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Status Report On The Terrestrial Mollusca of Jamaica - Rosenberg & Muratov (2006)
Status Report On The Terrestrial Mollusca of Jamaica - Rosenberg & Muratov (2006)
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ABSTRACT.—A total of 562 species of snails and slugs are known from Jamaica, of which 505 (90%) are endemic. Both
sympatric and allopatric species diversity are high and the terrestrial mollusk fauna is as diverse as any in the world for the
size of the island. Although less than 15% of the original forest cover remains and deforestation continues, many snail
species survive in the secondary forest that covers more than 25% of the land area. Perhaps as much as 90% of the mollusk
fauna still exists, although some species have suffered great contractions of range. Tree snails of the genus Anoma
(Urocoptidae) are the most critically endangered and some are probably already extinct. Many Jamaican species have
naturally small ranges, so even with intensive sampling, establishing that a species is no longer extant is difficult, especially
if the type locality is no more precise than “Jamaica”. A few endemic species, particularly those adapted to dry conditions,
are expanding their ranges.
Eight taxa formerly considered synonyms or subspecies are confirmed as full species: Priotrochatella pulchra
(Helicinidae), Geomelania gracilis and G. magna (Truncatellidae), Leidyula trichroma (Veronicellidae), Aerotrochus
mcnabianus (Sagdidae) and Pleurodonte amabilis, P. catadupae and P. candescens (Pleurodontidae). Fourteen taxa are
synonymized: Choanopoma dislocata tentatively with Parachondria shepardiana (Annulariidae); Geomelania affinis with
G. jamaicensis, G. exilis with G. beardsleana, G. jarvisi with G. minor, and G. striosa pumila with G. striosa (Truncatellidae);
Varicella leucozonias striatella with V. leucozonias (Oleacinidae); Urocoptis lata with U. producta and Pupa lata tentatively
with U. sanguinea (Urocoptidae); Helix albicans with Stauroglypta anthoniana (Sagdidae); Helix lindsleyana tentatively
with Pleurodonte atavus, Helix soror var. latior with Pleurodonte peracutissima; Pleurodonte tridentina chittyana and P.
t. schroeteriana with P. tridentina (Pleurodontidae); and Dialeuca jacobensis with D. conspersula platystyla. Pleurodonte
carmelita (Férussac, 1821) is preoccupied by Helix carmelita Lichtenstein, 1794 and is replaced by Pleurodonte mora
(Griffith & Pidgeon, 1834). Bulimus macrospira is transferred to Sigmataxis (Oleacinidae), Helix brevis to Happiella
(Systrophiidae) and Helix alveus to Stauroglypta (Sagdidae). Gastrocopta polyptyx (Vertiginidae), Drymaeus rufescens
(Bulimulidae) and Lacteoluna selenina and Aerotrochus turbiniformis (Sagdidae) are rejected from the Jamaican fauna.
Ten introduced species are newly reported from Jamaica: Helicodiscus parallelus (Helicodiscidae), Paralaoma servilis
(Punctidae); Karolus consobrinus (Ferussaciidae), Beckianum beckianum (Subulinidae), Bulimulus diaphanus and Drymaeus
multilineatus (Bulimulidae), Zachrysia provisoria (Pleurodontidae), Huttonella bicolor (Streptaxidae), Happiella decolorata
(Systrophiidae), and Bradybaena similaris (Bradybaenidae).
reported 385 species, but omitted the species named the central Cockpit Country, and the higher altitudes
by Chitty (1857a, b), stating (p. 361)“they do not ap- of the Blue Mountains and the John Crow Mountains
pear to have been adopted [by other workers]”. If he in the east had been poorly sampled, whereas easily
had included the 83 names that Chitty introduced at accessible areas such as Montego Bay, Mandeville,
species rank in 1857, it would have raised his total for Bogwalk and Fern Gully had been sampled repeatedly.
Jamaica to 468. Subsequent estimates of diversity have Micromollusks (under 5 mm) had not been sampled in
ranged from 400 to more than 500 species: Vendryes many areas. Even now most peaks in the Blue Moun-
(1899), 533; Simpson (1895), 464; Russell-Hunter tains have never been sampled. Our field work, coupled
(1955), 450 ± 100; Goodfriend (1986a), 400-450; Kay with examination of museum collections and review
(1995:60), “over 500”. We demonstrate herein that the of the literature, forms the basis of this report on the
highest previous estimates of terrestrial molluscan di- status of the Jamaican terrestrial mollusks.
versity in Jamaica are too low. More than 560 described
species of snails and slugs are currently known and METHODS
ongoing exploration continues to reveal new species.
The total number of species is likely to exceed 600, Observations on the Jamaican fauna were made
with about 90% endemism. during 12 collecting trips from 1986 to 2002, during
A number of museums have strong collections of which more than 1000 sites were sampled. Intensive
Jamaican land mollusks. The Museum of Compara- collecting, including soil and leaf litter sampling, was
tive Zoology holds most of C. B. Adams’ primary type done at 607 sites between 1999 and 2002; station num-
material (Clench & Turner 1950a, Jacobson & Boss bers for these sites have the prefix “JBS”, for Jamai-
1973), which was collected and named between 1844 can Biotic Survey <http://data.acnatsci.org/jamaica>.
and 1851. Adams rarely gave a locality more specific In addition to mollusks, other invertebrates, particu-
than “Jamaica” when describing his new species. He larly millipedes, were collected as part of the survey.
traded and sold specimens to a number of scientists The molluscan material is housed at the Academy of
and collectors, so his type material can also be found Natural Sciences, Philadelphia (ANSP); not all of it
in various other institutions. After Adams died, Ed- has yet been sorted. Sorting includes sifting soil and
ward Chitty–who had supplied some of Adams’ mate- leaf litter through nested sieves with mesh openings of
rial–described a number of species between 1853 and 4 mm, 2 mm and 0.5 mm ((Hubbard Scientific Co.,
1857; most of his type material is preserved in the Northbrook, Illinois; #5, #10 and #35), with the 2 and
Natural History Museum, London (Clench & Turner 0.5 mm fractions being examined under a microscope.
