-32). A common feature of with the pariétal callus ridge (Figs 92-102) because,
shallow water. rock dwelling species is large ocellated lumps and kinks which occur, particularly on the
spots, most noticeable when the mantle is extended second and third plication, are located on the projected
(Fig. 31). Ocellated spots were not observed on Une ofthe pariétal ridge, and also because excavation
species dredged in deep water from mud or sand ofcolumellar plications is aligned with the inner edge
substrates. Marks on the metapodium and tentacles, of the ridge. They are very constant within each
when magnified to around X50. were observed, in live species and are useful at spécifie level.
animais, to be three dimensional and to be comprised The morphology of the foot is problematical and
of many minute irregularly shaped marks, apparently of little use for spécifie assessment because of
floating within the semi-transparent membrane at difficulty in determining when a metapodium is fully
différent le\els and not on the surface. As a extended. It was noticed that there was a tendency for
metapodium is extended it becomes thinner and the the individuals in live sample groups to behave
marks which at tlrst may appear to be ofsolid colour, similarly when being photographed. For example, if
become stretched and often quite diffuse (Figs 33-34). one spécimen in a group was observed to hâve a
Iridescent marks are found on the foot of some rounded metapodium, then ail spécimens in that group
Gnvmlina (Fig. 20). Thèse were first observed in live tended to show the same state, but on other occasions
animais collected by dredging off the mountainous, some individuals of the same species might behave
north western coast ofVenezuela where the substrate differently. Sometimes mantles were not visible and
included a noticeable proportion of gravel containing on other occasions they were fully extended. It is
quartz and pyrites which sparkled when magnitled in believed that thèse incosistencies may reflect the
strong light. It seems possible that the présence of degree of traumatism suffered by the animais during
thèse iridescent markings is an adaptive feature. coUecting and sorting, the time delay before
However, similar iridescent marks were subsequently photographing took place or the water conditions in
found in another Gramdma species (Figs 28-29) in the the aquarium. There were incidents when one or more
extensive area of muddy substrates to south and west individuals in such a group would extend its
of Isla Margarita, Venezuela, where there was no metapodium to become very elongated and thin
évidence of any gravel particles. This contradicts the distally. This made it impossible to accurately assess
hypothesis and leaves the matteropen to further study. foot length uniess it was extended to be very long and
Melanism was noted to be very common in some narrow distally - definitely the fully extended state.
Gramdina species. For example, in Gramdina From thèse observations it follows that ail Caribbean
monjesensis from the harbour, Monjes del Sur, Gramdma may be able to extend the metapodium to a
Venezuela (Fig. 25), and G. ocella from East very elongate state, but more work on live animais is
Holandes Cays, San Blas, Panama (Fig. 32), where necessary in orderto résolve this matter.
approximately half the samples collected were With the large number of features to consider
melanistic. Many other species collected in the Gramdma species are often relatively easy to identify
Southern Caribbean showed no sign of melanism. In by shell morphology alone. They are certainly more
this respect, the genus Gramdina and the genus easily identified than shells of Gihbenda species in
Gihbenda are very similar. which there are fewer variables and in which it is
often essential to examine both shell and animal in
Identification of species. This has been based on order to achieve accurate spécifie assessment
shell morphology, and animal morphology and (McCleery, 2008, 2009). However, for example, in
chromatism when live animais were available. Several this article G. monjesensis, G. ocella, and G. plagula
radulae were extracted, and were noted to be Type 4 would not hâve been described as separate new
(Coovert and Coovert, 1995: 56), typical ofthe genus species without observation of animal chromatism.
