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All content in this area was uploaded by Juan Manuel Diaz
Content may be subject to copyright.
Caribbean Journal of Science,
Vol. 31, No. 1-2, 104-121, 1995
Copyright 1995 College of Arts and Sciences
University of Puerto Rico,
Mayaguez
Zoogeography of Marine Gastropod in the Southern
Caribbean: A New Look at Provinciality
JUAN M. DIAZ
Instituto de Investigaciones Marinas de Punta de
Betin
-lNVEMAR-,
Apartado 1016, Santa Martar Colombia
ABSTRACT . – Recorded occurrences of 266 species of higher Gastropoda were arranged in nine subareas
along the coastal areas of southern Central America and northern South America. The value of different
gastropod families as zoogeographic indicators and the degree of faunal “singularity” of subareas were
inferred respectively from the mean value of the index of “Breadth of Geographic Range” (BGR) of species
involved. BGR was lower among families exhibiting predominantly direct development. About 43% of
species are widely distributed in the tropical Western Atlantic, whereas approximately 40% are endemic
to the southern Caribbean – enough to consider this area as a separate zoogeographic province. Northern
Venezuela, Santa Marta, and the Leeward Islands are the subareas richest in species, whereas the subarea
between the Orinoco delta and Surinam is the most depauperate. The number of species occurring in a
given subarea is more closely related to environmental heterogeneity than to the shelf extent of the subarea.
A similarity level of 50% distinguishes five zoogeographic areas within the lower Caribbean, two of them
as transitional to other tropical Western Atlantic Provinces and the other three are proposed as subprov-
inces. Trade wind-induced upwelling along the coasts of northern Colombia and Venezuela on the one
hand, and zoogeographic links of the present molluscan fauna to the Eastern Pacific on the other, are the
main factors explaining the present distribution patterns of marine gastropod in the southern Caribbean.
RESUMEN. – Los registros de 266 especies de gasteropodos superiors fueron agrupados en 9 subareas a
lo largo de las costas del Caribe entre el sur de
Centroamerica
y el norte de
Suramerica.
El valor de las
diversas familias de
gasteropodos
como indicadores zoogeograficos y el grado de "singularidad"
faunistica
de las subareas fue deducido a partir del valor promedio del indite de “Amplitud del Rango Geografico”
(ARG) de las especies incluidas. El ARG fue menor en las familias con desarrollo larval directo. El 43% de
las especies
estan
ampliamente distribuido en el
Atlantico
Occidental tropical y el 39.8% son
endemicas
del sur del Caribe – suficientes para considerar esa area como una provincia zoogeografica aparte. Venezuela,
Santa Marta y las Islas de Sotavento son las subareas con mayor numero de especies, mientras que la
subarea comprendida entre el delta del Orinoco y Surinam es la mas pobre en especies. El
numero
de
especies en una determinada
subarea
esta mas relacionado con el grado de heterogeneidad ambiental que
con las dimensioned de la misma subarea. Un nivel de similaridad del 50% define cinco areas zoogeograficas
en el sur del Caribe, dos de ellas consideradas zonas de transition hacia otras provincial del
Atlantico
Occidental tropical y otras tres propuestas como subprovincias. La surgencia inducida por los vientos
alisios en las costas septentrionales de Colombia y Venezuela, y las relaciones zoogeograficas de la ma-
lacofauna actual con la del Pacifico Oriental, son los factores que mejor explican los esquemas actuales de
distribution de los
gasteropodos
marines en el sur del Caribe.
INTRODUCTION
The coastal and shelf areas of southern
Central America and northern South
America as far as the Orinoco delta are
known as the southern Caribbean marine
region. This area was frequently disre-
garded from biotic and biogeographic
studies, and its fauna considered as “typ-
ical” West Indian-Caribbean (Houbrick,
1968; Bayer et al., 1970; Briggs, 1974), al-
though Rehder (1962) and Work (1969)
stated that better knowledge of the fauna
of the lower Caribbean could result in di-
viding the region into zoogeographical
subregions. Meyer (1973) and Moore (1974)
first referred to the coexistence of wide-
ranging Caribbean species with remark-
ably high numbers of poorly known en-
demic elements along the shores of north-
ernmost South America. More recently,
based on molluscan distributions, Cosel
(1976, 1982, 1986), Petuch (1976, 1981,
104
SOUTHERN CARIBBEAN GASTROPOD
105
1982a, b, 1987, 1990), Gibson-Smith and
Gibson-Smith (1979), and Diaz and Getting
(1988) provided further evidence that cer-
tain areas in the southern Caribbean ex-
hibit a somewhat “anomalous” faunal
composition. Many endemic molluscan
species have been described in the last two
decades from the southern Caribbean (e.g.,
Bayer, 1971; Altena, 1975; Petuch, 1979,
1987, 1990; Diaz and
Velasquez,
1986; Jong
and Coomans, 1988).
The purpose of this paper is to document
the faunal heterogeneity along the south-
ern Caribbean coasts, and to test the some-
what intuitively claimed existence of mol-
luscan faunules or “anomalously” com-
posed areas, and its significance as zoo-
geographic areas. The analysis is based on
the recorded occurrences of species from
selected families of prosobranch gastro-
pod that can be considered zoogeographic
“indicators.” The origin of the present bio-
geogeographic relationships of molluscan
faunas in the southern Caribbean are dis-
cussed according to the emerging ideas.
METHODS
Selection of Taxa. —Marine molluscs, par-
ticularly gastropod, have been tradition-
ally used for determining paleobiogeo-
graphic and zoogeographic patterns (e.g.,
Coomans, 1962; Hall, 1964; Woodring, 1974;
Petuch, 1982a, b; Kohn, 1990). Many neo-
gastropods exhibit a non planktonic de-
velopmental mode or have lecithotrophic
larvae with very low dispersal capacity (cf.
Radwin and Chamberlain, 1973), so they are
often very restricted in their habitat pref-
erences. The durable shells of most pro-
sobranch gastropod preserve well as fos-
sils and allow direct access to the paleo-
biogeographical record. Furthermore, gas-
tropod are common collecting objects, thus
systematic lists or species inventories are
available for many areas.
Twenty-two families of higher caeno-
gastropods were selected. All but two, Cy-
praeidae and Ficidae, belong to the Neo-
gastropoda. Some families, such as Nassar-
idae and Turridae, were not included be-
cause most of their members are too small
(less than 5 mm) and taxonomically very
controversial, so that they are often dis-
regarded or misnamed in species lists. This
criterion was also used to exclude several
species belonging to the families Colum-
bellidae, Vexillidae, and Marginellidae.
The species chosen inhabit exclusively
coastal habitats and the upper shelf zones
to 100 m, for at least part of their range.