1950b). The Academy of Natural Sciences in Phila- Synoptic material has been and will continue to be
delphia (ANSP) holds specimens of more than 80% of deposited at the Institute of Jamaica, Kingston. Most
the species known from Jamaica, including material of the species are illustrated in an online interactive
collected by Amos Peaslee Brown in 1910 (Pilsbry & key (Rosenberg & Drumm 2004).
Brown 1910), Horace Burrington Baker in 1933 (Baker As material is identified and catalogued, data are
1934a-b, 1935a-d), Glenn Goodfriend (1983) and our- being made available through the website for ANSP’s
selves from 1986 to 2002. Baker was the first worker malacology collection <http://erato.acnatsci.org/
to routinely record the altitude, latitude and longitude biodiversity_databases/snails.php>. Other collections
of his sites and to preserve Jamaican mollusks in etha- examined include the American Museum of Natural
nol. In 1927 and 1928, Charles R. Orcutt collected Ja- History, New York (AMNH); Museum of Compara-
maican mollusks for the United States National Mu- tive Zoology, Cambridge (MCZ); the United States
seum (Coan 1966). His material is mainly dead-col- National Museum, Washington, D.C. (USNM); the
lected shells, often from along stream banks and often Florida Museum of Natural History, Gainesville
with subfossil material mixed in. Goodfriend’s Jamai- (FLMNH); the Natural History Museum, London
can collections, mostly made between 1976 and 1979, (BMNH); and the private collection of the late Glenn
were the first with locality data precise enough to al- Goodfriend.
low particular sites to be relocated. Goodfriend’s alco- All known literature on Jamaican terrestrial mol-
hol-preserved material and his dry specimens of lusks was surveyed to generate a species-level classi-
Pleurodonte are held at ANSP, along with a copy of fication and synonymy that summarizes the current
his field notes. status of each nominal species. Many Jamaican taxa
Until our intensive surveys from 1999 to 2002, have not been treated since the 1930s or earlier, so in
many remote areas such as Dolphin Head in the west, some cases the most recent status is uninformative,
TERRESTRIAL MOLLUCA OF JAMAICA 119
1) Synonym or questionable synonym. Status ac- Our surveys revealed several introduced species
cepted, unless morphology gave strong reason for dis- not previously reported from Jamaica (Rosenberg &
agreement, in which case evaluated for subspecific or Muratov 2001), all in cultivated or otherwise disturbed
specific status. areas: Helicodiscus parallelus (Helicodiscidae) in the
2) Infrasubspecific rank. Synonymized if clearly Blue Mountains north of Kingston; Karolus
based on size or color alone; treated as a questionable consobrinus (Ferussaciidae), widespread; Beckianum
synonym if based on shape or sculpture, unless mor- beckianum (Subulinidae), scattered occurrences;
phology gave strong reason for disagreement, in which Bulimulus diaphanus (Bulimulidae), widespread, es-
case evaluated for subspecific or specific status. pecially in areas disturbed by cattle; Drymaeus
3) Subspecies. Synonymized if sympatric with and multilineatus (Bulimulidae), in southeast
intergrading to the nominotypical subspecies; ranked Westmoreland; Zachrysia provisoria (Pleurodontidae),
as a full species if no intergrades observed; retained as in the Rio Cobre Gorge, St. Catherine; Huttonella bi-
a subspecies if allopatric with intergrades to the nomi- color (Streptaxidae), scattered occurrences; and
nate subspecies, or if information was insufficient to Happiella decolorata (Systrophiidae), scattered occur-
reevaluate the status. Because of the complexity of the rences, usually in association with bamboo. The last
taxonomy, subspecies in Neocyclotidae and Anoma of these is the same species reported from Barbados as
(Urocoptidae) were automatically listed as synonyms Happiella cf. decolorata by Chase and Robinson
of the nominotypical form. (2001); its discovery in Jamaica confirms it as a synan-
4) Species. Status as full species was accepted, thropic species and leads us to remove the “cf.” from
except for names overlooked by subsequent revisors, its designation. Bradybaena similaris (Bradybaenidae)
in which case evaluated based on examination of type has been intercepted with produce from Jamaica by
material if available or by interpretation of the origi- the United States Department of Agriculture (David
nal description. Robinson, pers. comm.), but the source population
within Jamaica has not been located. Paralaoma
RESULTS servilis (Punctidae) has been collected in St. Thomas
Parish (Natalie Blake pers. comm.).
At present, the Jamaican terrestrial fauna contains Geostilbia aperta (Swainson, 1840)
562 valid molluscan species, to which 928 available (Ferussaciidae) was first reported from Jamaica by
names have been applied, as listed in the Appendix. Bland (1861, p. 356, as Achatina gundlachi, a syn-
These species are distributed among 123 genera and onym). Pilsbry’s report (1908) is based on the same
subgenera and 34 families (Table 1). Of the valid spe- specimens, ANSP 3753, from Bland via the A. D.
cies, 505 are endemic to Jamaica (Table 1), 10 are na- Brown collection. There have been no subsequent re-
tive but occur elsewhere, 25 are introduced by humans, ports of the species in Jamaica, but its presence can
and 22 are of uncertain status (Table 2), i.e., cryptoge- now be confirmed based on specimens from Lowden
nic. We consider several species to be native because Hill, St. Thomas (Natalie Blake, pers. comm.) and from
of their occurrence as Pleistocene fossils in Jamaica. our station JBS 2 in the Caymanas area of St. Andrew
Helicodiscus apex and Lamellaxis monodon were re- Parish.
ported as fossils by Goodfriend & Mitterer (1988), Gastrocopta polyptyx (Vertiginidae), Drymaeus
Orthalicus undatus jamaicensis by Goodfriend (1986a) rufescens (Bulimulidae) and Lacteoluna selenina and
and Vertigo gouldii, V. milium, Ptychopatula macneili Aerotrochus turbiniformis (Sagdidae) are rejected from
and Punctum vitreum by Paul & Donovan (2006). The the Jamaican fauna. The species know as A.
endemic species are distributed among 19 families and turbiniformis in Jamaica is referred to A.