Gixifudina. but the number examined was considered Morphological variations alone between shells of
too small to be useful for species assessment. Some thèse three new species would not hâve been
features ofshell morphology are dépendent on the âge considered suftlciently différent for positive
of the animal, being weak or absent in young adults séparation. Now that it has been clearly demonstrated
40
T. McCleery NOVAPEX 1 1(2-3): 37-71, lOjuin 2010
that they are distinct species, largely due to différences live animais inhabit the surface layer and loose algal
in animal chromatism, the true importance of very material on top ofthe solid mud.
small différences in their shell morphology can be Samples from live material were photographed in a
recognized. small aquarium below a microscope with a digital
caméra mounted on top. The same equipment was
Discussion. Différences in shell morphology between used for detailed imaging of dried shells and was
Gramilina species coUected in shallow rocky or reef calibrated so that shell dimensions could be obtained
areas and those dredged in muddy substrates are from data displayed by the software. Dimensions of
significant. Shells from the former habitat tend to be shells are accurate to plus or minus 2 %, and those of
small, and hâve more rounded ends (Figs 61-69), live animais to plus or minus 4%. Ail relevant data,
whereas, those dredged from deeper, muddy habitats including a chosen shell image were entered in a
are frequently larger, very heavily callused, and database. One spécial feature of the database is a
slightly produced at both ends (Figs 40-45). comparator which enables a simple and very effective
Shells of Granulina spp. are normally semi- means for comparing two or more shell images. This
transparent when fresh, but tend to graduaily become proved to be very useful in highlighting small
translucent white when dried. Dead collected shells morphological différences.
are frequently opaque, and occasionally extremely Shell images are presented at X25 magnification,
hyaline, particularly when collected in fine mud - giving a true impression of relative sizes. Animal
probably long-dead shells. images are presented at various magnifications in the
As was found to be the case in the genus range lOX to 20X and images ofplications (Figs 91-
Gibberula Swainson, 1840 (McCleery, 2008, 2009), 108) at a uniform image size of 5 cm which equates
Gramilina species also appear to form species groups with approximately X50 to XI40 depending on shell
with wide geographical ranges. For example, G. length. A number ofS.E.M. images ofsurface texture
producera n. sp. from Curaçao appears to be closely are presented at various magnifications and hâve
related to G. molinai Espinosa and Ortea, 2005, from intégral scale bar.
Cuba, and the authorhas in his collection several other
undescribed species belonging to this group, Abbreviations
represented by only one shell, from widely separated
locations. It is expected that many species within such MNHN: Muséum national d'Histoire naturelle, Paris.
species groups will be found to hâve small géographie
ranges and to be endémie to their type localities. No AWC: Andrew Wakefield Collection.
attempt has been made herein to appraise the varions TMC: Tony McCleery Collection.
groupings within the genus Granulina. ad.: adult spécimen.
juv.:juvénile spécimen.
MATERIAL and METHODS Iv.: live collected.
dd.: dead collected.
Hand dredging in sand or muddy substrates and the L.: shell length.
use of a hand operated suction pump on rocks and W.: shell width.
rubble substrates were the most productive methods of
collecting Granulina in shallow waters down to SYSTEMATICS
approximately 30 mètres. Night diving yielded some
positive results as spécimens could be picked up from Family CYSTISCIDAE Stimpson, 1865.
sand and rubble, or off rocks. Many species were Subfamily GRANULININAE Coovert and Coovert,
collected by dredging from the author's yacht with the 1995.
aid of a small hydraulically operated réel. The Genus GA-oMi/Z/nfl Jousseaume, 1888.
résultant grit from ail methods of collection was Type species: Marginella pygmaea Issel, 1869 (non
screened into four grades. The finer screenings were Marginella pygmaea G. B. Sowerby II, 1846), =
&
placed in bowls of sea water and covered. Live Marginella isseli G. H. Nevill, 1875 (nom. nov.).
animais then crawled up the sides where they could be
picked up. The finergrades ofgrit from deep dredging Granulina volcana n. sp.
were also sorted visually for dead shells, which Figs 1-2,5-6,37-39,91
comprised an average ofapproximately 95 percent of
ail shells collected by dredging. As dredging Type material. Off Islas Los Testigos, to north,
techniques improved so did the percentage of species Venezuela, 1 1°28'N, 63°06'W, 73 m.
