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The palaeozoogeography of Oligocene to Recent marine Ostracoda from

the Neotropics (mid- and South America) and Antarctica

Article in Marine Micropaleontology · September 1999

DOI: 10.1016/S0377-8398(99)00024-9

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 Marine Micropaleontology 37 (1999) 345–364
www.elsevier.com/locate/marmicro

The palaeozoogeography of Oligocene to Recent marine Ostracoda

from the Neotropics (mid- and South America) and Antarctica
Adrian M. Wood a,Ł , Maria Inês F. Ramos b , Robin C. Whatley c
a
Centre for Quaternary Studies, School of Natural and Environmental Sciences, Coventry University,
Priory Street, Coventry, CV1 5FB, UK
b Instituto de Geociências, UFRGS, Caixa Postal 15001, CEP 91501-970, Porto Alegre, RS, Brazil
c Micropalaeontology Research Group, Institute of Geography and Earth Sciences, University of Wales, Aberystwyth SY23 3DB, UK

Received 31 July 1997; accepted 11 February 1999

Abstract

Classic biogeographical research has shown that the continent of South America supports a diverse and priceless biota,
of which ostracods are an important component. The distribution patterns of Oligocene to Recent shelf ostracods, from
the Neotropics to Antarctica, are explained in terms of dispersal and vicariant events. The quantitative examination of a
newly constructed database, containing over 140 genera, has allowed the measurement of generic similarity and endemicity
between biotas of different geographical regions. The measurement of these parameters has aided the construction of a
series of palaeoendemicity and communality maps. These maps emphasise changes in the spatio-temporal distribution of
mid to Late Tertiary ostracods, and can aid in the recognition of abiotic mechanisms that modify genera distribution. It has
been demonstrated that changes in the oceanic currents and water-mass temperature are significant in the formation and
maintenance of zoogeographical domains in the Oligocene–Recent of the Neotropics and Antarctica. South America was
an important centre of origin for ostracods during the Oligocene, however, few genera appear able to disperse northwards
towards the Caribbean. The migratory success or failure of benthonic ostracods is closely linked to oceanographical and
climatic conditions, and their physiology. Within the Meso-American region, filter and corridor pathways have allowed
rapid dispersal of shallow water ostracods which has lead to decreased endemism. Although a distinctive ostracod
assemblage was established in the Oligocene of Antarctica, the expansion of the Drake Passage permitted a new suite of
cryophilic genera to emerge on the continent during the ?Mio-Pliocene. Within the Meso-American region the alteration of
oceanic circulation patterns, subsequent to the closure of the Panamanian portal, may have initiated the development of a
‘proto’ Panamanian Province in the Early Pliocene.  1999 Elsevier Science B.V. All rights reserved.

Keywords: Ostracoda; palaeozoogeography; Tertiary; Recent; Neotropics; Antarctica

1. Introduction importance as an issue of both scientific and political

concern (Wilson, 1997). It was appropriate that the
The science of biogeography, and especially the first Environmental Summit Meeting was held in
related discipline of biodiversity, have gained global Rio de Janeiro, Brazil for here in South America
an unparalleled and priceless biota now exists which
Ł Corresponding author. owes its diversity to ancient tectonic events and