86 genera and subgenera. Thirteen families (6 opercu- subpyramidalis. Reasons for these changes are given
late and 7 pulmonate) have three or more endemic spe- in the systematics section following the discussion.
cies and form the core of the fauna: Helicinidae,
Stoastomatidae, Proserpinidae, Neocyclotidae,
120 G. ROSENBERG AND I. V. MURATOV
Figure 1. Species area curve for Jamaica compared to Pacific islands with elevation greater than 300
m, showing that Jamaica is as species rich for its area as Lord Howe Island, Rapa, and Oahu. Pacific
data from Cowie (1995, 1996).
Figure 2. Map of Jamaica showing parishes and informal biogeographic regions. Regions are
Western, Central, North Coast, South Coast, Blue Mountains and John Crow Mountains.
Boundaries are approximate.
stone soils of the Blue Mountains to the west. Charac- ous caves and caverns. There is often rock rubble with
teristic species include Proserpina bidentata, Annularia good drainage and a neutral to alkaline pH extending
pisum, Geomelania striosa, Meiophysema 10-100 cm or more in a layer above solid limestone.
lamelliferum, and Pleurodonte ingens. The depressions contain mainly eroded bauxite soil and
b) The Blue Mountains include the highest and can have impeded drainage and acid pH (Asprey &
most rugged terrain in Jamaica. They have been up- Robbins 1953). Depressions are frequently flooded and
lifted as much as 1000 m since middle Pliocene lack a crevice system, whereas the hill tops are prone
(Steineck 1974), which presumably has increased oro- to desiccation by radiation. Also, the rock-crevice sys-
graphic rainfall. The limestone cap has eroded off, ex- tem has lower and less variable temperature and higher
posing shale, igneous and metamorphic rocks. Char- and less variable humidity than the forest floor and so
acteristic species include Alcadia affinis, Hyalosagda protects invertebrates from desiccation (Diesel & Horst
similis, and Pleurodonte mora. Other species are shared 1995). The karst areas thus have ecotonal gradients of
with the John Crow Mountains, such as Pleurodonte pH and moisture, which presumably increase site di-
chemnitziana, Varicella leucozonias and Pleurodonte versity by increasing the number of microhabitats.
valida. At high elevations in the Blue Mountains, dif- Characteristic species include Annularia fimbriatula,
ferent faunal elements appear, including an undescribed Geomelania minor, Pleurodonte bainbridgii,
radiation of Punctidae. Blue Mountain Peak, the high- Eutrochatella pulchella, Sagda grandis, and Apoma
est point in Jamaica at 2252 m, had not been sampled chemnitziana.
for mollusks prior to our survey in March, 2002: 9 of d) Western, a largely limestone area, with the
15 species collected are apparently undescribed. Oc- Dolphin Head region and the Negril Hills possibly once
casional winter frosts are recorded on Blue Mountain isolated as separate islands. Characteristic species in-
Peak (Raw 1985), so the appearance of a distinctive clude Priotrochatella pulchra, Geomelania
fauna at high altitude is not surprising. jamaicensis, Thelidomus cognata, Pleurodonte
c) Central Karst, a limestone area with rainforest, picturata and P. tridentina. The western fauna grades
which has the highest diversity of land snails in Ja- into the central karst fauna, with which it shares spe-
maica. Rainforest on limestone also correlates with the cies like Colobostylus humphreysianus and
highest diversity in Queensland, Australia (Stanisic Eurycratera jamaicensis.
1997). Karst is best developed in the Cockpit Country, e) North Coast, a dry limestone area with coastal
which has extremely rugged topography with numer- terraces. Characteristic species include Eutrochatella
TERRESTRIAL MOLLUCA OF JAMAICA 123
[on] softer geological formations...the shales and clays of and continued efforts must be dedicated to conserving
the Black Grounds of Trelawny, the uplands of Clarendon, the biota.
the Bull Head Range, we find that of the rich forests, standing
even within living memory, nothing remains. Over wide Introduced species
areas even the crops have gone--bananas, ginger, yams--
Before European colonization, Jamaica was home
leaving bare rolling uplands, with a little coarse grass and
low fern--no home for our woodland birds and other animals.
to only two species of native land mammals (exclud-
This is also true to a certain extent in the Blue Mountains, ing bats), the hutia, Geocapromys brownii, which is
although there are still large areas of forest on the higher endangered, and a monkey, Xenothrix mcgregori which
portions of the range. The lower slopes, however, have lost is extinct. Neither is a likely predator on snails;
their forest cover and the wild life of this region has suffered Geocapromys is herbivorous (Anderson et al. 1983)
severe set-backs. and Xenothrix is thought to have been frugivorous
(MacPhee & Horovitz 2004). Many mammal species,
Another discouraging quotation comes from Proc- domestic and wild, have since been introduced to Ja-
tor (1953:80): maica. Cattle have a directly deleterious effect on
snails, trampling and fouling the habitat, and only a
The cockpit country is itself fast losing its densely-forested few native snails can coexist with them. Goats and fe-
character because of extensive cutting. Although there are ral pigs have less effect than cattle, but contribute to
few roads (and these bad), the woodcutters have access to degradation of forest. Introduced rats and mongoose
most of the remote fastnesses by means of a network of trails,
presumably eat native snails: small middens of pre-
over which they carry heavy hand-sawn boards balanced on
their heads, for miles. With the larger trees being cut,
dated snail shells can often be found in sheltered spots
the...flora is suffering modifications due to changes in among rocks. Similar impacts have been observed in
amount of light and moisture. other island faunas, for example New Zealand (Meads
et al. 1984). To our knowledge predation on snails by
Although these quotations date from half a cen- introduced species has not been directly observed in
tury ago, the situation today is not quite as dire as one Jamaica, so the effect on the fauna is difficult to gauge.