containing live animais. Before collecting ceased, Holotype. 2.67 x 1.87 mm, W:L 70%, MNHN 21985;
approximately halfofail species collected by dredging paratype 1. 2.67 x 1.88 mm, W:L 70%. MNHN 21986;
contained live material, probably due to the dredges paratype 2. 2.70 x 1.88 mm, W:L 70%, AWC; paratype
skimming the surface of substrates rather than by 3. 2.73 X 1.94 mm, W:L 71%, AWC; paratype 4. 2.83 x
biting too deeply and becoming blocked - much dead 1.98 mm, W:L 70%, TMC; paratype 5. 2.44 x 1.61 mm,
material appears to lie in the solid, settled mud, but W:L 66%, TMC.
41
T. MtCli:ery Eighteen iicw spocios o\'(Jriiiiiiliiia
Othcr material. 2 ad. I\.. 2 jii\. I\., a luiinbcr of substantially off-while with some darker areas.
brokcn pièces, olT Islas Los Tcstigos. U) norlli. Approximately Iwelve short, yellowish, clongate
Vcne/iicla, 1 1 28'N, 63^()6'W, 73 m, tÎvIC". marks t)n darker patch can be detected emerging from
under anterior dorsal callus, fanning oui posteriorly.
Type locality. OtT Islas Los Testigos, to north. Almost horizontal, narrow, transverse whitish band
Vene/Liela. 1 I 2K'N. 63^'06'W (Map réf. 16). localcd posteriorly.
Description. Shell wilhoiit coloiir. obovate, tending to Distribution. Only known from the type locality.
mm
pm\mri,tbrWin:.LSi6/6e-7r1an%g.e B2.o4d4yxwh1.o6r1i trantsolu2c.e8n3t ,wxhi1t.c9.8 Remarks. Gniniilina volcanu n. sp. is closest to G.
weak striâtions close to external varix, covered by cal/a n. sp. with which it is compared. Thèse two
closely related species were collected from localities
almost smoolh, \ery light calkis wash. Lip strongly
about seven miles apart. The type locality of
and e\enly ciir\ed, curlcd invvards, evenly wide,
Grcmiilina volcana, 73 m, is subject to the very strong,
moderateiy strong. Foiirteen denticles almost tlll inner westward flowing Equatorial current which enters the
edge. widely spaced posteriorly. In side view, lip Caribbean between Trinidad and Tobago, and
evenly convex. External varix wide, widest and raised
Grenada, whereas, the type locality ofG. calla, 28 m,
on dorsum medially. narrowest anteriorly, sweeps
is close to Isia Grandi, Los Testigos islands, and
around posterior canal spreading slightly over dorsum,
considerably sheltered from strong currents.
highest above insertion point, forms lumpy ridge Gnvmlina volcana is consistently lightly callused, the
ventrally, merges vvith pariétal callus ridge.
varix weakens and sweeps around posterior canal, the
Weakening \arix sweeps around anterior canal,
posterior ridge is smooth, whereas G. calla is
merges with anterior callus, labial edge merges with
consistently very heavily callused, has very strong
raised tlrst columellar plication. Four plications fill callus at the apex, the varix retains a strong profile
approximately one third of aperture. First moderateiy
fully around the posterior canal, and the posterior
deep. narrow. Second strongest, strongly kinked,
ridge is relatively long and lumpy. Chromatism aIso
merges with anterior callus. Third strong, short,
distinguishes between thèse two species: G. volcana
rounded lump distally. Fourth weakest, smaller
has two strong red lines on the metapodium, a siphon
rounded lump distally. Weak pariétal ridge
with distinct orange-red edges and no white marks
commences at distal end of fourth plication, extends (Figs 2, 5), whereas, G. calla has random red spots on
posteriorly and merges with posterior, lumpy ridge.
the metapodium, the siphon with three white spots
Ali plications excavated, particularly second and third. basely, no orange-red colouring (Figs 4, 7). The
Aperture moderateiy and uniformly wide. Surface of
différences described in the shell morphology between
ail callus deposits textured with minute pustules. thèse two species were consistent in ail mature adult
shells examined.