0377-8398/99/$ – see front matter  1999 Elsevier Science B.V. All rights reserved.
PII: S 0 3 7 7 - 8 3 9 8 ( 9 9 ) 0 0 0 2 4 - 9
346 A.M. Wood et al. / Marine Micropaleontology 37 (1999) 345–364
biological processes such as speciation, dispersal
and extinction.
Within the Neotropics, the majority of recent
biogeographical articles have focused on the Neo-
gene of Meso-America, and the consequences of an
emergent Central American Isthmus (Savage, 1982;
Stehli and Webb, 1985; Woods, 1989; Jackson et
al., 1996). Many of the resulting theories of species
dispersal were based primarily upon the analysis
of terrestrial organisms (Raven and Axelrod, 1974;
Rosen, 1976; Gomez, 1982; Savage, 1982; Good-
friend, 1989; Miller and Miller, 1989). Although
exceptions exist (Petuch, 1988; Collins, 1996), many
of the basic patterns in the marine realm have yet
to be described or discovered (Jackson and Budd,
1996).
The central aim of this paper is to describe, in
terms of dispersal events, key patterns in the distri-
bution of Oligocene–Recent South American marine
Ostracoda. The disposition and quality of sample
sites are never perfect, however, the presently as-
sembled database of over sixty Recent and fossil
locations is unequalled. The identification of over
140 ostracod genera has enabled the present authors
to identify dispersal events within two major palaeo-
migratory pathways. One path lies to the south be-
tween Antarctica and southern Argentina, and the
second, the Central American Isthmus, to the north.
By measuring the level of generic communality and
endemicity across these pathways, we are able to
measure changes in the spatio-temporal distribution
of ostracods, and allude to the (a)biotic mechanisms
that have modified them. A list of primary data
sources and genera presence–absence data can be
found in Appendixes A and B 1 .
2. The conflicting philosophies of ‘dispersal’
biogeography
The study of species distribution patterns has
led to the development of two schools of thought
which emphasise divergent processes and philoso-
phies (Nelson and Platnick, 1984; Hallam, 1988).
1Appendix B can be found at http:==www.elsevier.nl=locate=
marmicro or http:==www.elsevier.com=locate=marmicro.
2.1. Vicariant events
Vicariant species are closely related species that
occupy different geographical or ecological areas.
However, the term vicariant biogeography has de-
veloped a more specialised meaning than originally
intended by Croizat (1958). Accepting that dispersal
can take place, vicariant biogeographers believe that
the successful crossing of barriers is a rare event and
that most species have evolved in situ rather than as
a consequence of dispersal (Rosen, 1976).
Leon Croizat (1952) noted that whereas a disper-
sal event would affect only a single species, vicariant
species should affect groups of different taxa which
had a similar disjunct distribution pattern. The in-
terpretations of vicariant biogeography have been
enormously strengthened by the widespread accep-
tance of continental drift and the theory of plate
tectonics.
Throughout the Phanerozoic Eon, numerous vi-
cariant and combinatory biotas have been recog-
nised using ostracods. In “Ostracoda and Continental
Palaeogeography” Whatley (1988, p. 109) has intro-
duced a number of classic examples.
2.2. Dispersal events (sensu stricto)
The central hypothesis of dispersal biogeography
is that species originate in a particular area or cen-
tre and, if successful, spread out from that area to
colonise new habitats. Charles Darwin (1859) was
first to write about such centres which he called
“single centres of creation”. Many of the arguments
associated with centres of origin have focused on
the marine East Indies, and the East Indian Trian-
gle which is thought to exhibit the greatest species
diversity in the marine world (Briggs, 1984, 1992;
Whatley, 1987; Witte, 1993).
3. Tectonic history of the southern continents
The tectonic history of South America and es-
pecially the Caribbean is extremely complex (Bar-
ron, 1987; Perfit and Williams, 1989; Coates and
Obando, 1996). Since the initial fragmentation of
the once unified super-continent of Gondwanaland in
the Early Cretaceous, these southern continents have
 A.M. Wood et al. / Marine Micropaleontology 37 (1999) 345–364
undergone numerous, and geologically rapid vicari-
ant and convergent events. There are many problems
concerning the mutual positions of Africa, South
America and Antarctica during the Tertiary. How-
ever, based upon palaeontological and structural evi-
dence the most reliable reconstruction of continental
outlines for the Cretaceous and Tertiary periods can
be found in Tarling (1972, 1980), Smith and Briden
(1977), and Golonka et al. (1995).
A detailed review of continental migration in the
Southern Hemisphere is beyond the scope of this
paper. However, a short résumé of major tectonic
episodes, and a simplistic reconstruction of palaeo-
continental movements, based upon geological evi-
dence have been included (Fig. 1):
Mesozoic Andean Orogeny and the development
of subduction zones to the southwest
and west of South America.
Eocene Subduction to the west and within the
Caribbean region itself, the latter lead-
ing to the formation of the Lesser and
Greater Antilles.
?Oligo-Miocene Separation of Antarctica from South
America, and the subsequent develop-
ment of the Drake Passage.
Mio-Pliocene Northward migration of South America,
localised shoaling of the Caribbean Sea.
Formation of an extended Central Amer-
ican archipelago then Isthmus.
4. Previous ostracod research
Ostracod research of the South Atlantic Ocean
was initiated by Brady (1870, 1880, 1907) in the lat-
ter half of the Nineteenth Century, but only recently
have these founding taxonomic masterpieces been
augmented. A preponderance of ostracodological re-
search has focused on older Tethyan migration routes
(Dingle, 1988; Whatley, 1988; Whatley and Ballent,
1994; Boomer and Ballent, 1996), the juxtaposition
of southern continents (Whatley, 1988; Reyment and
Aranki, 1991), and mid-Tertiary faunas of Meso-
America (Van den Bold, 1957, 1958, 1963a,b, 1964,
1965, 1966a,b,c,d,e, 1968a,b, 1969, 1970, 1972a,b,
1973, 1974, 1975, 1976, 1977, 1983, 1985, 1988).
It has also been established that during the early
Tertiary, while the South Atlantic was half its present
347
width, ostracods were still able to migrate between
South America and Africa (Reyment and Aranki,
1991). As a consequence, the Recent Ostracoda of
West Africa and South America still retain many
generic legacies of this early Tertairy union (Witte,
1993; Wood and Whatley, 1994).
Much of the research into mid-Tertiary to Re-
cent Meso-American, South American and Antarc-
tic shelf Ostracoda have tended to be descriptive
in nature, as workers strive to fill the taxonomic
void. Subsequent ostracod research is many tiered
with several operational scales of inquiry (Sepkoski,
1988): alpha, the analysis of taxa at a single lo-
cality; beta, differentiation of taxa between sites;
and gamma, the taxonomic differentiation between
geographical regions. In these terms the majority
of work has been undertaken at the alpha scale,
focusing on species (palaeo)ecology, bathymetry,
intra-regional biostratigraphy or localised eustasy.
Very little has been attempted at the larger beta or
gamma scales, although exceptions do exist for the
Caribbean (Van den Bold, 1977), Argentina (What-
ley et al., 1998a,b), Chile (Hartmann-Schröder and
Hartmann, 1962; Hartmann, 1966), Brazil (Pinto and
Ornellas, 1970, 1978) and Antarctica (summarised in
Hartmann, 1997).
4.1. Oligocene
The most note-worthy palaeobiogeographical
study of Cainozoic ostracods of Central–South
America was made by Van den Bold (1977), which
described faunal provinces and dispersal mecha-
nisms for a number of selected genera (Van den
Bold, 1970, 1974). Similarly, the distribution of
hemicytherid genera in the southern hemisphere has
been outlined by Valicenti (1977).
A register of Argentinean Palaeogene ostracods
has recently been compiled by Echevarrı́a (1995),
while additional references can be found in Ber-
tels (1975) and Kielbowicz (1988). Ostracods of
Oligocene age have only been described from one
Antarctic site on King George Island (Blaszyk,
1987). This fauna was thought by Blaszyk to re-
semble (in part) assemblages obtained from Isla
Grande de Tierra del Fuego, in southern Argentina
(Echevarrı́a, 1982, 1987).
348 A.M. Wood et al. / Marine Micropaleontology 37 (1999) 345–364