might expect. Except for the north coast, each biogeo- A number of snail species have been introduced,
graphic region has at least one remaining area of pri- but they do not seem to invade native forest, and so
mary forest. These include rainforests along the main are unlikely to out-compete native snails in undisturbed
ridge of the John Crow Mountains, the more remote areas. As yet, no carnivorous snails have been intro-
areas of the Blue Mountains, the heart of the Cockpit duced to Jamaica. Since the fauna has evolved with an
Country, the Dolphin Head area in the west, and dry array of native molluscivorous species, it may not prove
coastal forest in the Hellshire Hills in the south. The as vulnerable to such introduction as the snail faunas
first two of these areas are part of the Blue and John of Pacific Islands have been. The snail parasite
Crow Mountains National Park (79,666 ha) and the Angiostrongylus cantonensis was first reported in Ja-
others have been proposed as national parks or con- maica in 2000 and has spread to native species (Lindo
servation areas. et al. 2002), but the effect on snail populations is un-
In addition to the larger areas of good forest, small known.
patches are scattered through many of the limestone Naturalized introduced plants in general seem to
areas. Lewis & Swabey’s observations (1945:54) still have had little effect on Jamaican snails. Logwood
hold true today: “...in the limestone country of the cen- savannahs harbor few snail species, typically only 3
tral and western parishes...the natural vegetation has or 4 per site, perhaps because they are associated with
in many places been saved. The stony hills and hill- cattle, or perhaps because they do not provide consis-
sides, which are so characteristic of these parishes, are tent shade. Stands of Caribbean pine also have few
unsuitable for grazing or cultivation and afford valu- snail species, presumably because the needles are acidic
able sanctuary for wildlife.” Diesel and Horst (1995) and do not form good leaf litter. There appear to be no
found that endemic grapsid crabs inhabit man-made obligate relationships of Jamaican snails with particu-
stone walls in abandoned plantations overgrown by lar plant species, and it is not unusual to find a variety
secondary forest. Similarly, endemic snails are often of native snails living in areas with many introduced
found in areas of secondary growth and even in plan- broad-leaf plant species. Overall, deforestation and the
tations, so long as leaf litter is allowed to accumulate accompanying drying out of the landscape are much
(personal observation). All forested areas are never- more a threat to the native fauna than introduction of
theless under threat because of growing population, plant species.
TERRESTRIAL MOLLUCA OF JAMAICA 125
Colobostylus interruptus is known only from Long and posedly extinct species, the Jamaican Iguana was found
Dallas Mountains east of Kingston, an area of about alive in 1990 in the Hellshire Hills. We estimate that
24 km2. The geological setting, limestone with lots of perhaps as much as 90% of the terrestrial molluscan
unconsolidated rubble, makes it likely that those two fauna still survives, although many species have un-
mountains represent the species’ total original range: dergone great reduction in geographic range. Perhaps
there is alluvium to the west, non-limestone areas to the majority of species have part of their ranges in pri-
the north and northeast; massive limestone with virtu- mary forest, that, if protected, should ensure the long
ally no loose rubble to the east, and ocean to the south. term survival of the species. Areas outside the current
Gloyne (1872: 38) stated, “This species appears almost and proposed parks are under extreme threat of defor-
extinct. I found a great number of examples dead and estation, yet they harbor many species of restricted
worn on Long Mountain, but neither my friend, Mr. geographic range that do not occur in the conservation
Vendryes, nor myself, have ever been able to collect areas. Because the fauna is largely extant, no better
even one alive. Perhaps its disappearance may be at- place exists anywhere in the world for studies of land
tributed to the greater dryness of the climate in these snail evolution and speciation.
last years?” Jarvis (1903), said “Living specimens of
this species are very rare, and only occasionally found SYSTEMATICS
on Dallas Mountain, but weather-beaten shells are very
abundant...” No live material is among that collected In compiling the list of Jamaican species presented
by H. B. Baker in 1933. No original forest remains in in the Appendix, we have avoided taxonomic innova-
this area, although there is much secondary growth, tions, but it has not always been possible to do so. For
particularly on the northern slopes. This seems, there- example, the former subgenera of Hyalosagda have
fore, like a species for which little hope is left. now all been raised to full genera, but not all species
If our 1999-2002 survey had placed sites at ran- in the former subgenera have previously been cited in
dom, probably only one would have been within the the new combinations. We have not flagged such new
Long/Dallas Mountain area (607 sites/11,580 km2 = 1 combinations in the list, but other nomenclatural
site/19 km2), and our chances of finding Colobostylus changes are detailed below. A summary of the major
interruptus alive would have been minimal. Since we changes is given in the results section. Most of the taxa
preferentially sampled forested areas and areas of geo- treated below are illustrated in an interactive key by
logical interest, our original study design would have Rosenberg & Drumm (2004).
justified three sites. In an attempt to find C. interrup-
tus and another area endemic, Parachondria armata Helicinidae
(discussed earlier), we placed six additional sites on
Long and Dallas Mountains in March 2002, for a total Priotrochatella pulchra (C. B. Adams, 1851a).
of nine. Long dead shells of C. interruptus were found Adams described this as a variety of P. josephinae, but
at every site, shells with remnants of color at a few noted that it was “quite distinct”. Clench & Jacobson
sites, and fresh dead shells with operculum in place at (1970) synonymized it with P. josephinae on the basis
only one site, JBS576 on the east side of Dallas Moun- of unlocalized mixed lots. We recognize it as a full
tain opposite Dallas Castle. If we have the opportunity species. Priotrochatella pulchra is restricted to west-
to return to this site at night after rain during the rainy ern Jamaica, primarily in the Dolphin Head area; it
season, we will likely find the species alive. A similar reaches a larger size and has weaker spiral sculpture
or much greater level of effort would have to be de- and more prominent peripheral sculpture than
voted to every candidate species to determine its sta- Priotrochella josephinae, which is restricted to St.