Animal: Length of fully extended foot unknown,
width approximately same as shell, semi-transparent Etymology. The name was inspired by the impression
with several small off-white marks laterally, ofan erupting volcano given by the chromatism of
increasing in size posteriorly. Largest mark metapodium and its posteriorpapillae. Latin for
substantial, white, level with posterior end of shell. volcano is volcanus.
Small off-white marks on metapodium concentrated
medially forni distinct medial Une, adjacent bright red Granulina calla n. sp.
lines formed by contiguous red spots anteriorly, Figs 3-4, 7, 40-42, 92
reducing in density and fading out distally, further
Type material. Isia Grande, Islas Los Testigos,
small, scattered, off-white spots extend to latéral
emdagrekss. Pwrhoipcohdiufmadeoffo-wuhtited,istfaollrym.edSebmyi-mtarannyspasrmaelnlt HVoelnoetzyupeel.a,2.I5l7°2x2.18.'7N5,m6m3,°0a8d..rIWv.,, 2W8:Lm.68%, MNHN
tmeanrtkasc.lesB,lacloknge,yesthilnocawtietdhonthbraeseaiisrrweeglullianrglsy. Ssippahcoend 2M1N95H4N; p2a1r9a5t5y;pepa1r.at2y.p6e92.x 2.16.940xmm1.,82Wm:mL, 7W0:%L,
moderateiy long, thin, semi-transparent with many 69%, AWC; paratype 3. 2.62 x 1.80 mm, W:L 69%,
AWC; paratype 4. 2.65 x 1.84 mm, W:L 70%, TMC;
small off-white spots, distinctive orange-red edges
basally. Mantle not observed fully extended. Small paratype 5. 2.53 x 1.74 mm, W:L 69%, TMC.
spuwsetlullionsges. vSimsailbllearleaatseraolflyc,oltohuerrsefcoarne,aibseolibeeveddetetcotebd:e Other material. 2 ad. Iv., I juv. Iv., 4 ad. dd., Isia
Grande, Islas Los Testigos, Venezuela, 11°22.8'N,
off-white, yellowish brown, reddish brown, turquoise, 63°08.rW, TMC.
and black. Five simple papillae grouped posterior
medially on mantle. Mantle roof chromatism appears Type locality. Isia Grande, Islas Los Testigos,
diffuse through translucent white dorsum, Venezuela, I l°22.8'N, 63°08.rW (Map réf. 15).
42
T. McCleery NOVAPEX 1 1(2-3): 37-71, 10juin 2010
Description. Shell without colour, obovate, posterior off horizontal. A few minute irregular black marks
mm
slightly produced. Size range 2.53 x 1.74 to 2.69 which appear to be attached internally to mantle roof
X 1.90 mm, W:L 68-70%. Body whorl semi- can be detected.
transparent, some very weak irregularly spaced
striations, covered by very light callus wash, finely Distribution. Only known from the type locality.
textured with minute pustules. Lip evenly curved,
curled inwards strongly, wide, widest medially. Remarks. Gramdina calla n. sp. is closest to G.
Fourteen somewhat irregular denticles fil! inner edge, volcana n. sp. with which it is compared. Thèse two
closely spaced anteriorly, wider medially, more so closely related species were collected from localities
posteriorly. In side view, lip slightly convex. Very about seven miles apart. The type locality of G.
