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Zoogeography of Marine Gastropod in the Southern Caribbean: A New Look at Provinciality

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Recorded occurrences of 266 species of higher Gastropods were arranged in nine subareas along the coastal areas of southern Central America and northern South America. 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. 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 gastropods in the southern Caribbean. -from Author
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Caribbean Journal of Science,
Vol. 31, No. 1-2, 104-121, 1995
Copyright 1995 College of Arts and Sciences
University of Puerto Rico,
Zoogeography of Marine Gastropod in the Southern
Caribbean: A New Look at Provinciality
Instituto de Investigaciones Marinas de Punta de
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
y el norte de
El valor de las
diversas familias de
como indicadores zoogeograficos y el grado de "singularidad"
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
ampliamente distribuido en el
Occidental tropical y el 39.8% son
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
especies en una determinada
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
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
marines en el sur del Caribe.
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,
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
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.
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-
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
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
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
(Costa Rica)
(Southern Colombia)
M(Santa Marta)
(Leeward Islands)
(Central Venezuela)
Narrow; fringing cora-
Iine archipelago;
mud, sand, gravel
Rather narrow; predom-
inantly mud
Extremely narrow to
absent; steep; mud,
sand, gravel
Rather narrow to wide;
sand, gravel, mud
Narrow; sand, gravel,
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
Agitated to calm; rather
clear to turbid; seasonal-
ly affected by upwelling
Agitated; turbid; somewhat
brackish in the north;
permanently affected by
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
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
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;
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-
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-
Table 2 summarizes the number of gen-
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
18 45
10 33
1 1
3 6
2 3
89 266
36.1 3
26.7 0
22.9 11 15 11 17 18 10 18 24 15
34.6 4
38.5 1
2 2
31.3 19 15 19 14 19 18 18 18 4
36.4 1
20.5 7
10 11
11 0
23.1 4
7 5 10 8 8 9
10 4
44.4 0
0 0
14.7 2
2 2
19.0 1
20.0 1
0 0
26.5 2
21.7 5
712 6
10 5
20.0 5
7 9 10 7
13 14 93
16.7 0
0020 0
25.5 2
3 2
0 2
18.1 6
52 10 2
11 8
9.5 0
20.5 8 10 10 14 12 11 12 66
31.6 2
= 23.8 81
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).
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
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
Area (km)2species average
Costa Rica (R)
Panama (P)
Southern Colombia (C)
Santa Marta (M)
Aruba (A)
Leeward Islands (LI)
Central Venezuela (V)
Surinam-Guiana (0)
120 41.7
124 37.6
* Shelf width estimated from the shoreline to the
100 fathoms depth contour.
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
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.
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-
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;
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-
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-
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;
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;
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.
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
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-
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
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
(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.
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-
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;
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-
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.
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
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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).
Ficus comunis (Roding, 1798)
F. howelli Clench & Aguayo, 1940
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)
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)
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
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
Fulgurofusus brayi (Clench, 1959)
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)
Colubraria lanceolata (Menke, 1828)
C. obscura (Reeve, 1844)
C. swifti (Tryon, 1881)
Melongena melongena L. 1758
Pugilina morio L., 1758
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
Columbella mercatorial
(L., 1758)
Minipirene dormitor (Sowerby, 1844)
Conlla ovulata (Lamarck, 1822)
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
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
Voluta musics L., 1758
V. uirescens (Lightfoot, 1786)
V. demarcoi Olsson, 1965
V. lacertina Petuch, 1990
Scaphella evelina Bayer, 1971
Volutomitra erebus Bayer, 1971
APPENDIX . Continued.