Greater Antilles Lesser Antilles

Pliocene
Antarctica Australia
(3mya)
North America South America

Greater Antilles Lesser Antilles

Late Miocene Antarctica Australia
(6mya)
North America South America

CONVERGENCE

Greater Lesser
Antilles Antilles
Early Miocene
Antarctica Australia
(20mya)
North America South America

VICARIANCE

Greater Lesser
Antilles Antilles
Eocene/Oligocene Australia
(35mya) North South America
America Antarctica

VICARIANCE

Paleocene North South America/

(60mya) America Antarctica/Australia

Fig. 1. Simplistic synoptic maps of relative (vicariant and convergent) continental movements in the Southern Hemisphere during the
Tertiary, based upon geological evidence.

4.2. Miocene–Pliocene able post-Eocene biostratigraphical zonation scheme

(Van den Bold, 1983) based upon rapid evolutionary
Much of Van den Bold’s earliest work on Caribbe- (i.e., ‘Radimella’) events (Van den Bold, 1988). In
an ostracods was taxonomic but also included a reli- Brazil and Argentina nearly all research has been
 A.M. Wood et al. / Marine Micropaleontology 37 (1999) 345–364
essentially descriptive and=or has focused on the
stratigraphical application of ostracods (Echevarrı́a,
1987; Sanguinetti et al., 1991, 1992; Carreño et
al., 1999). Von Ohmert (1968, 1971) published two
important papers on the Pliocene to Recent radi-
ation of the genus Caudites, and the endemicity
of the subfamily Coquimbinae in Chile. More re-
cently, Cronin and Dowsett (1996) have used R- and
Q-mode cluster analyses in order to identify changes
in ostracod communities and palaeoceanography in
the Caribbean.
4.3. Pleistocene
The few papers on the subject of Pleistocene Os-
tracoda have dealt mainly with relationship between
faunal associations and sea-level (Bertels, 1975; Vi-
calvi et al., 1977; Bertels et al., 1982; Bertels and
Martinez, 1990; Aguirre and Whatley, 1995).
4.4. Recent
Research into Recent ostracods has focused upon
three geographical regions:
4.4.1. Meso-America
Numerous Recent samples were collected and
analyzed by Van den Bold (1964, 1972b, 1975) dur-
ing his exploration of Neogene successions. These
Recent data confirmed the presence of two zoogeo-
graphical (sub)provinces, the Gulf of Mexico and
the Caribbean (Van den Bold, 1977). While working
on sub-Recent samples from Belize, Teeter (1975)
recognised similarities in the generic associations of
the Recent and palaeo-Caribbean.
4.4.2. Argentina, Brazil and Chile
A number of small-scale, sub-provincial ostra-
cod communities has been recognised in northeast-
ern Brazil which contain species common to the
Lesser Antilles and Central America (Coimbra et al.,
1992). However, the Recent ostracod biotas along the
Pacific (Hartmann-Schröder and Hartmann, 1962,
1965) and Atlantic (Whatley et al., 1998c) coasts
of South America have been segregated into large
geographically restricted areas or provinces. The
subdivision of southwestern Atlantic ostracod as-
semblages into zoogeographical provinces is as a
349
result of a succession of publications by Whatley et
al. (1987, 1988, 1995, 1996, 1998a,b).
4.4.3. Antarctica
Before 1964 Antarctic ostracod research was
dominated by taxonomy. A brief history of research
has recently been presented by Hartmann (1997), and
only one publication (Whatley et al., 1999) postdates
this. In a short zoogeographical summary, Hartmann
(1997) suggested that the faunas from the Antarctic
were littoral Tertiary relics. The indigenous charac-
ter, and Recent distribution pattern of southern ocean
species were thought by Benson (1964) and Hazel
(1967) to be a product of climate and topography.
5. Methodology: binary similarity coefficients
and endemicity as aids in panbiogeographical
study
Panbiogeography is a philosophy that focuses
upon the measurement of communality (or similar-
ity) between biotas of different geographical regions
(Myers and Giller, 1994). Two contrasting measures,
binary similarity coefficients and percentage generic
endemicity, have been used to study the spatio-tem-
poral evolution of Meso-American, South American
and Antarctic ostracods.
5.1. Binary similarity coefficients
Binary similarity coefficients are essentially used
to measure beta diversity or the degree of association
or similarity between paired sample sites (Whittaker,
1977; Wilson and Shmida, 1984; Magurran, 1988;
Shi, 1993; Hallam, 1994; Rosenzweig, 1995).
A large choice of binary measures are available,
although the Jaccard and Sorenson indices are most
commonly used (Hazel, 1970; Janson and Vegelius,
1981; Neil, 1995). In the field of ostracodology, the
Simpson and Sanders coefficients were employed
by Van den Bold (1966a, 1972a) and Neil (1995)
respectively, as an aid to correlation. The simplicity
of such coefficients are their greatest advantage,
however, one must be reminded that all groups count
equally irrespective of their abundance. The Jaccard
binary measure was chosen for the current study:
C j D j=.a C b j/ (1)
350 A.M. Wood et al. / Marine Micropaleontology 37 (1999) 345–364
where: j D the number of species common to both
faunas; a D the number of species in fauna A, and b
D the number of species in fauna B.
The calculated levels of intra and inter-regional
generic similarity in the Oligocene, Early Miocene,
Late Miocene and Pliocene are presented on a num-
ber of palaeogeographical maps (Fig. 2a–d; Table 2).
Complete similarity matrices for fossil sites are pre-
sented in Appendix C 2 .
5.2. Endemicity
Stenotopic or endemic biota can be broadly di-
vided into two groups: neoendemics and palaeoen-
demics (Engler, 1882). Neoendemics are confined
in their distribution to the areas in which they
evolved, whereas palaeoendemics are relicts, iso-
lated geographically by extinction. However, both
types are influenced by contemporaneous ecologi-
cal factors and=or historical, and large-scale abiotic
processes. The study of endemics has proved to be
very useful, revolutionising biogeography with the
introduction of new methodologies (Humphries and
Parenti, 1986) and measures (Bykov, 1979). Most re-
cently, Boomer and Whatley (1996), Larwood et al.
(1996) and Larwood and Whatley (1993) have high-
lighted that prolonged spatial isolation is essential to
the generation of endemic ostracods, therefore, by
measuring the latter one can allude to the former.
Endemicity is measured as a percentage, where gt is
the total number of genera in a given region, and ge
the number of genera restricted to that region:
ge
Percentage endemism D t ð 100
g America (3.4%). These regions were considered to
Where data allows, the percentage of generic
endemicity has been calculated for Meso-America,
South America and Antarctica. Meso-America is
considered by the present authors to include the
Lesser Antilles, Greater Antilles, Central American
region between 23 and 8ºN, Colombia and Vene-
zuela. As with communality, the regional endemicity
of Neotropical to Antarctic ostracods is also pre-
sented on a series of reconstructed maps (Fig. 2a–d).
A complete list of endemic genera is also given in
Table 1.
2Appendix C can be found at http:==www.elsevier.nl=locate=
marmicro or http:==www.elsevier.com=locate=marmicro.
The existence of amphi-atlantic taxa, our inade-
quate knowledge of late Tertiary African faunas, and
dubious taxonomy has made the identification of a
small number of endemic genera presently impossi-
ble.
6. Results and discussion
6.1. Oligocene
6.1.1. Palaeogeography and oceanography
A detailed account of South American palaeo-
(bio)geography can be found in Hallam (1994) and
Jackson et al. (1996). At the end of the Eocene the
Central American Isthmus had formed a continuous,
but submerged structural unit. A number of large sea-
ways still existed between North and South America
(Pindell et al., 1988). To the south, the development
of the Drake Passage may well have been initiated
during the Oligocene (Hallam, 1994), although an
epicontinental sea still existed between Patagonia
and the West Antarctic Peninsula. The anticyclonic
gyre of surface currents in the South Atlantic was
much larger allowing the warmer Brazilian current
to extend farther south.
6.1.2. Ostracod endemicity and communality
(Fig. 2a)
Due to incomplete Oligocene coverage, percent-
age generic endemicity has only been calculated for
three regions, the Austral Basin of South America
(18%), Western Antarctic Peninsula (0%) and Meso-
be important migratory pathways during the Tertiary
(Simpson, 1940), however, the especially high levels
of ostracod endemicity is indicative of low levels of
faunal interchange between the Austral Basin and the
Western Antarctic Peninsula. As a comparison, Re-
cent ostracod endemicity values in equatorial West
Africa do not exceed 4.3%, while values for south-
ern and northern Europe are almost zero (Wood and
Whatley, 1994).
A significant number of new ostracod genera first
appeared in the Austral Basin of southern Argentina
during the Oligocene, making this an important cen-
tre of origin. Such levels of generic origination are
undoubtedly linked to a number of mutually in-
 A.M. Wood et al. / Marine Micropaleontology 37 (1999) 345–364 351