strong extemal varix, slightly raised on dorsum volcana, 73 m, is subject to the very strong, westward
medially, sweeps very strongly around posteriorcanal, flowing Equatorial current which enters the Caribbean
stops abruptly ventrally, forms very strong, lumpy, between Trinidad and Tobago, and Grenada, whereas,
callus ridge, merges with weak pariétal callus ridge. the type locality of G. calla, 28 m, is close to Isla
Varix, remaining wide, sweeps around siphonal canal, Grandi, Los Testigos islands, and is considerably
merges with small area of anterior callus and raised sheltered from strong currents. G. volcana is
first columellar plication. Four strong plications tlll consistently lightly callused and has a smooth
approximately one third of aperture. First moderately posterior callus ridge, whereas G. calla is consistently
deep, narrow. Second strongest, strongly kinked as it very heavily callused with very strong callus at apex,
émerges, merges with moderately heavy anterior the extemal varix retains a strong profile fully around
callus. Third strong, short, pointed lump distally. the posterior canal, and has a relatively long and
Fourth weakest, smaller pointed lump distally. Weak, lumpy posterior ridge. Chromatism also distinguishes
even, pariétal ridge extends posteriorly from second between thèse two species: G. volcana has two strong
plication, merges with posterior ridge. Ail plications red medial Unes on the metapodium, a siphon with
excavated, second and third strongly, fourth almost distinct orange-red edges and no white marks (Figs 2,
completely. Surface of ail callus deposits, including 5), whereas, G. calla has random red spots on the
lip and space between plications, textured with minute metapodium, the siphon with three white spots basely
pustules approximately three times larger than those and no orange-red colouring (Figs 4, 7). Différences
covering dorsum. Aperture, evenly and moderately described in shell morphology between thèse two
wide. species was consistent in ail mature adult shells
Animal: Length of fully extended foot unknown, examined.
width approximately same as shell, semi-transparent
with several off-white marks latérally. Random Etymology. The name refers to the heavy callus and
smaller white marks on metapodium form indistinct strong knobbly deposits. The Latin for callus is
medial Une extending distally. Adjacent to Une, on calliim.
each side, approximately tlfteen small, deep red spots
intermingled with off-white ones. Further small, off- GraniiUna colonensis n. sp.
white spots extend from red spots to latéral edges. Figs 8-9, 43-45, 93, 109-112
Propodium largely off-white with some minute red,
brown and white marks. Semi-transparent tentacles, Type material. OffColon, to east. Panama, 9°31.5'N,
long, thin with five or six irregularly spaced, off-white 79°52.0'W, 57 m.
marks. Black eyes located on basai swellings. Siphon Holotype. 1.98 x 1.31 mm, W:L 66%, MNHN 21958;
moderately long, thin for genus. Three or four diffuse paratype 1. 1.96 x 1.35 mm, W:L 69%, MNHN
white spots at base, fine off-white spots intermingled 21959; paratype 2. 1.95 x 1.27 mm, W:L 65%, AWC;
with fewer reddish brown spots, becoming numerous paratype 3. 2.00 x 1.35 mm, W:L 67%, AWC;
distally. Mantle not observed extended. Small paratype 4. 1.76 x 1.20 mm, W:L 68%, TMC;
swellings visible laterally, therefore, believed to be paratype 5. 1.96 x 1.33 mm, W:L 68%, TMC.
pustulose. Small areas ofcolours also detected - off-
white, yellowish brown, reddish brown, turquoise, and Type locality. OffColon, to east. Panama, 9°31.5'N.
black. Five simple papillae grouped posterior medially 79°52.0'W, 57 m (Map réf. 1).
on mantle (Fig. 7). Thèse were observed to extend and
retract rapidly, independently of mantle and each Description. Shell without colour, obovate, slightly
mm
other. Mantle roof chromatism visible through biconic. Size range 1.76 x 1.20 to 2.00 x 1.35
translucent dorsum, substantially off-white with mm, W:L 65-69%. Body whorl semi-transparent, light
numerous darker areas bearing small yellowish marks callus wash, slightly textured. Lip evenly curved,
and occasional duU reddish spots. Approximately curled inwards slightly, moderately narrow, widest
eight short, yellow, elongate marks on darker patch medially, completely filled by twenty two denticles. Ui
émerge from under anterior dorsal callus and fan out side view, lip convex. Extemal varix very strong,
posteriorly. Towards posterior, narrow transverse wide, slightly wider posterior medially, not raised on
whitish band slopes downwards to right at about 10° dorsum, retains profile and sweeps around posterior
43
T. McCl LHRY Eighteen new spccies oi'Graiiiiliiui
canal uith little apparent shoulder, siops abmpiK ai Short, pale, whitish, transverse band located posterior
apertiire, very strong, widc, lumpy, callus ridgc medially, sloping slightly downwards to right.
continues anteriorly. merges with pariétal callus ridge. Distribution. Only known from the type locality.