V. persephone Bayer, 1971
Lyria leonardi Emerson, 1985
Morum oniscus (L., 1767)
Cancellomorum lindae (Petuch, 1987)
Cancellaria reticulate (L., 1767)
Agatrix srnithi (Dall, 1888)
Aphera lindae Petuch, 1987
Turbinella angulata (Lightfoot, 1786)
T. Iaevigata Anton
Vasum muricatum (Born, 1778)
V. capitellum (L., 1758)
Oliva oblongs Marrat, 1871
O. bewleyi Marrat, 1871
O. reticularis Lamarck, 1810
O. scripts Lamarck, 1810
O. fulgurator
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 &
O. nivea (Gmelin, 1791)
O. dealbata (Reeve, 1850)
O. lactea (Marrat, 1871)
O. petiolita (Duclos)
O. floralia (Duclos, 1853)
1987 G
]aspidella blanesi (Ford, 1898)
J. jaspidea (Gmelin, 1791) P-C-M-V
Ancilla glabrata (L., 1758) M-G-V-A
A. bulteata (Sowerby, 1823) A
A. lienardi (Bernardi, 1858) A
A. tankervillei (Swainson, 1825)
Agaronia testacea (Lamarck) R-P
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)
M. cloveri Rios & Matthews, 1972 0
Dentimargo aureocincta Stearns, 1872 P-C
D. reducta (Bavay, 1922) C-M-G-V-A
APPENDIX. Continued.
D. sulcata (d'Orbigny, 1842)
D. eburneola (Conrad, 1834)
Marginellopsis serrei Bavay, 1911
Persicula muralis (Hinds, 1844)
P. interruptolineata
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
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
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
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
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
C. attenuates Reeve, 1844 LI
C. parascalaris Petuch, 1987
C. perprotractus Petuch, 1987
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)
... The marine sector of Colombia is in the south region of the Caribbean Sea [33,34] and presents different characteristics on its coasts that allow for the evaluation of different biogeographical hypotheses [17,[34][35][36]. This marine sector has been characterized by various historical events, such as changes in the geomorphology of the coastline due to variations in sea level during the last glaciation and the uplift and movement of Caribbean basin mountain systems, such as the Sierra Nevada de Santa Marta [35]. ...
... A second barrier is believed to be located at 74-71 • W, including Santa Marta and the La Guajira Peninsula [17,35,36], which is attributed to the almost permanent upwelling [55]. The predominant oceanic currents in the area include the CC and the CCC. ...
... Mollusks and fish are the most diverse and abundant species groups in the Colombian Caribbean [36,60] and serve as the biological models for investigating how the Magdalena River plume (MRP) and the combination of the absence of shallow rocky bottoms (ARB) with the almost permanent upwelling in La Guajira (PUG) affect the phylogeography of their populations. For this purpose, we chose three marine species with varying biological and ecological characteristics. ...
Full-text available
The comparative phylogeography of marine species with contrasting dispersal potential across the southern Caribbean Sea was evaluated by the presence of two putative barriers: the Magdalena River plume (MRP) and the combination of the absence of a rocky bottom and the almost permanent upwelling in the La Guajira Peninsula (ARB + PUG). Three species with varying biological and ecological characteristics (i.e., dispersal potentials) that inhabit shallow rocky bottoms were selected: Cittarium pica (PLD < 6 days), Acanthemblemaria rivasi (PLD < 22 days), and Nerita tessellata (PLD > 60 days). We generated a set of SNPs for the three species using the ddRad-seq technique. Samples of each species were collected in five locations from Capurganá to La Guajira. For the first time, evidence of a phylogeographic break caused by the MRP is provided, mainly for A. rivasi (AMOVA: ΦCT = 0.420). The ARB + PUG barrier causes another break for A. rivasi (ΦCT = 0.406) and C. pica (ΦCT = 0.224). Three populations (K = 3) were identified for A. rivasi and C. pica, while N. tessellata presented one population (K = 1). The Mantel correlogram indicated that A. rivasi and C. pica fit the hierarchical population model, and only the A. rivasi and C. pica comparisons showed phylogeographic congruence. Our results demonstrate how the biological traits of these three species and the biogeographic barriers have influenced their phylogeographic structure.
... The marine sector of Colombia is in the southern of the Caribbean Sea [ 25,26 ] and presents different characteristics on its coasts that allow the evaluation of different biogeographical hypotheses [ 17,[26][27][28] ]. This marine sector has been characterized by various historical events, such as changes in the geomorphology of the coastline due to variations in sea level during the last glaciation and the uplift and movement of Caribbean basin mountain systems, such as the Sierra Nevada de Santa Marta [ 27 ]. ...