Deeper occurring species have been often
recorded only from a few localities.
Recent works including lists of gastro-
pod species from the countries of this re-
gion and additional records, plus species
inventories from various localities were
used to develop a composite list of species
that occur in the region (see Appendix).
Emphasis was placed on revisionary works
such as Vokes and Houart (1986) and Vink
and Cosel (1985) rather than more general
works. Especial caution was devoted to se-
lecting the species described recently by
Petuch (1987, 1990), as some of them
(mainly in the families Olividae and Con-
idae) were described on the basis of subtle
shell characters of single specimens falling
into the variation range of known taxa.
Only records where a species was specifi-
cally listed from an area were used; distri-
bution maps suggesting that the range of
a species should include one of the areas
examined, but not specifying a locality,
were disregarded.
Subareas. —Although the term “Southern
Caribbean” refers conventionally to the
area south of latitude 15°N to the conti-
nental coasts of Central and South America
(from the Honduras-Nicaragua boundary
to Trinidad), for the purposes of this paper
the “Southern Caribbean” embraces the
shores and shelf areas along the continen-
tal coasts of Central and South America,
from northern Costa Rica to Surinam-
French Guiana, including the Leeward An-
tilles off Venezuela (Aruba, Bonaire, Cu-
racao, Margarita, and other smaller Vene-
zuelan islands). Nicaragua and Trinidad
have been disregarded, because data on
their molluscan faunas are very scarce.
However, since the molluscan fauna of the
area between the Orinoco delta and Suri-
nam has been adequately documented, it
has been included for this analysis.
According to broad environmental fea-
tures and the availability of faunal infor-
106
J. M. DIAZ
FIG.1. The nine subareas selected for the analysis. A, Aruba; C, southern half of the Colombian Caribbean
coast; G, Goajira and Paraguana peninsulas; LI, Leeward Islands; M, Santa Marta; O, Orinoco delta-Surinam;
P, Panama; R, Costa Rica.
mation, the region was partitioned into
nine subareas (Fig. 1). This paper compares
the molluscan composition of these sub-
areas. The data available from Costa Rica
and Panama does not allow the creation of
more subareas there, since most of the lit-
erature or specimen records list simply
“Panama” or “Costa Rica” as collecting
sites. Aruba (A) has been singled out from
the remainder Leeward Islands because “its
marine fauna is to certain degree distinct
from that of Curacao and Bonaire” (Jong
and Coomans, 1988). Furthermore, the sea
floor around these islands drops away rap-
idly to a depth of almost 1000 m, whereas
the maximum depth between Aruba and
the Venezuelan coast does not exceed 135
m. A small portion of the Caribbean coast
of Colombia from the vicinity of Santa
Marta eastwards to the sector where the
easternmost spurs of the Sierra Nevada de
Santa Marta drop into the sea has been also
singled out (M), as it represents a well doc-
umented distributional boundary for some
molluscs (Cosel, 1976, 1982, 1986; Diaz and
Getting, 1988; Diaz, 1990). Colombia and
Venezuela have or share three subareas,
which can be recognized on the basis of
both ecological and faunal features. An
outline of the environmental features of
each subarea is presented in Table 1.
Comparative Analysis.— I used Czechan-
ovsky’s presence-absence index of similar-
ity, which is widely used to determine af-
finities among biotic assemblages, and is
also employed to compare species overlap
between geographic areas (e.g., Crovello,
1981; Wells, 1990). The formula is:
2c x 100
S=a+b'
where a is the number of species in subarea
A, b the number in subarea B, and c the
number of species in common. The index
ranges from 0 (no species in common) to
100 (total overlap). A dendrogram was pro-
duced from the similarity matrix using the
group-average sorting strategy.
Since most species do not occur through-
out the region and many are even endemic
to a subarea (or their range in the Lower
Caribbean embraces only one or a few of
the subareas), every species does not have
the same value as indicator for defining
SOUTHERN CARIBBEAN GASTROPODS
107
TABLE 1. Major environmental features of the shelf, shores, and water masses in the subareas examined
(extracted from IUCN [1979] and Wells [1988]).
Shelf, bottom Water Habitats
Narrow; mud, sand
R
— —
(Costa Rica)
P(Panama)
C
(Southern Colombia)
M(Santa Marta)
G(Goajira-Paraguana)
A(Aruba)
LI
(Leeward Islands)
V
(Central Venezuela)
O(Orinoco-Guiana)
Narrow; fringing cora-
Iine archipelago;
mud, sand, gravel
bottoms
Rather narrow; predom-
inantly mud
Extremely narrow to
absent; steep; mud,
sand, gravel
Rather narrow to wide;
sand, gravel, mud
Narrow; sand, gravel,
rocks
Absent; steep; sand,
gravel, rocks
Very irregular; absent
to very wide; mud,
sand, gravel, rocks
Very wide; predomi-
nantly mud
Calm to agitated; turbid
and somewhat brackish
in the north, rather clear
in the south
Predominantly clear; rath-
er calm
Calm; predominantly tur-
bid; influenced by dis-
charges of major river
systems
Agitated to calm; rather
clear to turbid; seasonal-
ly affected by upwelling
Agitated; turbid; somewhat
brackish in the north;
permanently affected by
upwelling
Clear; agitated to calm
Clear; agitated to calm
Rather turbid; agitated; af-
fected by upwelling in
the eastern part
Turbid; agitated; somewhat
brackish; affected by dis-
charges of major river
systems
Estuaries; scattered man-
groves sandy shores; sea-
grass; poorly developed
coral reefs
Well developed coral reefs;
mangroves; seagrass;
scattered rocky and
sandy shores
Estuaries; mangroves; scat-
tered seagrass-meadows
and coral reefs around
offshore islands; sandy
shores
Rocky shores; fringing
reefs; scattered seagrass
meadows; mangroves,
and sandy shores
Sandy shores; seagrass;
scattered mangroves; al-
gae meadows
Sandy shores; coral reefs;
mangroves; seagrass
Rocky and sandy shores;
well developed reefs;
mangroves; seagrass
Rocky and sandy shores;
scattered seagrass and
mangroves; poorly de-
veloped coral reefs
Sandy and gravel shores;
estuaries
the degree of faunal “singularity” of a giv-
en subarea. Consequently, each species was
given an equitable value according to the
extent of its geographic range within the
region. The “Index of Breadth of Geo-
graphic Range” (BGR) of a species is de-
fined as the percent of the total number of
subareas (or sites) considered in the anal-
ysis (nine in this case) occupied by a spe-
cies.