Varix, remaining wide. sweeps around anterior canal,
weakening, merges inlo anterior callus. labial edge Remarks. Ciramiliiui colonemis n. sp. appears to be
merges with slightly raised first cokimellar plication. most closely related to G. darienensis n. sp. (Figs 46-
Four plications fill approximately 30% of aperture. 48) with which it is compared. The shell of G.
First sinuous, slightly raised. Second, third and fourth colonensis is heavier and more biconic in shape, the
plication discontinuons due to very deep pariétal lip wider, the varix stronger, posterior and siphonal
excavation. Distal portion of second stepped canals are less flared (Fig. 110) than G. darienensis
dovvnwards at distinct medial lump, extends onto (Fig. 118). The tlrst plication not thickened medially
anterior callus with wide axial dimension, widely as in G. darienensis (Fig. 94). Body whorl surface is
rounded distally. Distal portions of third and fourth te.xtured with small round granules of various sizes
combine in substantial vertical ridge with tlne distal (Figs 110-112), but covered by an even scaly texture
end turned outwards onto anteriorcallus. Posteriorend in G. darienensis (Figs 118-120). Chromatism has
weakens and merges with weak pariétal ridge. Inner much in common between the two species, but the
portion of fourth plication, located on pariétal wall strong off-white medial line on the metapodium ofG.
deep inside aperture. Aperture moderately and eolonensis (Figs 8-9), is absent in G. darienensis (Figs
uniformly wide. Surfaces generally lightly textured: 10-11). Pale annular rings highlight the eyes of G.
callused surfaces with scales and ridges, adjoining darienensis but are absent in G. colonensis. Thèse two
body whorl with random sized, roundish lumps (Figs new species are separated geographically by
110-112). approximately seventy miles.
Animal: Length of fully extended foot more than
twice length of shell, width approximately same as Etymology. Colon city is located at the Caribbean end
shell, semi-transparent. Small off-white marks of the Panama canal. Graniilina colonensis was
intermingled with many smaller reddish brown spots collected a few miles to the east and takes its name
laterally. Metapodium finely elongate distally. Solid, from the city ofColon.
wide, off-white, medial line extends posteriorly,
approximately one third length of metapodium, GranuUna darienensis n. sp.
Figs 10-11,46-48,94, 117-120
widening and becoming less solid medially,
weakening and fading distally. Weak marks between Type material. OffIsla Chichime, San Blas, Panama,
medial line and edge of metapodium diffuse, off-
9°37.3'N, 78°53.2'W, 75 m.
white, comprised of many small spots. Marks reduce Holotype. 2.14 x 1.39 mm, W:L 65%, MNHN 21960;
in size posteriorly, intermingle with smaller off-white paratype 1. 1.99 x 1.34 mm, W:L 67%, MNHN
and reddish brown spots. Edges of metapodium, 21961; paratype 2. 1.88 x 1.30 mm, W:L 69%, AWC
except distally, highlighted by small, solid, off-white, paratype 3. 2.17 x 1.37 mm, W:L 63%, AWC
dashes, generally separated by reddish brown spots. paratype 4. 2.02 x 1.31 mm, W:L 65%, TMC
Off-white spots densely concentrated on propodium, paratype 5. 2.06 x 1.36 mm, W:L 66%, TMC.
particularly close to head. Semi-transparent tentacles,
long, thin, rust coloured over whole length with Other material. 16 ad. dd., off Isla Chichime, San
diffuse off-white marks, strongest distally. Eyes black. Blas, Panama, 9°37.3'N, 78°52.4'W, 75 m; 7 ad. dd.,
Siphon and mantle not observed fully extended. off Isla Limon, Panama, 9°31.5'N, 79°52.0'W, 64 m,
Siphon with concentrated small off-white spots. TMC.