... A second barrier has been believed to be located in 74 -71°W, including Santa Marta and the La Guajira Peninsula [ 17,27,28 ], which is attributed to the permanent upwelling that transports upwelled water toward the center of the Caribbean Sea and that would limit larval dispersal toward the southwestern Caribbean coast of Colombia [ 48 ]. This putative barrier possibly affects the genetic and phylogeographic structure of species associated with coral reefs and shallow rocky bottoms, whose marine ecosystems are absent for more than 300 km of coastline between Cabo de la Vela (La Guajira) and Tayrona National Natural Park, where upwelling occurs (Figure 1). ...
... Mollusks and fish are the most diverse and abundant species groups in the Colombian Caribbean [ 28,49 ], and serve as the biological models for investigating how the Magdalena River plume (MRP) and the combination of the absence of shallow rocky bottoms (ARB) with permanent upwelling in La Guajira (PUG) affect the phylogeography of their populations. For this purpose, we selected three marine species with different PLDs: the reef fish Acanthemblemaria rivasi (with PLD < 22 days) [ 50,51 ], the exposed rocky shore snails Cittarium pica (with PLD < 6 days) [ 52,53 ] and Nerita tessellata (with PLD > 60 days) [54][55][56]. ...
Full-text available
The comparative phylogeography of marine species with contrasting dispersal potential across the southern Caribbean Sea was evaluated by the presence of two putative barriers: the Magdale-na River plume (MRP) and the combination of the absence of a rocky bottom and permanent upwelling in the La Guajira Peninsula (ARB+PUG). Three species of rocky shallow bottoms were selected with different dispersal potentials: Acanthemblemaria rivasi (PLD < 22 days), Cittarium pica (PLD < 6 days), and Nerita tessellata (PLD > 60 days). We generated a set of SNPs for the three species using the ddRad-seq technique. Samples of each species were collected in five locations from Capurganá to La Guajira. For the first time, evidence of a phylogeographic break caused by MRP is provided, mainly for A. rivasi (AMOVA: ФCT = 0.420). The ARB+PUG barrier causes an-other break for A. rivasi (ФCT = 0.406) and C. pica (ФCT = 0.224). Three populations (K = 3) were identified for A. rivasi and C. pica, while N. tessellata presented one population (K = 1). The Mantel correlogram indicated that A. rivasi and C. pica fit the hierarchical population model, and only the A. rivasi and C. pica comparisons showed phylogeographic congruence. Our results demon-strate how the biological traits of these three species and the biogeographic barriers have influ-enced their phylogeographic structure.
... That author also sustains that the north boundary of the Brazilian Province is the Western Indian Province, near the Venezuela coast. Diaz (1995) and Boschi (2000 a,b), studying gastropods and decapods, respectively, discussed the zoogeographical significance of these groups in the Caribbean and in Americas. Diaz (1995) worked with beach and shelf samples from a wide region between Costa Rica and Suriname, including large islands off Venezuela, such as Aruba, Bonaire, Curaçao and Margarita, as well as smaller ones. ...
... Diaz (1995) and Boschi (2000 a,b), studying gastropods and decapods, respectively, discussed the zoogeographical significance of these groups in the Caribbean and in Americas. Diaz (1995) worked with beach and shelf samples from a wide region between Costa Rica and Suriname, including large islands off Venezuela, such as Aruba, Bonaire, Curaçao and Margarita, as well as smaller ones. He concluded that the region between the Orinoco Delta and Suriname is abruptly impoverished in Caribbean gastropods, with gradual enrichment of Brazilian species eastward and, therefore, being a transitional area toward the Brazilian Province. ...
... It is noteworthy that Briggs (1974), Boltovskoy (1976), Madeira-Falcetta (1977), Forti-Esteves (1984), Diaz (1995), and Boschi (2000a,b) did not sustain the occurrence of a Transitional Zone between 23°S and 15°S, as indicated by ostracods. Whatley et al. (1998b) proposed a provincial model for Argentina, Uruguay, and southern/southeastern Brazil based on the latitudinal distribution of 128 species of benthonic ostracods (mostly in open nomenclature) between 55°S and 20°S. ...