RESULTS
The 266 species considered in this study
(see Appendix) inhabit diverse coastal and
shelf environments, including rocky
shores, coral reefs, mangroves, seagrass
meadows, sand, muddy, gravel, and rubble
bottoms. Most species range in depth be-
tween 2 and 50 m, and only few inhabit
exclusively shallower (e.g., Purpura patula
and Terebra salleana) or deeper waters (e.g.,
Fulgurofusus brayi and Fusinus couei). The
great majority of the species are predators
on worms and other molluscs, some are
scavengers, and a handful occasionally eat
plant material. Shell size ranges from less
than 5 mm (several Columbellidae and
Marginellidae) to more than 400 mm (Tur-
binellidae).
Table 2 summarizes the number of gen-
108
J. M. DIAZ
TABLE 2. Number of genera, mean value of the index of Breath of Geographic Range (BGR) of the species,
total number of species, and distribution of the number of species in the nine subareas examined.
Families Gen- Spe-
era ties
Number of
Mean Number of species in subareas
BGR
RPCM GALIV O
Cypraeidae
Ficidae
Muricidae
Thaididae
Colubrariidae
Columbellidae
Melongenidae
Buccinidae
Fasciolariidae
Columbariidae
Volutidae
Harpidae
Cancellaridae
Turbinellidae
Olividae
Marginellidae
Volutomitridae
Mitridae
Vexillidae
Costellariidae
Conidae
Terebridae
2
6
1
2
18 45
27
1
3
10 33
2
2
818
817
1 1
3 6
2
2
33
2
4
5
24
9
30
1
2
1
3
419
2 3
1
28
38
89 266
36.1 3
4546
44
41
26.7 0
000
1
1
0
21
22.9 11 15 11 17 18 10 18 24 15
34.6 4
57
4
1
4
4
43
38.5 1
2 2
3
1
2
2
11
31.3 19 15 19 14 19 18 18 18 4
36.4 1
111
1
11
22
20.5 7
879
10 11
8
11 0
23.1 4
7 5 10 8 8 9
10 4
44.4 0
1
1
0
1
0 0
10
14.7 2
2 2
11
1
1
20
19.0 1
0
1
2
1
11
10
20.0 1
0
1
2
1
0 0
21
26.5 2
222
2
2
1
21
21.7 5
9
9
11
712 6
10 5
20.0 5
7 9 10 7
13 14 93
16.7 0
1
0020 0
10
25.5 2
3 2
1
0 2
1
21
18.1 6
52 10 2
11 8
51
9.5 0
0
0
0
30
0
10
20.5 8 10 10 14 12 11 12 66
31.6 2
545
4
5
4
62
x
= 23.8 81
102
100
120 108 117 112
124 51
era, the average for the species in each fam-
ily, as well as the number of species in the
nine subareas. The 22 families considered
for the analysis yielded 266 species be-
longing to 89 genera. Five families con-
tributed more than 60% of the species:
Muricidae (16.9%), Columbellidae (12.4%),
Marginellidae (11.3%), Conidae (10.5%),
and Olividae (9%). The genera Steironepion,
Nassarina, Aesopus (Columbellidae), Volvar-
ina, and Gibberula (Marginellidae) were
omitted because they are too small and
poorly understood.
Of the 266 species, 115 (43%) are widely
distributed in the tropical Western Atlan-
tic. Five occur also in the Eastern Atlantic
(Amphiatlantic species) and five more oc-
cur also in the Eastern Pacific (Amphiam-
erican species). Endemic to the lower Ca-
ribbean are 106 species (40%), plus three
Amphiamericans so far recorded only from
the lower Caribbean (e.g., Anachis varia).
Thirty-three species (12%) can be regarded
essentially as Antillean or West Indian, i.e.,
their distributions embrace predominantly
the Antillean arc but overlap partially one
or more of the subareas considered in the
analysis. In contrast to this, six moderately
widely distributed Caribbean species are
exclusively bound to the continental
shelves; i.e., their range does not include
the Antillean arc (e.g., Turbinella angulata).
Although western Atlantic species are
more or less homogeneously distributed
among all subareas, making 61–70%. of the
total species, both lower Caribbean and
“Antillean” species exhibit a tendency to
concentrate in certain subareas (Fig. 2).
The families Costellaridae, Volutidae,
Volutomitrididae, and Vexillidae obtained
the lowest BGR’s (their members tend to
have narrow distributions), whereas Col-
umbariidae, Colubrariidae, Melongenidae,
Cypraeidae and Thaididae achieved the
highest BGRs (the members of these fam-
ilies usually exhibited wide distributions).
SOUTHERN CARIBBEAN GASTROPODS
109
FIG. 2. Distribution of numbers of “Southern Caribbean” (upper number), “Antillean” (middle number),
and exclusive (=endemic, lower or third number) species of higher gastropod among the subareas.
Average BGR of all species was 23.8; i.e.,
most species in the lower Caribbean ex-
hibit a distribution range that encompasses
slighty more than two of the nine subareas
examined here.
Table 3 summarizes the number of spe-
cies recorded and their average BGR, as
well as shelf extent of the nine subareas.
Clearly, the shelf extent of the subareas is
not correlated with the number of species
recorded. The subareas richest in species
are V (47% of the total), M (45%), and A
(44%). LI (42%) and G (41%) are slightly
less diverse. In contrast, subareas R (31%)
and O (19%) definitely have less diverse
faunas.
The lowest average BGR was attained by
the species recorded from G, followed by
the species from LI and A. The highest
values are achieved by species occurring
in R, P, and C, whereas the species record-
ed from V and O attain moderate values.
Figure 3 shows subarea affinity based on
untransformed presence-absence data, ap-
plying the conventional Czechanovsky
measure of similarity and group-average
sorting. A broken line at the arbitrary sim-
ilarity level of 50% delineates two major
groups of subareas (A-LI-V-M and C-P-R)
and leaves subareas O and G detached.
The gastropod fauna of subarea O is the
most dissimilar in the southern Caribbean.
It is definitively impoverished, exhibits a
very low endemism degree, and 53%. of the
species occurring there are wide-ranging
in the Tropical Western Atlantic (53%). Al-
though species endemic to the southern
Caribbean are well represented (35.3%), the
distribution area of several “Brazilian en-
demics” begins elsewhere between the
Orinoco delta and French Guiana (e.g., Tur-
TABLE 3. Shelf extent, number of recorded species,
and average Breath of Geographic Range (BGR) of the
species recorded in the nine subareas analyzed.
Shelf Num-
extent* ber of
BGR
Area (km)2species average
Costa Rica (R)
Panama (P)
Southern Colombia (C)
Santa Marta (M)
Goajira-Paraguana
(G)
Aruba (A)
Leeward Islands (LI)
Central Venezuela (V)
Surinam-Guiana (0)
4400
26,000
22,000
140
45,000
250
2000
65,000
180,000
81
44.6
101
43.3
100
43.5
120 41.7
108
31.6
117
35.7
112
34.0
124 37.6
51
36.5
* Shelf width estimated from the shoreline to the
100 fathoms depth contour.