Mantle: poorly observed. Cap-like posterior,
suggestive of papillae bears one strong pustule Type locality. Off Isla Chichime, San Blas, Panama,
medially (Fig. 9). Mantle roof chromatism 9°37.3'N, 78°52.4'W (Map réf. 2).
substantially off-white with some pale orange spots.
Figures 1-11
1-2, 5-6. GranuUna volcana n. sp. OffIslas Los Testigos, to north, Venezuela, 1 1°28'N, 63°06'W, 73 m.
1. Holotype. 2.67 x 1.87 mm, W:L 70%, MNHN 21985; 2. Young adult spm. 2.71 x 1.86 mm, W:L 69%, TMC; 5.
Young adult spm. 2.49 x 1.76 mm, W:L 71%, TMC; 6. Young adult spm. 2.68 x 1.86 mm, W:L 69%, TMC.
3-4, 7. GranuUna callan. sp. Isla Grande, Islas Los Testigos, Venezuela, 1 1°22.8'N, 63°08.rW, 28 m.
3. Holotype. 2.57 x 1.75 mm, W:L 68%, MNHN 21954; 4, 7. Paratype 5. 2.53 x 1.74 mm, W:L 69%, TMC.
8-9. GranuUna Colonensis n. sp. Holotype. OffColon, to east, Panama, 9°31.5'N, 79°52.0'W, 57 m. 1.98 x 1.31
mm, W:L 66%, MNHN 21958.
10-11. GranuUnadarienensis n. sp. OffIsla Chichime, San Blas, Panama, 9°37.3'N, 78°53.2'W, 75 m;
10. Holotype. 2.14 x 1.39 mm, W:L 65%, MNHN 21960; 11. Adult spm. 2.19 x 1.41, W:L 64%, TMC.
44
A
T. McCleery NOVAPEX 1 1(2-3): 37-71, lOjuin 2010
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45
T. McCLLliRY Eightecn ncw species o\'Gidiiiiliiia
Description. Shell w ilhout colour, obo\ate. Size yellowish-while and rust spots, larger marks
range I.KS x 1.30 mm to 2.17 ,\ 1.37 mm. W:L 63- intermingled with Une black spots. Mantle roof
69"o. Body whorl iranslueent white wiili light callus chromatism substantially yellowish-white with pale
wash, lighlly lextured. Lip evenly curved, ciirls orange spots and some darker areas. Approximately
inv\ards slightly, narrow, twenty threc irregular, vveak six indistinct, short, whitish, elongate marks on darker
denticles almost fill inner edge, fading out slightly patch émerge from under anterior dorsal callus,
above anterior canal, in side vievv, lip conve.\. fanning out poSteriorly. Pale whitish, transverse band
Extemal varix strong, moderalely wide. widest located posteriorly, slopcs slightly downwards to
medially, weakens. sweeps round slightly llared right.
posterior canal, fades out o\er immersed spire. Callus
remains wide, de\clops lumpy extemal edge ventrally, Distribution. Known from the type locality and offIsia
merges with very weak pariétal callus ridge. Posterior Linton, Panama.
callus extends slightly over dorsum. Weakening varix
sweeps around slightly tlared siphonal canal, merges Remarks. Graniilina clarienensis n. sp. is closest to G
with anterior callus. Labial edge of weakening varix, cokmemis n. sp. with which it is compared. G.