no endemics
0.28 0.33
1.7% 0.36 0.07
0.41 0.44 0.54
0.34

0.21
no data
0.15

11%
0.26
18%
0.25
0.11

0%

a OLIGOCENE b EARLY MIOCENE

0.42
2.6% 0.29 0.36 1.2% 0.39
0.28
0.57

0.14
0.09

11% 0.27
0.5 11.7%

0.08

9%

c LATE MIOCENE d PLIOCENE

Fig. 2. Palaeogeographical reconstruction of the South American region during the Oligocene, Early Miocene, Late Miocene and
Pliocene, including percentage ostracod endemicity values for critical regions and mean genera communality (Jaccard Coefficient) values
for selected Neotropical to Antarctic sites. Mean Jaccard Coefficient values indicate similarity between adjacent localities in each N–S
transect.
352 A.M. Wood et al. / Marine Micropaleontology 37 (1999) 345–364

Table 1
Oligocene to Recent endemic genera from Meso-America, South America and Antarctica

Oligocene endemics Early Miocene endemics Late Miocene endemics

Meso-America South America Meso-America South America Meso-America South America Antarctica
Orionina A. (Ambostracon) no endemics A. (Patagonacythere) Caribella Argenticytheretta no data
A. (Patagonacythere) Argenticytheretta Pseudoceratina Australicythere
Argenticytheretta Brasilicythere Bensonia
Australicythere Papillosacythere a Brasilicythere
Australicytheridea Australicythere a Australicytheridea a
Bensonia Soudanella a Papillosacythere a
Brasilicythere Bensonia a Soudanella a
Coquimba Australicytheridea a
Papillosacythere
Soudanella
Antarctica
Antarctica no data
no endemics
Pliocene endemics Pleistocene–Recent endemics
Meso-America South America Antarctica Meso-America North Brazil South Brazil=Argentina Antarctica
Pseudoceratina Papillosacythere Meridionalicythere Caribella Tanella Argenticytheretta Antarcticythere
Soundanella Ruessicythere Whatleyella Australicytheridea Austrocythere
Brasilicythere a Austroaurila Macroscapha
Australicytheridea a Brasilicythere Pelecocythere
Argenticytheretta a Falklandia Pontocypria
Papillosacythere new genus B
new genus A
a Lazarus genera which have not been included in the calculation of percentage endemicity.