110
J. M. DIAZ
FIG. 3. Classification of nine subareas in the south-
ern Caribbean based on presence-absence data of 266
gastropod species, using the Czechanovsky measure
and group-average sorting. Two main groups of sub-
areas and two detached subareas are distinguished at
an arbitrary similarity level of 50%.
binella laevigata, Marginella cloveri). Zoogeo-
graphically, this subarea should be consid-
ered to be outside the Caribbean Sea.
Subareas V, M, A, and LI join together
in the dendrogram (Fig. 3) forming a some-
what diffuse group characterized by a spe-
cies-rich fauna. The close faunal affinity
between Aruba and the remaining Lee-
ward Islands off the Venezuelan coast is
evident, in spite of the apparent environ-
mental differences among both subareas.
This resemblance is principally due to the
relative high number of “Antillean” spe-
cies occurring there. Of a total of 33 An-
tillean species “invading” the Lower Ca-
ribbean, 20 and 26 have been recorded re-
spectively in subareas A and LI, making 17
and 22%. of the whole caenogastropod fau-
na in these subareas (“Antillean” species
yield only 4-10% of the gastropod fauna
in the remaining subareas). The main dif-
ference between these two subareas results
from a handful of endemics to one of them
(nine species in A, seven in LI) and higher
incidence of sand-mud dwelling species in
A (e.g., Ficus communis, Antillophos candei)
versus a higher number of rock-coral
dwelling gastropod in LI (e.g., Pygmaep-
teris lourdesae, Dermomurex pauperculus).
Subareas V and M share many wide
ranging Tropical Western Atlantic and
Lower Caribbean species, and some faunal
elements apparently restricted to upwell-
ing areas, i.e., occurring only in subareas
M, G, and V (e.g., Calotrophon velero, Fusinus
caboblanquensis). They also share several
“Antillean” species, which are in part
shared with subareas A and LI.
Subarea G is quite detached from the ad-
jacent subareas. Almost half of the species
endemic to the lower Caribbean occur
there, many of them being restricted to it
(15 species, or 14.7% of its caenogastropod
fauna).
Subareas R, P, and C form a group hav-
ing comparatively many wide-ranging
species and a lower degreee of endemicity.
Amphiamerican species are numerically
concentrated in these subareas.
DISCUSSION
Concerning their habitat preferences,
feeding habits, and sizes, the 266 species
analyzed are diverse enough to be consid-
ered a representative sample of the pro-
sobranch gastropod fauna in the region (cf.
Rosenberg, 1993).
The duration of planktonic life has zoo-
geographic significance, since the dispers-
al capability of a species partially deter-
mines its range of distribution. Thus, gas-
tropod species with direct development or
whose larvae have short free-swimming
stages show generally more restricted dis-
tribution than those with greater dispersal
capability (Perron and Kohn, 1985; Schel-
tema, 1989). As stated above, average BGR
was lowest among the families Costellar-
iidae, Volutidae, Volutomitridae, and Vex-
illidae, all them with predominantly direct
development (see Radwin and Chamber-
lain, 1973; Penchaszadeh, 1988). Single spe-
cies of these families have very limited dis-
tribution within the Lower Caribbean, so
that they are in part responsible for the
incidence of endemism in certain subareas,
such as G and P. On the other hand, av-
SOUTHERN CARIBBEAN GASTROPODS
111
erage BGR's are high among families as
Cypraeidae, Thaididae, Colubrariidae, and
Melongenidae, which have mostly plank-
tonic development (Radwin and Cham-
berlain, 1973; Bandel, 1976a).
Conversely, as one might expect from
traditional island biogeographic theory
(e.g., MacArthur and Wilson, 1967), the
species-area effect alone offers little to ex-
plain why some of the analyzed subareas
are currently species-richer than others.
As stated by Williamson (1988), the im-
portance of environmental heterogeneity
and historic biogeographic (antecedent)
factors must be taken into account in eval-
uating biogeographic studies involving
species-area relationships. In structurally
diverse habitats, coexisting species often
seem to specialize and to avoid competitive
exclusion by differential use of the phys-
ical structure. Although it is difficult to
evaluate all components of environmental
heterogeneity and whether it is a cause or
merely a correlate of species richness, it is
possible to take the degree of co-occur-
rence of various shelf-bottoms types, water
qualities, and habitats within a given sub-
area as a measure of its environmental het-
erogeneity. A coarse comparison of envi-
ronmental features (Table 1) suggests a
closer relationship with the total number
of species recorded from each subarea than
that resulting from consideration of shelf
extent. Subareas R and O stand out clearly
as the least environment-diverse and also
the least species-rich.
The gastropod fauna of the southern Ca-
ribbean may be conveniently divided into
four categories on the basis of their broad
distribution patterns: (A) Species widely
distributed in the Caribbean Sea or even
in most parts of the tropical Western At-
lantic (43% of the species); (B) amphiatlan-
tic species (3%); (C) species restricted to the
southern Caribbean, i.e., their whole range
falls within the studied region (40%); and
(D) Antillean species which extend into
the southern Caribbean (12%).
Briggs (1974) subdivided the tropical
Western Atlantic into three provinces ac-
cording to his “10 percent-rule” (i.e., when
10% or more of the species are endemic to
a given area, it is designated as a separate
province): Caribbean, West Indian, and
Brazilian. According to this scheme, the
studied area falls entirely within the “Ca-
ribbean Province,” which extends along
the continental coastline of Central and
South America from Tampico (Mexico) to
eastern Venezuela, including allopatrical-
ly the southern portion of the Florida Pen-
insula. Although the theoretical basis for
the establishment of these provinces has
been questioned (Voss, 1975), the scheme
has been adopted by Caribbean zoogeog-
raphers (e.g., Spivey, 1981;
Gonzalez
et al.,
1991 ). Briggs’ scheme contrasts with Pe-
tuch’s statement (1982a, b) that the Carib-
bean Molluscan Province may be divided
into northern and southern components
by a line of abrupt faunal shift at about
latitude 15°N. Further arguments in favor
of a north-south faunal division of the Ca-
ribbean Region were provided by Acero
(1985) on the basis of fish distribution pat-
terns.