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This study analyzes the geographical distribution of 131 podocopid ostracod species recovered from the Brazilian continental shelf between Cabo de São Roque (lat. 05°30’S) and Cabo Frio (lat. 23oS). This very large area corresponds to the northeastern and eastern Brazilian marine regions. The 104 samples studied were collected in water depths ranging from 12 to 110 m as part of the legs 4 and 7 of the REMAC Project. The cosmopolitan species, as well as those shared with the Caribbean and/or Gulf of Mexico region, represent a small part of the ostracods herein studied and it is assumed that their dispersion was prompted by processes linked to events of relative sea level changes during the Neogene and Quaternary. The fossil record of some species spans to the Neogene, mostly from the Atlantic coast of North and Central America, while one species has Tethyan origin. Three species known from the Neogene of the Caribbean have been recorded as relicts in the study area. From the 131 species herein identified, 36.5% are more widespread in temperate waters south of Cabo Frio town, 46.5% of warm waters north of Cabo Frio town, 4% are present only in the studied area, and 11.5% are rare and probably restricted to the E region. A new province – the Brazilian Province – is herein proposed based on the species occurrence.
... 35) cited as from Barbados seems to be L. ansatus. Díaz (1995) and Capelo and Buitrago (1998) mentioned F. closter from off the Orinoco delta in Guayana; we have not seen specimens from that area, but it is relatively near the confirmed eastern range of L. ansatus. Other reported localities are patently incorrect: Hadorn and Rogers (2000: 11) rightfully questioned specimens labeled "Honduras" and "Yucatán Peninsula, Mexico"; there is no credible evidence that L. ansatus occurs north of continental Colombia in the western Caribbean region. ...
... Altena proposed that the Horst and Schepman shells differed by "their brownish-purple color and details of their form" from those obtained by the Coquette, surmising that the latter "probably belong to one variable species which may be F. closter." Díaz (1995) included "Orinoco delta to Suriname" in the range of F. closter. Hadorn and Rogers (2000) did not distinguish F. ansatus from the later-described F. carvalhoriosi, so their records from the Guianas could represent either taxon; specimens of both species from French Guiana are contained in the Rogers collection. ...
... Aristofusus benjamini seems to be common in deep waters off the Guianas, as indicated by material we examined and by other published records: French Guiana (Hadorn & Rogers 2000;Massemin et al. 2009), Suriname (Okutani in Takeda & Okutani 1983;Hadorn & Rogers 2000), Guyana (Hadorn 1997;Hadorn & Rogers 2000;Mallard 2001;Mallard & Robin 2017), and the Guayana region of Venezuela off the Orinoco Delta (Díaz 1995). ...
The fasciolariid fauna from two expeditions to French Guiana is examined and augmented with published records and material of other collections from the Guianas and northeastern Brazil. Twelve species of Fasciolaria and Aurantilaria (Fasciolariinae), Aristofusus, Lyonsifusus and Fusinus s.l. (Fusininae), and Lamellilatirus and Polygona (Peristerniinae) are reported and discussed. Nine species are represented in expedition collections, and reports of three other species are evaluated. Two morphologically distinct species of Lamellilatirus are described as new; type localities of both are off French Guiana, 114–118 m. Ten Guianan fasciolariids range variously northward to Caribbean South America and the Lesser Antilles and southward to Ceará, Brazil; one other extends into the northern Caribbean, and one extends southward to Rio de Janeiro, Brazil.
... The proposed nomination is located in the southern Caribbean (Fig. 2), an area recognized as a distinct marine biogeographic province for both marine molluscan diversity (Diaz 1995) and fish diversity (Spalding 2007, Robertson & Cramer 2014. The terrestrial habitat includes two plant ecoregions: Venezuelan Mangrove and Aruba-Bonaire-Curaçao cactus shrub (Sotomayor 2003). ...
... With about 650 marine fish species, 745 marine molluscs, 430 marine algae and 201 marine sponges documented for the proposed nomination area, comparative analysis shows that the species richness of these important marine groups surpasses that of other biotically rich areas of the Caribbean (such as Puerto Rico in the case of marine algae) and even the Pacific. For fishes and for molluscs the southern Caribbean has furthermore been demonstrated to be among the top centres for endemism (Diaz 1995, Smith et al. 2002. ...
... The proposed nomination is centrally located in the southern Caribbean, an area recognized as a distinct marine biogeographic province for both marine molluscan diversity (Diaz 1995) and fish diversity (Spalding 2007, Robertson & Cramer 2014. The differences may reflect the fact that these areas represent distinct continental faunas that were separated by the Inter-American Seaway during most of the history of the Caribbean Basin. ...