clusive factors including available area, relative sea
level and habitat patchiness. The continental shelf
of southeastern South America covers a huge area
of some 1 million km2 . The greatest opportunity for
speciation was probably associated with the subdi-
vision of this epicontinental sea (and the nonplank-
totrophic ostracod population) by changing sea levels
during the Early Oligocene (Valentine and Jablonski,
1983; Prothero and Berggren, 1992).
Nine neo-endemic (evolved in situ) ostracod gen-
era and subgenera have been identified from south-
ern Argentina. Only Soudanella Apostolescu can be
considered a palaeo-endemic. A complete listing of
endemics is given in Table 1. Soudanella is an inter-
esting genus for it remains the only palaeo-endemic
recorded from the upper Tertiary of South Amer-
ica. This genus was originally described from the
Palaeocene of Senegal, however, it is now known
to have occurred in the Palaeogene of the Middle
East (Bassiouni, 1969), North Africa (Reyment and
Reyment, 1981), Caribbean (Van den Bold, 1975)
and South America (Bertels, 1969, 1975; Neufville,
1979). However, since the Eocene its distribution
has contracted southwestwards to southern Argentina
(Bertels, 1975; Echevarrı́a, 1995) where it remained
as a palaeo-endemic until its final extinction in
the Pliocene (Echevarrı́a, 1988). The accounts of
Soudanella from the Neogene of West Africa are
erroneous. This genus appears to have been confused
with both Ruggieria and Keijella (Carbonnel, 1992).
Only one neo-endemic, Orionina has been de-
scribed from the Oligocene of Meso-America (Van
den Bold, 1963a, 1965). Its status as an endemic was
short lived for rapid migration, via shoals in the An-
tilles and along the coast of the emerging Central
American Isthmus, enabled it to colonise the southern
coasts of North America in the ?Early to mid-Miocene
(Swain, 1951). However, McKenzie (1987) suggests
that as few as 5% of species occurred in both the
Caribbean and Gulf Coast in the Early Oligocene.
Similar filter routes were envisaged by Van den
Bold (1974) to account for the northward migration
 A.M. Wood et al. / Marine Micropaleontology 37 (1999) 345–364
of shallow water species of Cativella, Pellucistoma
and Costa from the Oligocene of Venezuela and
Colombia. Van den Bold also revealed that the dis-
persal rates of ostracod genera were quite variable
(Van den Bold, 1974, fig. 3). Puriana was considered
to be a neo-endemic of the Meso-American region,
however, it appears to have first appeared simultane-
ously in the Oligocene of Puerto Rico (Van den Bold,
1965), the Lesser Antilles (Van den Bold, 1966c) and
Southern Mississippi (Hazel et al., 1980).
Mean intra-regional similarity values of between
0.28 and 0.41 for generic associations on island
sites within the Greater and Lesser Antilles, would
also support the idea of unrestrained interchange.
However, these figures contrast considerably with
low inter-regional values between north and south.
Although a passive transport agent existed in the
form of the warm Brazilian Current, the average
similarity value of 0.14 would indicate restricted dis-
persal along the southwestern Atlantic shelf during
the Oligocene. Paradoxically, the absence of a cold
Falklands Current, and a less formidable temperature
gradient, should have encouraged ostracod dispersal
along the shelf of southern Brazil.
The King George Island assemblage of Antarctica
(Blaszyk, 1987) has a low diversity and no endemics;
the assumption of remoteness is also supported by a
low communality score (0.15).
6.2. Early Miocene
6.2.1. Palaeogeography and oceanography
The severing of Antarctica from Patagonia, and
the establishment of a sea-way between Australia
and Antarctica lead to the development of the cir-
cum-Antarctic oceanic circulation system, and its
northerly branch the Falklands (Malvinas) Current.
The precise timing of this event is difficult to pin-
point, however, Hallam (1994) suggests the Early
Oligocene. Additional evidence from planktonic fo-
raminiferal assemblages in the South Atlantic in-
dicate ocean cooling across the Oligocene=Mio-
cene boundary (Spezzaferri, 1995), in the Middle
Miocene (Flower and Kennett, 1994), and latest
Miocene (Boltovskoy, 1979, 1980). These Neo-
gene events may denote major changes in deep
ocean circulation related to the gradual development
of the circum-Antarctic oceanic circulation system
353
and East Antarctic Ice Sheet (Frankes et al., 1992).
The closure of the E–W-trending deep water con-
nection occurred in the Caribbean in association with
a regional shift from deep to mid bathyal-neritic con-
ditions (Coates and Obando, 1996).
6.2.2. Ostracod endemicity and communality
(Fig. 2b)
Ostracod endemicity values for the Austral Basin
of South America are complicated by the presence
of Lazarus genera: taxa which seem to suffer ex-
tinction but then reappear later in the stratigraphical
record (Jablonski, 1986). Neogene neo-endemics,
including Papillosacythere, Australicythere, Benso-
nia, Australicytheridea, Brasilicythere and Argenti-
cytheretta Rose, 1975, appear to become extinct only
to re-emerge from the dead (see Table 1). These
genera undoubtedly existed within this region but are
as yet undiscovered. If Lazarus endemics are not in-
cluded in the calculation, Early Miocene endemicity
appears to decrease to 11%.
However, the eurythermal genus Coquimba does
appear to be an authentic escapee from the confine-
ments of the Austral Basin, having being recorded
from the Miocene of the Caribbean (Van den Bold,
1972a, 1973). In order to eliminate the need for pos-