tus. To perform such surveys, it may be possible to Thomas Parish in eastern Jamaica. Both species live
enlist the help of local environmental groups, of which on limestone outcrops in forested areas. A third popu-
there are a growing number in Jamaica. lation of Priotrochatella, possibly representing an
Jamaica is comparable to the Hawaiian islands as undescribed species occurs in the remote karst area in
a site of endemic radiation and diversification of land southwestern St. Ann.
snails. Thirteen families have endemic radiations, some
of which include endemic genera. Unlike Hawaii and
other oceanic islands in the Pacific, most of the spe-
cies are still extant. Even among Jamaican birds and
reptiles, only a few species are extinct, and one sup-
TERRESTRIAL MOLLUCA OF JAMAICA 127
Pilsbry took it for Jamaican. It is unlikely that the lo- that its systematic position was uncertain. Pilsbry did
cality refers to St. Thomas Parish, Jamaica: Brown not examine any specimens of this species; there are
obtained the specimens from Thomas Bland, who re- none at ANSP; he noted that the type material was not
ported Pupa jamaicensis from the island of St. Tho- in the Adams collection at Amherst but probably in
mas in 1861 (p. 359). Pupa jamaicensis is now con- the Chitty collection as Pfeiffer (1859) had redescribed
sidered a synonym of Gastrocopta pellucida a close the species from material at the British Museum. We
relative of G. polyptyx. As there are no authentic records were unable to locate type material of this species at
of G. polyptyx from Jamaica, we exclude it from the BMNH on recent visits (by Rosenberg, February and
fauna. December 2005). Based on the original description, it
is unlikely that this species is a spiraxid. It is 17 mm
Bulimulidae (Pfeiffer 1859), larger than members of any spiraxid
genus except Pseudosubulina (size ranges given by
Drymaeus rufescens (Gray, 1825) and Drymaeus Schileyko 2000), and has scattered stripes of dark
erubescens (Pfeiffer, 1847). Drymaeus erubescens was brown, a coloration not otherwise known among Ja-
described without locality; it was first reported from maican spiraxids. Size and coloration are both consis-
Jamaica by C. B. Adams (1849c). Pilsbry (1930) noted tent with Jamaican oleacinids; and the “nearly straight
that the species had not recently been collected in Ja- columella” and “distant striae” lead us to place the spe-
maica, but accepted it as Jamaican on the strength of cies in Sigmataxis.
two records in the ANSP collection and the Jamaican
type locality of Drymaeus rufescens (Gray, 1825), Urocoptidae
which he considered a senior synonym of D.
erubescens. One of the ANSP specimens (25845), as Mychostoma Albers, 1850. The type species of
Pilsbry noted, says on the original label “probably from this genus is Helix subula Férussac, 1832 by subse-
Honduras”, the other (ANSP 3553) is actually a slen- quent designation of Martens (1860). Pilsbry (1903 in
der juvenile specimen of the Jamaican endemic, 1902-1904) demonstrated that Helix subula is a conti-
Drymaeus immaculatus. If one supposes that C. B. nental species of Brachypodella that subsequent au-
Adams’s 1849 record was based upon pale, slender thors had misidentified as a Jamaican species. No ge-
specimens of Bulimulus guadalupensis, a common neric name is available for the Jamaican species usu-
species conspicuously absent from his list, then there ally placed in Mychostoma. A new name is not intro-
are no records of this species from Jamaica subsequent duced here, since we think this group of species may
to 1825. We therefore consider the type locality of prove to be paraphyletic to Apoma. Pending resolu-
Drymaeus rufescens to be erroneous and exclude the tion of relationships, we maintain use of the name
species from the Jamaican fauna. The only authentic Mychostoma since it is not confusing, although it is
locality for the species is Old Providence Island (Isla incorrect.
de Providencia, Colombia).
Spirostemma intermedium (Sowerby II, 1875).
Oleacinidae This is a primary homonym of Cylindrella intermedia
C. B. Adams, 1849a. If further study of Spirostemma
Varicella griffithii (C. B. Adams, 1845). Origi- confirms that this is a good species, a replacement name
nally spelled “griffithsii”, but emended to “griffithii” will be needed.
by Adams (1851a). The latter spelling is in prevailing
use and is here accepted as the correct original spell- Urocoptis lata (C. B. Adams, 1850a). Preoccu-
ing under ICZN Article 33.3.1. pied by Pupa lata C. B. Adams, 1849b, a Urocoptis
(see below). Urocoptis producta (C. B. Adams, 1851b)
Varicella leucozonias striatella Pilsbry, 1907. is the next available name.
Named as a variety by Pilsbry (1907 in 1907-1908),
but H. B. Baker (1941) recognized it as a subspecies. Pupa lata C. B. Adams, 1849b. Pilsbry (1926)
It intergrades with the nominotypical species in sym- doubtfully classified this as a Columella (Vertiginidae),
patry and so is synonymized. but it proves to be the protoconch of a Urocoptis, prob-
ably Urocoptis sanguinea (Pfeiffer, 1845c).
Sigmataxis macrospira (C. B. Adams, 1851).
Placed in Spiraxis by Pilsbry (1907: 31), who stated
TERRESTRIAL MOLLUCA OF JAMAICA 129
treated under Microphysa are now placed in several simson (Pfeiffer, 1852), which was treated as a sub-
different genera. We classify Adams’ species as species of P. invalida by Baker (1935d). No intergrades
Stauroglypta alveus as it has greatest similarity to occur, so we recognize P. candescens as a full species.
Stauroglypta spreta among Jamaican sagdids. The It differs from P. invalida in having the outermost of
specimens we have examined (ANSP 820) are some- the four apertural teeth shaped like a rounded trapezoid
what worn but show traces of the diagonal threads that (Figure 6) rather than being sharply triangular (Figure
characterize Stauroglypta conchologically. 7), and in reaching a greater maximum size. The spe-
cies are apparently allopatric, although they have
Helix albicans Pfeiffer, 1850. Apparently not broader ranges than illustrated by Jarvis (1902b).
treated since Vendryes (1899); placed here as a ques- Pleurodonte candescens is found in Trelawny, St. Ann,
tionable synonym of Stauroglypta anthoniana (C. B. St. Mary and northern St. Catherine parishes; P.