... This marine ecosystem, discontinuously distributed in Colombia, is absent for more than 300 km of coastline between Cabo de la Vela (Guajira) and TNNP, which probably also influences the genetic and phylogeographic structure of rocky shore species. Mollusks are the most diverse and abundant group in those ecosystems, turning their species into a biological model to investigate how the absence of a given habitat may affect the genetic structure of its populations (Díaz, 1995). ...
... This study demonstrated that C. pica presents high levels of genetic substructuring, exhibiting three well-differentiated populations in the southern Caribbean sector of Colombia. This spatial pattern coincides with three of the five zoogeographic subareas proposed by Díaz (1995), who investigated the biogeography of marine gastropods in the southern Caribbean. These subareas are Goajira (Cabo de la Vela), Samarian (Santa Marta-TNNP) and Isthmian (Cartagena, Isla Fuerte and Capurganá). ...
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It has been proposed that species associated with shallow rocky bottoms in the southern Colombian Caribbean Sea are affected by two biogeographic barriers: the Magdalena River plume (MRP) and the combination of the absence of rocky bottoms and the almost permanent upwelling in the Guajira peninsula (ARB+PUG). We evaluated whether these barriers had generalized genomic effects on the phylogeographic structure of the West Indian top shell (Cittarium pica), a crucial fishery resource in many areas of the Caribbean Sea. Ten species-specific microsatellite loci, 55,112 SNPs from ddRADseq, and COI gene were used on samples of C. pica collected at five Colombian localities (Cabo de la Vela; Santa Marta; Cartagena; Isla Fuerte; and Capurganá). Genetic structure analyses performed for microsatellite and SNP loci indicated that there are three genetic groups of C. pica in Colombia (pop 1: Cabo de la Vela; pop 2: Santa Marta; pop 3: Cartagena+Isla Fuerte+Capurganá). However, both SNPs and COI gene were congruent in showing a phylogeographic break caused by only the ARB + PUG (AMOVA: Φ CT-SNP =0.224, p < 0.05; Φ CT-COI =0.722, p < 0.05), which was confirmed by the maximum-likelihood and network trees. In contrast, MRP was shown to be a permeable barrier to gene flow. Demographic history analysis indicated that C. pica experienced historical changes in population size during the last glaciation period. Cittarium pica is a biological model to demonstrate how the ARB + PUG barrier could affect marine organisms living in shallow rocky habitats, mainly those lacking larvae or having a short-lived larval phase in the southern Caribbean Sea. Finally, some recommendations for fisheries management and conservation C. pica populations are discussed.
... The Caribbean Sea covers an area of about 2,754,000 km 2 and is characterized by a tropical half open sea, belonging to the Atlantic Ocean, bounded to the south of North America, east of the Central America and the north of South America (Diaz 1995). This region has a high marine biodiversity, distributed in many different environments, presenting numerous endemic mollusk species (Diaz 1995;Diaz-Ferguson et al. 2011). ...
... The Caribbean Sea covers an area of about 2,754,000 km 2 and is characterized by a tropical half open sea, belonging to the Atlantic Ocean, bounded to the south of North America, east of the Central America and the north of South America (Diaz 1995). This region has a high marine biodiversity, distributed in many different environments, presenting numerous endemic mollusk species (Diaz 1995;Diaz-Ferguson et al. 2011). ...
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The widespread occurrence of marine gastropods in coastal regions is a straightforward evidence of successful adaptation to different environments. In the Caribbean Sea, as one of Conservation International’s biodiversity hotspots, little is known about the gastropod fauna, especially in the Eastern Caribbean. The present study contributed to bridge this gap by studying the biodiversity of gastropods from Accra Beach, Barbados. Throughout random collections in September 2015, we collected 321 gastropods, comprising eight species, distributed in three families (Neritidae: Nerita tessellata , N. fulgurans , N. versicolor and N. peloronta ; Littorinidae: Echinolittorina ziczac , E. angustior and E. tuberculata ; and Muricidae: Plicopurpura patula ). Nerites were more abundant and diverse, highlighting N. tessellata , representing 66% of the sampled gastropods. This paper also reports the first record of N. fulgurans and E. angustior for the island of Barbados.