tulating a single, epic, migratory event of some 3500
miles, we speculate that intermediate stations may
have existed in Brazil.
No endemic ostracod genera were recorded from
Meso-America. The relative ease of genera redis-
tribution in the Caribbean appears to confirm a re-
gional shift to shallower oceanic conditions and the
continued shoaling of the Antilles. Cronin (1987)
considered dispersal among Caribbean islands was
passive, and not directly related to specific abiotic
events. Indeed, the colonisation of shallow water
habitats within the Caribbean could easily have been
achieved via natural rafts of drifting debris.
A number of low similarity values has been
recorded between faunas of the Lesser and Greater
Antilles (0.07 between Trinidad and Puerto Rico).
These low figures have arisen because one is com-
paring bathymetrically divergent faunas, bathyal ver-
sus neritic. The former depth zone is characterised
throughout the Caribbean by the genera Cardobair-
dia, Krithe, Ambocythere, Bradleya and Henry-
howella (Van den Bold, 1963b, 1969).
354 A.M. Wood et al. / Marine Micropaleontology 37 (1999) 345–364
Most significantly the communality between
southern Brazil and the Caribbean reaches its acme
within the Miocene with a similarity value of 0.21.
An important fauna containing the genera Bair-
doppilata, Pellucistoma and the species Orionina
vaughani (Ulrich and Bassler), were described from
the Pelotas Basin (32ºS) by Sanguinetti (1979). In
the Recent, Orionina vaughani occurs as far south
as Espirito Santo State (20ºS). The southward dis-
placement of ‘equatorial’ genera into southern Brazil
during the Early Miocene was probably linked to tec-
tonism in the south, and the gradual development of
the circum-Antarctic oceanic circulation system. The
subsequent decoupling of the South Atlantic gyre
from Antarctic waters resulted in the intensification
of the warm Brazilian Current (Kennett, 1980).
6.3. Late Miocene
6.3.1. Palaeogeography and oceanography
Partial uplift of the submerged Central Ameri-
can Isthmus occurred with a resultant disruption of
the California Current (Duque-Caro, 1990; Coates
and Obando, 1996). Major tectonic activity in the
Caribbean Sea also resulted in the rapid uplift and
subsidence of parts of the ocean floor (Van den Bold,
1968b, 1988; Steineck et al., 1984).
6.3.2. Ostracod endemicity and communality
(Fig. 2c)
Both endemicity and communality parallel the
Oligocene, although two neo-endemics, Caribella
and Pseudoceratina are recorded from Meso-Amer-
ica. High similarity values (0.5) were obtained
(as expected) between sites from Venezuela and
Trinidad, but as in the Oligocene, the similarity
between genera of the north and south remains low
(0.14). Only common cosmopolites such as Aurila,
Bradleya, Cytherella, Krithe and Xestoleberis occur
in both regions. Sadly, no ostracods have yet been
described from the Miocene of Antarctica.
6.4. Pliocene
6.4.1. Palaeogeography and oceanography
The Central American Isthmus emerged in the
Late Pliocene (Jackson et al., 1996) in association
with renewed uplift of the Andes.
6.4.2. Ostracod endemicity and communality
(Fig. 2d)
As the Central American Isthmus emerged in the
Late Pliocene a ‘corridor’ type pathway was estab-
lished which aided the migration of benthonic ostra-
cods. The result was a further decline in Caribbean
endemism to only 1.2%. With mean similarity values
of between 0.36 and 0.42, intra-regional commu-
nality remained high within the Caribbean Sea, and
between the Caribbean and the southern part of the
Pacific coast of North America. Before the forma-
tion of the isthmus two Oligocene endemics of Ar-
gentina, Ambostracon (Ambostracon) and Coquimba
were able to enter the Gulf of California (Valentine,
1976; Carreño, 1985), whether this was achieved
via a Central American portal or the west coast of
South America remains unclear. One younger en-
demic, Pseudoceratina, which emerged in the Late
Miocene remains confined to the Caribbean.
It is likely that the distinctive species character
of the Pacific coast and Caribbean–Gulf Coast ostra-
cod communities was established before the Early–
mid-Pliocene (Carreño, 1985). However, the isola-
tion of ostracod populations by the Central Amer-
ican Isthmus appears not to have aided speciation
among pre-isthmus species of the genera Puriana
(Cronin, 1987) or Orionina (Gunther and Swain,
1976; Cronin and Schmidt, 1988).
The percentage of endemics remained high in
South America (11.7%) as the southward migration
of Antarctica lead to the expansion of the Drake Pas-
sage. The computed values for endemicity (9%) and
communality (0.08) of the Cockburn Island assem-
blage in Antarctica (Szczechura and Blaszyk, 1996)
affirm this progressive disconnection.
6.5. Recent
6.5.1. Ostracod endemicity and communality
(Fig. 3; Table 2)
A recent succession of geographically diverse
publications has improved our basic knowledge of
ostracod endemicity and communality in the south-
western Atlantic (Fig. 3; Table 2; Whatley et al.,
1987, 1988, 1995, 1996, 1998a,b). The communal-
ity values are presented in a similarity matrix that
is subdivided on the basis of mean regional sim-
ilarities of approximately 0.3 (see inset, Table 2).
Table 2
Jaccard Coefficient similarity matrix for Pleistocene and Recent sample sites. On the basis of regional trends in similarity the Caribbean, Brazilian, Subantarctic (in part)
and Antarctic provinces are recognised (Whatley et al., 1998c). Mean inter-regional similarities values have also been supplied (see inset). Primary data sources are given in
Appendix A