The data discussed in this paper show
that the southern Caribbean harbors a gas-
tropod fauna with 40% of endemism,
enough to consider it a separate zoogeo-
graphic subregion. Nevertheless, Rosen-
berg (1993), by comparing gastropod lists
of scattered and limited areas in the West-
ern Atlantic, stated recently that no local
region of the Western Atlantic has more
than 4% endemics and subsequently no
faunal province subdivisions could be made
within this region. Since endemic species
of a particular province or subprovince are
not always restricted to a single locality or
country, and similarities between local
faunas are partly determined by habitat
availability, in order to determine zoogeo-
graphic differences within a wide region
faunal lists should be grouped first accord-
ing to similarities among them and then
one should examine similarities between
the groups. In this way, groups of allied
faunas are justly or better compared and
actual differences between broad areas
within the region become more evident.
The similarity analysis makes evident the
faunal heterogeneity and variation among
subareas within the southern Caribbean. A
similarity level of 50% seems adequate to
define biogeographic areas or subprov-
112
J. M. DIAZ
inces within the Southern Caribbean Prov-
ince. Clearly, faunal shifts between adja-
cent subareas are not always sharply de-
fined and the boundaries could be drawn
anywhere, at least statistically. However,
there may be valid ecological or zoogeo-
graphical reasons for locating the bound-
ary in a certain place.
Subarea O (Fig. 1) should not be consid-
ered as a subprovince itself but as a tran-
sition area towards the Brazilian Province.
The Orinoco delta is a zone of abrubt im-
poverishment of the Antillean-lower Ca-
ribbean fauna, whereas southwards from
it a gradual enrichment of Brazilian ele-
ments takes place (cf. Rios, 1985;
Cervigon
et al., 1992).
Although subareas A and LI are both al-
lied to M and V, the former exhibit about
the same number of southern Caribbean
and “Antillean” species (23 and 17% re-
spectively in A; 19 and 22’% in LI). Hence,
the Leeward Islands should be regarded as
a gradual transition area to the Antillean
or West Indian Province, Aruba being the
first step.
Subareas V and M represent a subprov-
ince (I propose the name “Samarian-Ven-
ezuelan Subprovince”) that harbors the
species-richest molluscan fauna in the
southern Caribbean. It is seasonally affect-
ed by trade wind-induced upwelling of
cold water (Bula-Meyer, 1977;
Cervigon
et
al., 1992) and is disjuncted by subarea G,
the latter being affected almost perma-
nently by upwelling (Bula-Meyer, 1977;
Corredor, 1979). Towards the east, this sub-
province has a fairly well defined bound-
ary at the eastern end of the Peninsula of
Paria, coinciding with a coarse change in
the environmental features (cf.
Cervigon
et al., 1992). On the other side, subarea M,
representing the allopatric western portion,
has a well defined faunal and environ-
mental boundary to the south. An abrupt
faunal shift at Santa Marta, caused possibly
by the combined barrier effect of an ex-
tremely narrow shelf and upwelling-in-
duced cold waters, has been documented
(Cosel, 1976, 1982; Diaz and
Gotting,
1988;
Diaz, 1990). The boundaries of this sub-
province to subarea G are not so well de-
fined (see below).
The shelf areas off La Goajira and Para-
guana
and the Gulf of Venezuela (subarea
G) are definitely a detached subprovince (I
propose the name “Goajira Subprovince”).
The uniqueness of the molluscan fauna of
this area was first revealed by Petuch (1976),
who announced subsequently the exis-
tence of a “Colombian-Venezuelan Neo-
gene relict pocket holding the oldest
known intact shallow water molluscan
fauna in the Western Atlantic” (Petuch,
1981:311). The same author later described
many of the former “living fossils” as new
species (Petuch, 1987), some of which ap-
pear dubious because they were described
from single specimens that may be only
geographic varieties of wide-ranging spe-
cies (cf. Diaz, 1990; Tursch and Huart, 1990).
The boundaries of this subprovince are ill
defined, but they can be conveniently set
at the northern tip of the
Paraguana
Pen-
insula (Cabo San Roman) and near Palo-
mino (about 60 km eastward from Santa
Marta). This is coincidentally about the
distribution range of Syphocypraea mus (cf.
Petuch, 1979).
The “Isthmian Subprovince” embraces
at least the coasts and shelf areas from near
Cienaga
(Colombia) to the Costa Rica-
Nicaragua boundary. It extends probably
to northernmost Nicaragua (Cabo Gracias
a Dies), where apparently a faunal shift
takes place (cf. Petuch, 1982b; Acero, 1985).
As pointed out by Cosel (1986), due to ma-
jor river discharges most coastal and shelf
areas in the southern Colombian Carib-
bean and Panama are characterized by
muddy bottoms and turbid waters with
lowered salinity. This could lead to the
presence of another zoogeographically iso-
lated area in the southern Caribbean. Ev-
idence of the presence of this faunal pocket
is the high number of mollusks, particu-
larly bivalves, having sibling species in the
eastern Pacific (Cosel, 1986). More recent-
ly, when describing some new gastropod
from Panama, Petuch (1990) proposed the
existence of a further relict pocket or faun-
ule along the Caribbean coast of Panama
and Costa Rica, the “Blasian Biogeographic
Subregion,” which, as shown above, en-
compasses also the southern part of the
Colombian Caribbean.
SOUTHERN CARIBBEAN GASTROPODS
113
FIG. 4. Proposed spatial arrangement of molluscan subprovinces in the southern Caribbean: 1, Isthmian;
2, Samarian-Venezuelan; 3, Goajira; 4, transition area to the Antillean province; 5, transition area towards the
Brazilian province.
Figure 4 summarizes the spatial arrange-
ment of molluscan subprovinces in the
lower Caribbean as proposed above. The
most plausible way to explain such zoo-
geographic schemes and faunal anomalies
is to search for vicariant events that cor-
respond to geological and paleoclimatic
episodes of known age.
The fact that most species endemic to the
southern Caribbean concentrate along the
continental shelf of northern Colombia and
Venezuela (Fig. 2) may be adequately ex-
plained from ecological factors causing vi-
cariance. As pointed out by Petuch (1976),
Meyer et al. (1978), and Vermeij (1978), the
trade wind-induced upwelling of cold wa-
ter along the coasts of Venezuela and
northernmost Colombia may restrict to
shallow water the distribution of many
species of molluscs and crinoids of this re-
gion. The isolating “cold water” condition
seems to have prevailed since the Tertiary
as can be deduced from the existence of a
Colombian-Venezuelan-Trinidad faunal
subprovince during the Miocene (Wood-
ring, 1974; Petuch, 1982a).
Shifts in oceanographic conditions after
the Pliocene emergence of the Isthmus of
Panama (Maier-Reimer et al., 1990), as well
as sea level fluctuations and changes in pat-
terns of upwelling and nutrient distribu-
tion in northern South America during the
Pleistocene (Jackson et al., 1993), caused
not only high rates of extinction (Olsson,
1972; Vermeij, 1978; Vermeij and Petuch,
1986) but also high rates of speciation (All-
mon et al., 1993; Jackson et al., 1993) and
further disjunction of the molluscan fauna
of that subprovince into three geographi-
cally discrete pockets, which are presently
evidenced as a Goajira Subprovince and a
disjoined Samarian-Venezuelan Subprov-
ince.