Adams, 1849b) based on the original description. invalida is found along the south coast from Portland
and St. Thomas through St. Elizabeth parishes.
Pleurodontidae With the recognition of P. candescens as distinct
from P. invalida, a question arises as to the position of
Pleurodonte amabilis (C. B. Adams, 1850c). P. invalida consanguinea, another subspecies recog-
Adams described this as a full species but subsequently nized by Baker, and its two synonyms, Helix
placed it as a variety of P. cara (Adams 1851b). The consanguinea vars. carinata and imperforata. The den-
lectotype selected by Jacobson & Boss (1973, ANSP tition of H. consanguinea and H. carinata places them
9013) is invalid. There is no evidence that it is part of with P. candescens, but they are generally flatter and
the original material studied by Adams, having come more broadly umbilicate, and appear to be allopatric
from the Swift Collection, nor does it match the origi- in distribution. As shown in the Appendix, both names
nal description, being too high spired. The supposed are preoccupied, so they are listed as doubtful syn-
lectotype is a specimen of P. catadupae (see below); a onyms of P. candescens and the question of the status
supposed paralectotype in the same lot is P. cara. A of these populations is left for future study.
specimen in the Chitty collection (BMNH 54.4.3.30),
labeled Helix cara var. amabilis, exactly matches the Pleurodonte carmelita (Férussac, 1821). The
measurements given by C. B. Adams in the original original combination is Helix carmelita which is pre-
description (Figures 3-5). It is here designated the lec- occupied by Helix carmelita Lichtenstein, 1794. Gei-
totype of that species to end confusion about its iden- ger (2003) called Lichtenstein’s taxon a nomen dubium,
tity. a conclusion that we second. Unfortunately, Férussac’s
Specimens exactly matching the original descrip- name does not have enough usage to qualify for rever-
tion of Pleurodonte amabilis have been found in the sal of precedence (ICZN Article 23.9). The next avail-
central Cockpit Country and the species is here recog- able name, Helix mora Griffith & Pidgeon, 1834 was
nized as valid. Pleurodonte amabilis differs from P. first synonymized with H. carmelita by Pfeiffer (1846,
cara in that the reflected aperture covers less than half p. 73). Subsequent authors such as Pilsbry (1889 in
of the umbilicus rather than most or all of it, the outer 1889-90, p. 99), have supported this synonymy al-
two apertural teeth are distinct rather than merged at though the species was described without locality.
the base, the labrum meets the body whorl below the Griffith & Pidgeon’s illustration of Helix mora
peripheral keel rather than exactly at it, and white (pl. 36, fig. 1) was likely based on material at BMNH;
flammules mark where the periostracum is loose be- a footnote on p. 595 says, “Most of the inedited shells
cause of underlying grooves on the shell, dorsally and figured in this work are from the collection of the Brit-
ventrally. This last feature serves also to distinguish P. ish Museum”, and the name is attributed in the plate
amabilis from P. catadupae. caption to [John Edward] Gray, who worked there. One
of us (Rosenberg) examined two lots containing five
Pleurodonte candescens (C. B. Adams, 1850a). specimens of Pleurodonte carmelita of appropriate
Helix candescens was considered a variety of vintage from BMNH, but none of the labels demon-
Pleurodonte invalida by Pfeiffer (1853) and Pilsbry strated a connection to Griffith & Pidgeon, and none
(1889 in 1889-1890), but apparently neither of them of the specimens were a convincing match for the il-
saw Adams’ type material. Examination of the lecto- lustration. The type material of Helix mora is thus re-
type (MCZ 155775, designated by Jacobson & Boss garded as lost. The description says only “Chestnut
1973) proves P. candescens to be an older name for P. brown, edge of lips white” (p. 597), which is consis-
TERRESTRIAL MOLLUCA OF JAMAICA 131
Figures 3-7. Pleurodonte species. 3-5. Lectotype of Pleurodonte amabilis, BMNH, 54.4.3.30, 26.7
mm. 6. Lectotype of Pleurodonte candescens, MCZ 155775, 20.0 mm, showing trapezoidal outer tooth.
7. Pleurodonte invalida, ANSP 763, 19.9 mm., showing triangular outer tooth
tent with P. carmelita, but hardly diagnostic. The il- at the base; if there are two inner teeth, they are closer
lustration, a ventral view only, is consistent with P. together than in P. cara; the labrum meets the body
carmelita in size and more importantly in the distinc- whorl below the peripheral keel rather than exactly at
tive ridge of callus around the inner peristome. Among it, the apertural lip is brown rather than usually white,
Jamaican pleurodontids, this ridge occurs only in some and the whorls are bordered by a spiral brown band
fully adult specimens of P. carmelita. We thus accept both dorsally and ventrally that does not reach the pe-
Pleurodonte mora as the valid name for this taxon. ripheral keel, rather than being monochromatic or grad-
ing gradually from darker at the keel to lighter cen-
Pleurodonte catadupae (H. B. Baker, 1935c). trally. P. catadupae has been found only in St. James
Baker named this as a subspecies of P. cara, but no Parish on the western side of the Cockpit Country, P.
intergrades occur, so we recognize it as a full species. amabilis only in Trelawny Parish in the central Cock-
Pleurodonte catadupae differs from P. cara in that the pit Country, and P. cara only in Manchester Parish
outer two apertural teeth are distinct rather than merged south of the Cockpit Country.