A.M. Wood et al. / Marine Micropaleontology 37 (1999) 345–364

355
356 A.M. Wood et al. / Marine Micropaleontology 37 (1999) 345–364
?2.4%
4%
8.8%
9.6%
RECENT
Fig. 3. Recent regionalised endemicity values of ostracod shelf
genera from the southwestern Atlantic sea board.
Four major zoogeographical regions are recognised:
the Caribbean, Brazilian, Subantarctic and Antarctic.
The spatial extent of these regions appear to parallel
those recently described by Whatley et al. (1998c).
The only deviation from their scheme is the apparent
absence of their Bonaerensian Province (43–36ºS).
It is certain that differences in the scale of sampling
and taxonomic investigation has caused this transi-
tional (ecotonal) province to be subsumed within the
Brazilian and Subantarctic provinces.
At 8%, levels of endemicity remain high in
Argentina=southern Brazil, while the continued iso-
lation of Antarctica appears to have assisted en-
demicity where values rise to 9.6%. However, dis-
tance alone would not suffice in maintaining iso-
lation and, therefore, endemicity. As within the
northeastern Atlantic system (Wood and Whatley,
1994), oceanic structures such as the circum-Antarc-
tic oceanic circulation system, Subantarctic and
Antarctic fronts (Tomczak and Godfrey, 1994) are
significant in the formation and maintenance of zoo-
geographical domains. The capacity of water masses
to control ostracod distribution can be seen in the
transitional Bonaerensian Province which lies at the
convergence of the cold Falklands and warm Brazil-
ian shelf currents.
The levels of ostracod endemicity on the con-
tinental shelf decline northwards as oceanic con-
ditions become more uniform. However, a second
transitional boundary (15–23ºS) exists between the
ostracods of the ‘Caribbean’ and Brazilian provinces.
North of this latitude typical Caribbean genera have
been described (Coimbra et al., 1992), while to the
south an admixture of northern and southern forms
can be found (Coimbra and Ornellas, 1989; Coim-
bra et al., 1995). Boltovskoy (1981) and Ramos
(1996) have suggested that a combination of sea-
sonal upwelling–downwelling, and the northward
penetration, at depth, of the Falklands Current would
aid the development of a seasonal thermal gradient
or barrier in this region. Endemicity in the Caribbean
remains low at 2.4%, and communality high at 0.3
due to the continued presence of easily traversed
(corridor and filter) migratory route ways.
7. Conclusion
The value of benthonic ostracods as tools in
palaeobiogeographical analysis have been exempli-
fied many times (reviewed in Whatley, 1988, p. 104).
It has been demonstrated that by measuring certain
biogeographical properties of faunal ‘nests’, one can
confirm modifications to both the structure and spa-
tio-temporal distribution of ostracods, and therefore
ascertain the mechanisms of change.
The philosophies of dispersal and vicariant bio-
geography advocate divergent mechanisms of dis-
persal, but neither can alone explain the distribution
patterns of Oligocene to Recent Ostracoda, from the
Neotropics to Antarctica. A compromise is required.
As with species, new genera also have centres
of origin, and it would appear that southern South
America represents such a region in the Oligocene.
Speciation in the Austral Basin was probably facil-
itated by the subdivision (vicariant event) of the
continental shelf by changing sea levels in the
Early Oligocene. Although a large number of genera
evolved on the southern peripheries of South Amer-
ica in the late Tertiary, few managed to disperse
northwards into the Caribbean; the exceptions were
 A.M. Wood et al. / Marine Micropaleontology 37 (1999) 345–364 357