Likewise, the present Caribbean bound-
aries of the Isthmian Subprovince coincide
with the Central American-northern South
American Miocene faunal Subprovince (cf.
Woodring, 1974). Many gastropod from
the Plio-Pleistocene of Costa Rica also have
a Holocene distribution in the western Ca-
ribbean, extending approximately from
Honduras to Colombia (Robinson, 1993).
The latter was disjoined by the final closing
of the Isthmus of Panama and underwent
drastic environmental changes during the
Pleistocene, leading to the extinction of
many molluscan genera and species in the
Caribbean (Olsson, 1972; Petuch, 1982a;
114
J. M. DIAZ
Vermeij and Petuch, 1986). The present
molluscan fauna of the Isthmian Subprov-
ince is composed mainly of wide-ranging
Western Atlantic or pan-Caribbean spe-
cies. On the other hand, as the amount of
endemic species is fairly low, it seems that
the birth rate of new taxa in this subprov-
ince has been lower than in the remaining
subprovinces in the southern Caribbean.
Hence, whereas the loss in diversity dur-
ing the Pliocene and Pleistocene in many
areas of the Western Atlantic was compen-
sated for, mainly by speciation (Vermeij
and Rosenberg, 1993), in the Isthmian Sub-
province the loss was apparently compen-
sated mostly by invading species which
achieved wide distributions in the Western
Atlantic (Vermeij and Rosenberg, 1993).
Nevertheless, a significant amount of forms
related to the Panamic fauna of Western
Central and South America (amphiameri-
can and sibling species: 29% after Radwin,
1969; 17.5% after Kruckow, 1980; up to 58%
after Cosel, 1986) underscore the zoogeo-
graphic links to the Panamic-Eastern Pa-
cific.
Caenogastropod species distributions in
the southern Caribbean suggest defined
zoogeographic tendencies. They can be ex-
plained as a whole from a combination of
historic biogeographic and dispersal fac-
tors, as well as regional environmental fea-
tures. The proposed arrangement of gas-
tropod distribution patterns in the lower
Caribbean into three major discrete sub-
provinces should be further evaluated by
workers studying other marine taxa.
Acknowledgments. -I thank F.
Chavarria,
P. Hoeblich, and M. Puyana for comple-
mentary records of gastropod from Costa
Rica, Venezuela and Colombia respective-
ly. I also thank Dr. F. J. Borrero, Dr. L.
Botero, and two anonymous reviewers for
their helpful and constructive comments
on earlier drafts of the manuscript. For their
support I acknowledge the Instituto Col-
ombiano para el Desarrollo de la Ciencia
y la
Tecnologia
-COLCIENCIAS- and IN-
VEMAR.
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APPENDIX. Species considered for the analysis and subareas in the lower Caribbean where they have been
recorded (R: Costa Rica; P: Panama; C: southern Caribbean coast of Colombia; M: Santa Marta; G: Goajira-
Paraguana; V: northern Venezuela; A: Aruba; LI: Leeward Islands; O: Orinoco delta to Surinam).
Ficidae
Ficus comunis (Roding, 1798)
F. howelli Clench & Aguayo, 1940
Muricidae
Murex messorius Sowerby, 1841
M. chrysostoma Sowerby, 1834
M. olssoni Vokes, 1967
M. donmoorei Bullis, 1964
M. thompsoni Bullis, 1964
M. consuelae Verrill, 1950
M. blakeanus Vokes, 1967
M. sunderlandi Petuch, 1987
M. tryoni Bullis, 1964
Chicoreus brevifrons (Lam., 1822)
C. spectrum (Reeve, 1846)
C. mergus Vokes, 1974
Phyllonotus pomum (Gmelin, 1791 )
P. margaritensis Abbott, 1958
Siratus springeri Bullis, 1964
S. beauii Fischer & Bernardi, 1857
Paziella oregonia Bullis, 1964
Calotrophon nelero (Vokes, 1970)
Panamurex gatunensis (Brown & Pils., 1911)
Actinotrophon actinophorus (Dan, 1889)
Typhis expansus Sowerby, 1874
T. bullisi (Gertman, 1969)
T. tytyrus Bayer, 1971
Pterotyphis pinnatus (Broderip, 1833)
Risomurex withrowi Vokes & Houart, 1986
R. cf. gilbertharrisi (Weisbord, 1962)
R. deformis (Reeve, 1846)
Muricopsis muricoides (C. B. Adams, 1845)
M. oxytata (Smith, 1938)
M. praepauxillus (Maury, 1917)
M. huberti Radwin & D'Atilio, 1976
Murexiella mcgintyi (Smith, 1938)
G-V-A
V-O
R-P-C-M-G-V-O
G-V-A-LI-O
R-P-C-M-G
M-G-V-O
G-V-O
M-LI
G-V
G
O
R-P-C-M-G-V-A-LI-O
A-LI
R
R-P-C-M-V
G-V-A-LI
V-O
G-V-O
G-V-O
M-G-V-A-LI
G-V
P-C
M-V-O
P-C-M-O
V
R-P-C-M-G
M-G-V-A-LI
G-V
R-P-C
R-P-M-V
P-C-V-LI
LI
LI
A-LI
SOUTHERN CARIBBEAN GASTROPODS
117
APPENDIX . Continued.