merges with slightly raised first columellar plication. ckirienensis has a somewhat lighter, more rounded
Four plications fill approximately 30% of aperture. shell, narrower lip, weaker varix around siphonal
First strong with significant lump medially. Second, canal, and posterior and siphonal canals considerably
strong with large, pointed lump as it émerges from more fiared than in G. colonemis. The first plication
deep excavation, thins distally, fades out on anterior bears a medial lump which is absent in G. colonensis.
callus. Third and fourth plications discontinuons due The body whorl surface is covered by an even scaly
to a very deep pariétal excavation. Emergent portion texture (Figs 1 18-120), whereas, in G. colonemis the
of third, strong and curved, anterior part continues body whorl is textured with small round granules of
outwards, stops before anterior callus, medial portion varions sizes (Figs 110-112). Chromatism has much in
continues posteriorly, merges with very weak pariétal common between the two species, but the strong off-
ridge. Fourth located deep inside aperture. Aperture white medial line on the metapodium ofG. colonemis
moderately and uniformly wide. Surfaces lightly is absent in G. darienensis. The eyes ofG. darienensis
textured with random sized scales (Figs 1 18-120). are highlighted by pale annular rings, but absent in G.
colonensis. Thèse two new species are separated
Animal: Length of fully extended foot approximately geographically by approximately seventy miles.
twice length of shell, width approximately same as
shell, semi-transparent. Latéral marks not observed. Etymology. This species is named after the mountains
Metapodium pointed distally, variety of shapes and of Darien which form the backdrop to the archipelago
sizes of marks give variegated effect in shades from ofSan Blas.
white to yellowish-white, intermingled with rust
brown spots. Concentration of marks shows as faint Graniilinagayracaensis n. sp.
medial line. Propodium with small off-white spots Figs 12-15,49-51,95
intermingled with some smaller rust spots. Semi-
transparent tentacles, long, thin, with moderately Type material. Gayraca Bay, Santa Martha, Colombia,
heavy off-white marks, intermingled with some rust 11°19.5'N, 74°06.3'W, 7 m.
marks, distinct rust coloured areas basally, black eyes Holotype. 1.75 x 1.22 mm, W:L 70%, MNHN 21962;
with off-white annular rings on slight swellings. paratype 1. 1.61 x 1.08 mm, W:L 67%, MNHN
Siphon moderately long, thick, semi-transparent with 21963; paratype 2. 1.92 x 1.29 mm, W:L 67 %, AWC;
many small off-white spots and two small rust spots. paratype 3. 1.64 x 1.17 mm, W:L 71%, AWC;
Mantle: papillae not observed, but may be présent, paratype 4. 1.82 x 1.26 mm, W:L 69%, TMC;
otherwise weakiy pustulose. Chromatism with fine paratype 5. 1.76 x 1.23 mm, W:L 70%, TMC.
Figures 12-23
12-15. Granulinagayracaensis n. sp. Gayraca Bay, Santa Martha, Colombia, 1 1°19.5'N, 74°06.3'W, 7 m;
12-13. Paratype 1. L61 x 1.08 mm, W:L 67%, MNHN 21963; 14. Holotype. 1.75 x 1.22 mm, W:L 70%, MNHN
21962; 15. Paratype 5. 1.76 x 1.23 mm, W:L 70%. TMC.
16-17. Granulina cartagenaensis n. sp. OffCartagena, Colombia, 10°22.4'N, 75°35.8'W, 25-41 m. Paratype 5.
2.20 X 1.63 mm, W:L 74%, TMC.
18-19. Granulina waltergomezi n. sp. OffIsia Chichime, San Blas Archipelago, Panama, 9°37.3'N, 78°53.2'W,
75-95 m. Holotype. 1.47 x 1.16 mm, W:L 79%, MNHN 21989.
20-23. Granulina iridisa n. sp. Cabo Codera, Venezuela, 10°35.2'N, 66°03.9'W, 18 m; 20, 22-23. Holotype. 2.37
X 1.66 mm, W:L 70%, MNHN 21967; 21. Adult spm. 2.46 x 1.67 mm. W:L 68%.
46