132 G. ROSENBERG AND I. V. MURATOV
Helix (Lucerna) lindsleyana Vendryes, 1899. This Pleurodonte tridentina chittyana (C. B. Adams,
name was introduced by Vendryes (1899) in a list and 1849a) and P. t. schroeteriana (Pfeiffer, 1845a). Both
partial key to Jamaican species of land and freshwater were described as species, but treated as subspecies of
shells; there is no illustration or locality and only a P. tridentina by Baker (1935d). They intergrade in sym-
brief description. Vendryes attributed the name to patry with the nominotypical subspecies, and are here
Chitty, but we can find no evidence that Chitty intro- synonymized with it.
duced the name; it is not listed by Clench & Turner
(1950b). Helix lindsleyana is one of the last four spe- Xanthonychidae
cies in a section of the key treating what are now clas-
sified as Pleurodonte. Helix jacobensis C. B. Adams, 1851b. Listed as a
Helix lindsleyana is described by “Outer two teeth valid species by several authors, but known to them
scarcely or not at all united at base” but this phrase only from the original description. The type material
might also have been intended to apply to some or all is lost according to Jacobson and Boss (1973). We con-
of the three succeeding species, as Vendryes’ key was sider a synonym of Dialeuca conspersula platystyla
not dichotomous in structure. Each of the other three Pfeiffer, 1850, based on the original description and a
species, Helix pallescens, H. atavus and H. picturata comparison with a specimen identified as Helix
has additional descriptive phrases added. This arrange- jacobensis in the Chitty collection at BMNH
ment may be interpreted as a differential diagnosis, so (54.3.18.587). Adams did not compare his taxon to
we consider the name Helix lindsleyana to be avail- Pfeiffer’s, which was ascribed with doubt to the
able. Other characteristics of the species based on its Molucca Islands.
position in the key are 3-4 apertural teeth, globose body
whorl, and lack of a furrow above the periphery be-
hind the aperture. ACKNOWLEDGMENTS
We have not been able to trace type material of
Helix lindsleyana; it may be in the Vendryes collec- This paper is dedicated to the memory of Glenn
tion at the Institute of Jamaica in Kingston. Jarvis Goodfriend, who passed away on 15 October 2002.
(1902b) also attributed the name to Chitty and showed He was always supportive of our work on the Jamai-
distribution coextensive with Helix atavus, both being can fauna and willing to share his knowledge.
restricted to Western Westmoreland. We have not been At the University of the West Indies, Eric
able to locate Jarvis’s specimens; he may have been Garraway and Catherine Murphy provided invaluable
working from information communicated by Vendryes, logistic and moral support. His graduate students,
whose help he acknowledged in a related paper (Jarvis Lauren Munroe and Natalie Blake, joined us for se-
1902a). Pleurodonte pallescens does not occur in lected field work, and along with Sarah Watson and
Westmoreland (it is apparently a Cockpit endemic) and David Drumm, assisted with sorting and identifying
P. picturata has the outer two teeth somewhat united the samples in Philadelphia. Many other faculty and
at the base. This leaves the most likely identity for Helix staff at the University aided us in various ways: Ralph
lindsleyana as being a form of P. atavus, so we tenta- Robinson, Byron Wilson, Peter Vogel, Ian Brown,
tively synonymize it with that species, although it is Simon Mitchell, Dunbar Steel, and George Proctor
possible that it will turn out to be a form of P. picturata (who told us stories of Henry Pilsbry when we first
if type material is ever discovered. No other species of met him botanizing in Barbecue Bottom). At the Insti-
Pleurodonte are consistent with the characteristics de- tute of Jamaica, Elaine Fischer, Tracy Commock, and
tailed by Vendryes. Elizabeth Morrison encouraged our interest in the fauna
and in their collections. Elaine Donaldson willingly
Helix soror var. latior Deshayes, 1851. This name assisted with permits at NRCA (now NEPA). Tommy
was introduced in a plate caption and apparently has Allen, Jason Bond, Peter Clarke, Paul “Herlitz” Davis,
not been treated by any subsequent author. Pfeiffer Rohan Esty, Tom Grace, Davile “Raymond” Holmes,
(1853) listed the figure on which the name was based Sam James, Mark Kitson, Susan Koenig, Michael
under Helix peracutissima C. B. Adams, 1845, a place- “Claude” Lewis, Leon Linton, Sylvia Macia, Marcia
ment with which we concur. Helix latior is thus a syn- Mundle, Michael Norris, Daniel Perez-Gelabert,
onym of Pleurodonte peracutissima. Dwight Pryce, Mike Robinson, Granville Spence,
Menocal Stevenson, Christoph Schubart, Michael
Schwartz, Ann Sutton, Marius van der Merwe, Jason
TERRESTRIAL MOLLUCA OF JAMAICA 133
Weintraub, and Clinton West all assisted us in the field, -----. 1851a. Description of new species and varieties
in various capacities of guides, assistants, colleagues of shells, which inhabit Jamaica. Contributions
and friends. John Taylor and Kathie Way (BMNH), to Conchology 8: 129-140.
Fred Thompson and Roger Portell (FLMNH), Paula -----. 1851b. Descriptions of new species and varieties
Mikkelsen and Jay Cordeiro (AMNH), Kenneth J. Boss of the land shells of Jamaica, with notes on some
and Adam Baldinger (MCZ), and M. G. Harasweych, previously described species. Contributions to
Raye Germon and Paul Greenhall (USNM) readily Conchology 9: 153-174.
loaned specimens or allowed us access to collections -----. 1851c. Catalogue of the land shells which inhabit
under their care. G. T. Watters commented on the no- Jamaica. Contributions to Conchology no. 9: 179-
menclature of Annulariidae. David Robinson willingly 186.
advised on the identification of introduced species. Dan Adams, C. D. 1972. Flowering Plants of Jamaica. Uni-
Otte and Paul Callomon helped with preparation of versity of the West Indies: Mona, Jamaica. 848
the figures. C. R. C. Paul and an anonymous reviewer pp.
provided valuable comments on the manuscript. This Albers, J. C. 1850. Die Heliceen, nach natürlicher
research was supported NSF Grants DEB-9870233 and Verwandtschaft systematisch geordnet. Enslin:
DEB-9408089 to Gary Rosenberg. Any opinions, find- Berlin. [ii], 262 pp.
ings, and conclusions or recommendations expressed Anderson, S., C. A. Woods, G. S. Morgan and W. L. R.
in this material are those of the authors and do not nec- Oliver. 1983. Geocapromys brownii. Mammalian
essarily reflect the views of the National Science Foun- Species 201: 1-5.
dation. Asprey, G. F. and R. G. Robbins. 1953. The vegetation
of Jamaica. Ecological Monographs 23:359-412.