M. edwardpauli Petuch, 1990 P
Favartia cellulosa (Conrad, 1846) C-M-V-A-LI-O
F. alveata (Kiener, 1842) M-V-A-LI
F. germainae (Vokes & D'Atillio, 1980)
LI
Trachypollia nodulosa (C. B. Adams, 1845) R-P-C-M-V-A-LI
T. didyma (Schwengel, 1943) P-M-V-A-LI-O
Dermomurex pauperculus (C. B. Adams, 1850) R-P-M-LI
D. kaicherae Petuch, 1987
G
D. alabastrum (A. Adams, 1864) R-P
Pygmaepteris juanitae Gibson-Smith, 1980 M-V
P. lourdesae Gibson-Smith, 1980 M-LI
Murexsul harasewychi Petuch, 1987
G
Columbariidae
Fulgurofusus brayi (Clench, 1959)
Thaididae
Purpura patula (L. 1758)
Thais deltoidea (Lamarck, 1822)
T. rustics (Lamarck, 1822)
T. haemastoma haemastoma (L. 1767)
T. haemastoma floridana (Conrad, 1837)
T. coronata coronata (Lamarck, 1822)
T. coronata trinitatensis (Guppy, 1869)
Colubrariidae
Colubraria lanceolata (Menke, 1828)
C. obscura (Reeve, 1844)
C. swifti (Tryon, 1881)
Melongenidae
Melongena melongena L. 1758
Pugilina morio L., 1758
Buccinidae
Bailya parva (C. B. Adams, 1850)
B. marijkae Jong & Coomans, 1988
B. intricata Dall, 1889
Cantharus tinctus (Conrad, 1846)
C. auritulus (Link, 1807)
C. karinae Usticke, 1959
Pisania pusio (L., 1758)
Engina turbinella (Kiener, 1835)
E. stootsi Jong & Coomans, 1988
E. demani Jong & Coomans, 1988
E. willemsae Jong & Coomans, 1988
Antillophos candei (d’Orbigny, 1842)
A. chazaliei (Dautzenberg, 1900)
A. cf. adelus (Schwengel, 1942)
Engoniophos unicinctus (Say, 1826)
Metula agassizi Clench & Aguayo
M. lintea Guppy, 1882
Mohnia kaicherae Petuch, 1987
Columbellidae
Columbella mercatorial
(L., 1758)
Minipirene dormitor (Sowerby, 1844)
Conlla ovulata (Lamarck, 1822)
P-C-G-V
R-P-C-M-G-V-A-LI
R-P-C-M-V-A-LI
R-P-C-M-V-A-LI
C
R-P-C-V-A-LI
C-O
P-C-M-V-O
R-P-C-M-LI-O
P-C-M-A-LI
M-A
R-P-C-M-G-V-A-LI-O
V-O
R
LI
R-P
P-C-M-G-V-A-LI
R-P-M-V-A-LI
M-G-V-A-LI
R-P-C-M-G-V-A-LI
R-P-C-M-V-A-LI
A
A
A
R-C-M-G-V-A
M-G-V
P-C-G
C-M-G-V-A-LI
V
R-P
G
R-P-C-M-G-V-A-LI
V-A-LI
R-P-V-A-LI
118
J. M. DIAZ
APPENDIX. Continued.
C, ovuloides (C. B. Adams, 1850)
Anachis obesa (C. B. Adams, 1845)
A. lyrata (Sowerby, 1832)
A. coseli Diaz & Mittnacht, 1990
A. cf. fraudans Jung, 1969
A. hotessieriana (d'Orbigny, 1842)
A. demani Jong & Coomans, 1988
A. sparsa (Reeve, 1859)
A. catenata (Sowerby, 1844)
A. sertulariarum (d’Orbigny, 1839)
A. pretri (Duclos, 1846)
A. pulchella (Blainville, 1829)
A. dicomata Dall, 1899
A. varia (Sowerby, 1832)
A. plicatula (Dunker, 1853)
Cosmioconcha nitens (C. B. Adams, 1845)
C. calliglypta (Dall & Simpson, 1901)
C. hurmfreyi Jong & Coomans, 1988
Mitrella ocellata (Gmelin, 1791)
M. lunata (Say, 1826)
M. nycteis (Duclos, 1846)
M. dichroa (Sowerby, 1844)
M. idalina (Duclos, 1840)
Nitidella nitida (Lamarck, 1822)
N. laevigata (L., 1758)
Decipifus sixaolus Olsson & McGinty, 1958
D. kirstenseni Jong & Coomans, 1988
Mazatlania aciculata (Lamarck, 1822)
Strombina pumilio (Reeve, 1859)
S. francesae Gibson-Smith, 1974
Fasciolariidae
Fasciolaria tulips (L., 1758)
Latirus infundibulum (Gmelin, 1791)
L. mcgintyi Pilsbry, 1939
L. cariniferus (Lamarck, 1822)
L. angulatus (Roding, 1798)
L. eppi Melvill & Shepman, 1891
Teralatirus ernesti (Melvill, 1910)
T. cayohuesonicus (Sowerby, 1878)
Dolicholatirus pauli (McGinty, 1955)
Leucozonia nassa (Gmelin, 1791)
L. ocellata (Gmelin, 1791)
Fusinus closter (Philippi, 1850)
F. couei Petit, 1853
F. helenae Bartsch, 1939
F. caboblanquensis Weisbord, 1964
F. eucosmius (Dall, 1889)
Harasewychychia harasewychi Petuch, 1987
Volutidae
Voluta musics L., 1758
V. uirescens (Lightfoot, 1786)
V. demarcoi Olsson, 1965
V. lacertina Petuch, 1990
Scaphella evelina Bayer, 1971
Volutomitra erebus Bayer, 1971
P-A-LI
R-P-C-M-G-V-A-O
R-P-C-M-G
C-M-G
M-G
M-LI
A
R-P-C-M-G-V-A-LI
R-P
C-M-G-V-O
P-M-V-A-LI
R-P-C-M-V-A-LI
A-LI
R-P-C
V-A-LI
P-G-V
C-M-O
A
R-P-C-M-G-V-A-LI
R-P-C-M-G-A-LI-O
P-M-A-LI
P-C-M-V-LI
A-LI
R-P-C-M-G-V-A-LI
R-P-C-M-G-V-A-LI
P-M-G-V-A-LI
LI
R-P-C-M-V
G-V
V
R-P-C-M-G-V-A-LI
P-C-M-G-V-A-LI-O
P-C-M-G-LI
R-P
M-V-A-LI
LI
A-LI
P-C-M-V-A-LI
M
R-P-C-M-G-V-A-LI
R-P-C-M-V-A-LI
M-G-V-A-O
V
G-V
M-G-V
O
G
M-G-V-A-LI
R-P-C
R
P
C
G
SOUTHERN CARIBBEAN GASTROPOD
119
APPENDIX . Continued.