... During the Pleistocene, which extended from 2.6 million years to 11,700 years ago, the Caribbean Sea experienced a series of changes in its oceanographic processes due to factors such as sea level variation (i.e., dropping in sea level of about 150 m), volcanic activity and island formation (Ludt and Rocha, 2015). It has long been proposed that the patterns of genetic variation in many species in the Caribbean Sea have been strongly influenced by geologic processes (Avise et al., 1987;Baums et al., 2006;Díaz, 1995;Meschede and Frisch, 1998). Likewise, the present day effect of ocean currents such as the Caribbean current, the Antilles current, and a semi-permanent cyclonic Panama-Colombia Gyre (PCG) located in the southwestern Caribbean have affected the population structure of many invertebrates in the Caribbean Sea Jossart et al., 2017;Lopera et al., 2020;Miloslavich et al., 2010). ...
The Caribbean Sea is characterized by an incredible biodiversity, including several endemic mollusk species, most of which have larval dispersal as their main mechanism of gene flow. It is known that the present-day population structure of a species reflects the combination of oceanographic currents, life-history traits, and historical events. Nerita tessellata is a common gastropod species in coastal, rocky, and exposed shores across the Caribbean Sea; however, little is known about the mechanisms that have shaped its distribution. Here, we tested potential barriers to gene flow and the associated genetic structure of N. tessellata in 12 populations across the Caribbean using digest restriction-site associated DNA sequencing (ddRAD-seq). Likewise, we evaluated demographic history and migration patterns through two coalescence approaches using Approximate Bayesian Computation (ABC) combined with supervised machine learning. The global analysis, based on 26,421 single nucleotide polymorphism (SNP) markers derived from ddRAD-seq, revealed low nucleotide diversity with high levels of gene flow and a lack of significant population differentiation within the survey area, indicating that N. tessellata is panmictic across the Caribbean Sea. Our results reveal that a pattern of north to south movement along Caribbean currents could potentially explain the limited genetic structure and high gene flow among N. tessellata populations. The scenarios simulated with ABC showed a constant effective population size for the ancestral population. Additionally, the four populations tested experienced demographic expansions that date to the Pleistocene. An extended pelagic larval stage could explain the large-scale genetic uniformity of this species across its distribution. Our findings contribute to understanding genetic diversity and population structure in a common gastropod within the Caribbean Sea.
... Other possible causes of connectivity may include similar environmental features shared between Rosario and Tintipán, such as the narrow shelf and predominantly mud bottom, calm and turbid waters influenced by river discharges and similar habitats (e.g. mangroves and coral reefs around offshore islands) (Díaz 1995;Díaz et al. 2000). ...
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The Caribbean king crab Maguimithrax spinosissimus is an important artisanal fisheries resource with a distribution range that includes the Florida Keys and Caribbean Sea islands. We carried out a phylogeographic analysis of M. spinosissimus in one oceanic (Old Providence) and two continental islands (Rosario and Tintipán). We analysed 89 and 49 Control Region (CR) and Cytochrome Oxidase I (COI) sequences, respectively. We found genetic differentiation between Old Providence and Rosario + Tintipán units (FST values of 0.84 for CR and 0.91 for COI gene), and gene flow between Rosario and Tintipán (FST values −0.0085 for CR, P > .88; −0.01 for COI gene, P > .99). Our analysis showed two genetic stocks and possibly an isolated biogeographic unit of M. spinosissimus. We suggest managing the species as different populations, and conducting more ecological and biological studies for the determination of possible cryptic species.
... En el Caribe Colombiano, la malacofauna asociada al manglar ha sido ampliamente estudiada, siendo la ciénaga Grande de Santa Marta y su vecindad una de las áreas más exploradas desde los años setentas (Cosel 1986, Díaz 1994, 1995, Palacio 1978, 1983. La bahía de Cartagena y el golfo de Morrosquillo también han sido históricamente estudiados (revisado por Ortiz & Blanco 2010, Blanco et al. 2010, mientras que el Archipiélago de San Andrés y Providencia ha recibido atención durante la última década (vilardy & Polanía 2002). ...