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TERRESTRIAL MOLLUCA OF JAMAICA 139
Helicinidae 40 74 54 40 100 11
Proserpinidae 4 8 50 4 100 2
Stoastomatidae 75 82 91 75 100 4
Neocyclotidae 68 85 80 68 100 10
Annulariidae 65 120 54 65 100 9
Truncatellidae 17 36 47 17 100 4
Operculates 269 405 66 269 100 40
Carychiidae 1 3 33 1 100 1
Veronicellidae 9 17 53 3 33 4
Succineidae 4 4 100 1 25 1
Lauriidae 1 3 33 0 0 1
Pupillidae 3 6 50 0 0 2
Vertiginidae 9 12 75 0 0 4
Strobilopsidae 1 2 50 0 0 1
Helicodiscidae 2 2 100 0 0 2
Punctidae 2 2 100 0 0 2
Ferussaciidae 3 4 75 0 0 2
Subulinidae 10 17 63 1 10 7
Spiraxidae 7 13 54 7 100 5
Oleacinidae 57 92 62 57 100 7
Bulimulidae 4 5 80 1 25 2
Strophocheilidae 1 1 100 0 0 1
Orthalicidae 1 1 100 0 0 1
Urocoptidae 73 153 48 72 99 10
Thysanophoridae 2 2 100 1 50 1
Sagdidae 56 80 70 56 100 12
Streptaxidae 1 1 100 0 0 1
Systrophiidae 3 3 100 1 33 2
Polygyridae 1 1 100 0 0 1
Pleurodontidae 31 78 40 30 97 4
Bradybaenidae 1 1 100 0 0 1
Xanthonychidae 5 13 38 5 100 2
Agriolimacidae 1 1 100 0 0 1
Euconulidae 1 3 33 0 0 1
Zonitidae 3 3 100 0 0 3
Pulmonates 293 523 56 236 81 83
Total 562 928 61 505 90 123
140 G. ROSENBERG AND I. V. MURATOV
Table 2. Distribution of non-endemic species of terrestrial mollusks from Jamaica. Of total of 57 species,
10 are native (N), 25 introduced by humans (I), and 22 are of unknown status (?).
This list gives the current classification of all named terrestrial molluscan species known from Jamaica.
Undescribed, unidentified, misidentified and extralimital species are excluded, as are species known as fossils
only. An asterisk indicates that the name is represented by Jamaican specimens in the malacology collection at
ANSP. Subspecies are not numbered separately, but are listed at the same level of indent as species (e.g., num-
bers 2 and 10). A plus sign (+) before a name indicates that it is a synonym or possible synonym (+?) of the name
in the numbered heading. Synonymies exclude names not applied to the Jamaican fauna. The original genus is
given in square brackets if it differs from the current genus.
NERITOPSINA
HELICINIDAE
Classification from Wagner (1907-11), Baker (1922, 1934a, b, 1940), Boss & Jacobson (1974a, b), Clench &
Jacobson (1970) and Richling (2004).
PROSERPINIDAE
STOASTOMATIDAE
Classification follows Chitty (1857b) as modified by Baker (1934a, b) and Boss (1972).
ARCHITAENIOGLOSSA
NEOCYCLOTIDAE
Classification follows Bartsch (1942) as modified by Baker (1943b) and Morrison (1955).
CAENOGASTROPODA
ANNULARIIDAE
Classification follows Henderson & Bartsch (1920) with some modifications by Baker (1934b, 1935a), omit-
ting subfamilies.
TRUNCATELLIDAE
Generic classification based on Clench & Turner (1948); specific classification based on Rosenberg & Muratov
(in prep.).
PULMONATA
CARYCHIIDAE
VERONICELLIDAE
SUCCINEIDAE
LAURIIDAE
PUPILLIDAE
VERTIGINIDAE
STROBILOPSIDAE
HELICODISCIDAE
PUNCTIDAE
FERUSSACIIDAE
SUBULINIDAE
Classification from Pilsbry (1906-1907), Baker (1935c), Robinson (1999) and Schilekyo (1999b).
SPIRAXIDAE
Classification from Pilsbry (1907-1908), Baker (1935c, 1939) and Schileyko (2000).
OLEACINIDAE
Classification follows Pilsbry (1907-1908), Baker (1935a, c; 1941) and Schileyko (2000).
BULIMULIDAE
STROPHOCHEILIDAE
ORTHALICIDAE
UROCOPTIDAE
Classification from Pilsbry (1902-1904), Sykes (1904), Baker (1935c, d), Paul (1982, 1983) and Schileyko
(1999a), omitting subfamilies.
THYSANOPHORIDAE
SAGDIDAE
Classification follows Baker (1935b, d, 1950), Richardson (1986), Goodfriend (1986b), and Schileyko (1998b).
STREPTAXIDAE
SYSTROPHIIDAE
POLYGYRIDAE
PLEURODONTIDAE
Classification follows Pilsbry (1889-1890), Baker (1935d) and Goodfriend (1983). Pleurodontidae is recog-
nized as a family, rather than a subfamily of Camaenidae following Bouchet et al. (2005).
ERRATUM
Rosenberg, G. 2006. Status Report on the Terrestrial Mollusca of Jamaica. Proceedings of the Academy of Natural Sci-
ences of Philadelphia 155: 117-161.
Due to a production error, the concluding page of the taxon list in the appendix to this paper was omitted. It is
provided below. Also, the running heading gave the date of publication as April 2005, but it was May 2006.