V. persephone Bayer, 1971
Lyria leonardi Emerson, 1985
Harpidae
Morum oniscus (L., 1767)
Cancellomorum lindae (Petuch, 1987)
Cancellaridae
Cancellaria reticulate (L., 1767)
Agatrix srnithi (Dall, 1888)
Aphera lindae Petuch, 1987
Turbinellidae
Turbinella angulata (Lightfoot, 1786)
T. Iaevigata Anton
Vasidae
Vasum muricatum (Born, 1778)
V. capitellum (L., 1758)
Olividae
Oliva oblongs Marrat, 1871
O. bewleyi Marrat, 1871
O. reticularis Lamarck, 1810
O. scripts Lamarck, 1810
O. fulgurator
Roeding,
1798
O. circinata Marrat, 1871
O. reclusa Marrat, 1871
O. goajira Petuch & Sargent,
Olivella olssoni Altena, 1975
O. minuta (Link, 1807)
O. myrmecoon Dall, 1912
O. ankeli Diaz &
Goetting,
O. nivea (Gmelin, 1791)
O. dealbata (Reeve, 1850)
O. lactea (Marrat, 1871)
O. petiolita (Duclos)
O. floralia (Duclos, 1853)
P-G-V
v
R-C-M-V-A
G
R-C-M-G-V-O
M-V
V
R-P-C-M-G
O
R-P-C-M-G-V-A
V-A-LI
M-G-V-A
P-C
R-A-LI
R-P-C-M-V-
V-A
G
A
1987 G
C-M-G-O
R-P-C-M-G-V-A-LI-O
R-P-C-M-G
1989
M-A-LI
P-C-M-V-A-O
LI
C-M-O
P-V
M-A-L1
]aspidella blanesi (Ford, 1898)
J. jaspidea (Gmelin, 1791) P-C-M-V
P-C-V-A-LI
Ancilla glabrata (L., 1758) M-G-V-A
A. bulteata (Sowerby, 1823) A
A. lienardi (Bernardi, 1858) A
A. tankervillei (Swainson, 1825)
V
Agaronia testacea (Lamarck) R-P
Marginellidae
Prunum prunum (Gmelin, 1791) C-M-G-V-LI-O
P. cf. rostratum Redfield M
P. labiatum Kiener P-C
P. apicinum (Menke, 1828) R-A
P. marginatum (Born, 1778) M-G-V-A-LI-O
P. guttatum (Dillwyn, 1817) P-C
Marginella carnea Storer, 1837 P
M. margarita (Kiener, 1834)
LI
M. cloveri Rios & Matthews, 1972 0
Dentimargo aureocincta Stearns, 1872 P-C
D. reducta (Bavay, 1922) C-M-G-V-A
120
J. M. DIAZ
APPENDIX. Continued.
D. sulcata (d'Orbigny, 1842)
D. eburneola (Conrad, 1834)
Marginellopsis serrei Bavay, 1911
Persicula muralis (Hinds, 1844)
P. interruptolineata
(Muhlfeldt,
1816)
P. maculosa (Kiener, 1834)
P. cordorae Jong & Coomans, 1988
P. porcellana (Gmelin, 1791)
P. fluctuata (C. B. Adams, 1850)
P. pulcherrima (Gaskoin, 1849)
P. catenata (Montagu, 1803)
P. chrysomelina (Redfield, 1848)
P. weberi Olsson & McGinty, 1958
Pachybathron tayrona Diaz & Vel., 1987
P. cypraeoides (C. B. Adams, 1845)
Cysticus jansseni Jong & Coomans, 1988
Volvarina avena (Kiener, 1834)
Cypraeolina ovuliformis (d’Orbigny, 1841)
C. antillensis Jong & Coomans, 1988
Mitridae
Mitra nodulosa (Gmelin, 1791)
M. barbadensis (Gmelin, 1791)
M. leonardi Petuch, 1990
Pusiolina veldhoveni Jong & Coomans, 1988
Pusia puella (Reeve, 1845)
P. pulchella (Reeve, 1844)
P. exigua (C. B. Adams, 1845)
P. venusta Sarasua, 1978
P. cubana Aguayo & Rehder, 1936
P. dermestina (Lamarck, 1811)
P. variata (Reeve, 1845)
P. histrio (Reeve, 1844)
P. sykesi (Melvill, 1925)
P. monilifera (C. B. Adams, 1845)
P. hendersoni (Dall, 1927)
P. laterculata (Sowerby, 1874)
P. bibsae (Usticke, 1969)
P. epiphaneum (Rehder, 1943)
Subcancilla leonhardhilli Petuch, 1987
Conomitra lindae Petuch, 1987
C. caribbeana Weisbord, 1929
Turricostellaria lindae Petuch, 1987
T. leonardhilli Petuch, 1987
Nodicostellaria kremerae Petuch, 1987
Conidae
Conus ermineus Born, 1778
C. spurius Gmelin, 1791
C. spurius lorenzianus Dillwyn, 1817
C. mus Hwass, 1792
C. jaspideus Gmelin, 1791
C. puncticulatus Hwass, 1792
C. mappa granarius Kiener, 1848
C. mappa trinitarius Hawass, 1792
C. regius Hwass, 1792
C-A-LI
R
A-LI
V-A-LI
C-M-G-V-A-LI
V-A
LI
C
R-M
M-G-V-A-LI
P-A-LI
A-LI
R-P
M
A-LI
M-G-LI
R-P-C-M-G-V-A-LI
P-C-M-G-V-A-LI
LI
R-P-C-M-V-A
R-P-C-V-A-LI-O
P
M-A-LI
P-M-A-LI
V-A
R-P-M-V-A-LI
M
M
R-P-M-V-A-LI
R-A
P-A-LI
R-A-LI
P-C-M-V-A-LI
O
R-C-M-A-LI
A
R
M-G-V
G
V
G-V
G
G
R-M-G-A-LI-O
P-C-M-G-V-A
R-P-C
R-P-C-M-G-A-LI
P-C-M-G-V-A-LI-O
R-C-M-G-V-A-LI
M-G
V
R-P-C-M-A-LI
SOUTHERN CARIBBEAN GASTROPOD
121
APPENDIX. Continued.
C. daucus Hwass, 1792 R-P-M-A-LI-O
C. penchaszadehi Petuch, 1986 M-G
C. centurio Born, 1778 M-G-A-LI-O
C. brunneofilaris Petuch, 1990 P
C. amphiurgus Dall, 1889 C-M
C. kevani Petuch, 1987
G
C. austini Rehder & Abbott, 1951 M-G-V-O
C. cingulatus Lamarck, 1810 P-C-M
C. forsteri Clench & Aguayo, 1942 V-O
C. bayeri Petuch, 1987 C
C. mindanus Hwass, 1792 P-A-LI
C. granulates L., 1758 R-P-C-LI
C. hieroglyphus Duclos, 1833 A-LI
C. curassaviensis Hwass, 1792 A
C. aurantius Hwass, 1792
LI
C. attenuates Reeve, 1844 LI
C. parascalaris Petuch, 1987
G
C. perprotractus Petuch, 1987
G
Terebridae
Terebra weisbordi Gibson-Smith, 1984
T. trispiralis Weisbord, 1964
T. protexta Conrad
T. salleana (Born, 1778)
T. dislocata (Say, 1822)
T. curacaoensis Jong & Coomans, 1988
T. hastata (Gmelin, 1791)
T. taurina (Lightfoot, 1786)
C-M-G-V
V
P-C-M-G-V-A-LI
R-P-C-M-G-V-A-O
R-P
A-LI
P-C-M-G-V-A-LI-O
P-C-M-G-V-A-LI-O