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Distribution of the mangrove gastropods Neritina virginea (Neritidae) and Littoraria angulifera (Littorinidae) within the Colombian Caribbean Darién Ecoregion. Gastropods are one of the most abundant groups within the Caribbean mangroves, however, little is known about the distribution of particular species at a regional scale. With this aim, we studied the geographic distribution of Littoraria (Littorinopsis) angulifera and Neritina (Vitta) virginea within the Darién Ecoregion in the Caribbean coast of Colombia, from 77 sampling stations along 609km between the Colombian-Panamá border and Córdoba State, Colombia. The fieldwork was conducted in June-August 2009, and a total of 3 963 individuals of both species were hand-picked from the ground, prop-roots and trunks along 50m transects, and shell sizes were measured. The description of geographic patterns considered surface water salinity, mangrove cover and gastropod distribution within the Gulf of Urabá. In the outer-most part of the Gulf, L. angulifera was present in 84.8% of the stations, while N. virginea was only present in 15.2% of the stations. In this part, mangroves areas were patchily distributed, and the gastropods (mainly L. angulifera) were found on woody debris along the supralittoral zone in sandy shores. In the inner-most part, in contrast, N. virginea occurred in 84.6% of the stations, mostly in estuaries, deltas and river margins, while L. angulifera only appeared in Turbo Bay (15.4%). Mean shell size also exhibited a clear geographic pattern: size range was 6-22mm in L. angulifera, and 6-12mm in N. virginea. L. angulifera was found in open-water stations with water salinities >10PSU, but it was absent in sites with lower salinities like the Atrato River Delta and other small rivers. Its presence on coastal woody debris suggests that despite of the recruitment of small individuals from the nearshore stock of larvae, populations are unable to establish due to the absence of mangroves protection. Oppositely, N. virginea was found under estuarine conditions on mangrove roots and ground. Our results confirm that L. angulifera is an esteno-tolerant marine species, and N. virginea is an eury-tolerant estuarine species, thus their geographic distribution is strongly shaped by the large freshwater discharge of the Atrato River. We hypothesize that absence or limited distribution of gastropods in various areas of the Darién Ecoregion may be further explained by the poor conservation state of mangroves.
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Fourteen species and subspecies of Conus from the Santa Marta area, Caribbean Sea of Colombia, are recorded and illustrated. The taxonomic status of some of them are briefly discussed; their habitat preferences (substrates, depths) and theirdistributional patterns inthe western Atlantic are comparatively analyzed.
This book had its origin when, about five years ago, an ecologist (MacArthur) and a taxonomist and zoogeographer (Wilson) began a dialogue about common interests in biogeography. The ideas and the language of the two specialties seemed initially so different as to cast doubt on the usefulness of the endeavor. But we had faith in the ultimate unity of population biology, and this book is the result. Now we both call ourselves biogeographers and are unable to see any real distinction between biogeography and ecology.
Distribution and ecology of the two species of Voluta Linné, 1758, V. virescens and V. música at the Caribbean coast of Colombia are treated. V. virescens occurs between Wounta Haulover, Nicaragua, and Taganga (Santa Marta), Colombia. It inhabits obligately soft bottom and lives in mud, fine sand and coraline sand from shallow water down to a depth of 90 m. V. música occurs in the eastern and southern part of the Caribbean sea. In Colombia it goes as far westward as Ensenada de Chengue, east of Santa Marta. It lives in sand with coraline lime and has not been observed in mud and mineral sand. Its vertical distribution is from shallow water down to about 40 m. Both species live completely buried in the sand and prey on gastropods and other invertebrates, they also eat carrion. The radula of V. virescens is described and figured. The function of a biological barrier east of Santa Marta is discussed, which forms a distribution limit for some mollusks.
Biogeography is currently in an exciting, challenging, revolutionary stage. Quantitative biogeography, both at the historical and ecological levels, will play an increasingly important role because rapidly increasing amounts of data will require quantitative methods and computer analysis. An effective way to comprehend quantitative methods and to determine when they can be effective is to consider any biogeographic study as a multistage decision process. Not only does this provide a framework for such activities, but it also emphasizes that one's conclusions are affected by decisions made at different stages of the study. The best perspective is that biogeography is an unending synthesis, both of many types of theories and of many types of data and analyses to test them. A corollary to this approach is that no single method or analysis can answer all questions of biogeographic interest. Finally, while computers and quantitative analyses can enhance the human mind, they can never replace it.