Fig. 4. (a–d) Ostracod palaeobiogeography and oceanography (after Kennett, 1980; Duque-Caro, 1990) of the Neotropics and
Antarctica during the Oligocene, Early Miocene, Late Miocene and Pliocene. A major centre of ostracod origination existed in the
Austral Basin during the Oligocene, however, subsequent Miocene dispersal events were rare. Notable exceptions include the genera
Ambostracon (Ambostracon) and Coquimba which succeeded in colonising the Caribbean, Californian coast and Japan by the Pliocene.
Aa D Ambostracon (Ambostracon); Ad D Australicytheridea; Ae D Australicythere; Ap D Ambostracon (Patagonacythere); Ar D
Argenticytheretta; Be D Bensonia; Br D Brazilicythere; Ca D Caribella; Co D Coquimba; Me D Meridonalicythere; Or D Orionina; Pa
D Papillosacythere; Ps D Pseudoceratina; So D Soudanella.
358 A.M. Wood et al. / Marine Micropaleontology 37 (1999) 345–364

Ambostracon (Ambostracon) and Coquimba. These Miocene may have initiated the development of a
two genera appear to possess some preadaptation or proto-Panamanian Province (sensu Valentine, 1976)
fortuitous co-option, possibly linked to thermal tol- prior to the emergence of the Isthmus itself.
erance, which assisted their rapid dispersal into the Other than the constraints of physiology, it has
Caribbean and western Pacific. The general pattern been demonstrated that changing ocean currents and
of late Tertiary ostracod distribution and dispersal is water-mass temperature have regulated the dispersal
presented in Fig. 4a–d. potential of Recent and fossil shelf genera in the
The migratory success of benthonic ostracods is southwestern Atlantic Ocean; the result is ostracod
closely linked to extrinsic factors such as oceanog- provinciality.
raphy and climate, and the intrinsic physiology of
the taxa, but not distance alone. Passive dispersal
appears to be a significant phenomenon for relatively Appendix A
few taxa in the shallow marine realm (McKenzie,
Primary data sources, author(s), date of publication and research
1973; Witte, 1993). Alternative means were available region, used to calculate communality and endemicity values
to the Neotropical species Cyprideis salebrosa (Van
den Bold) and Cyprideis beaconensis (Leroy). By the Author Region
end of the Miocene both species existed throughout 1 Van den Bold, 1963a Cuba
the Americas, their dispersal agent was undoubtedly 2 Van den Bold, 1965 Puerto Rico
dynamic and avian (Van den Bold, 1976). 3 Van den Bold, 1966c Lesser Antilles
4 Van den Bold, 1957 Trinidad
Within the Meso-American region the availability 5 Van den Bold, 1958 Trinidad
of both filter and corridor pathways enabled rapid 6 Van den Bold, 1972a Venezuela
dispersal of shallow water ostracods during the late 7 Valicenti, 1977 Argentina
Tertiary, thus reducing levels of generic endemism in Bertels, 1975
the Caribbean to <2.7%. 8 Echevarrı́a, 1991 Argentina
9 Echevarrı́a, 1995 Argentina
Two major vicariant events occurred in the 10 Kielbowicz, 1988 Argentina
Neotropics and Antarctica during the late Tertiary. 11 Blaszyk, 1987 Antarctica
Antarctica separated from South America in the 12 Van den Bold, 1973 Cuba
?Oligo-Miocene, and the Central American Isth- 13 Van den Bold, 1965 Porto Rico
mus emerged, separating ostracod populations in 14 Van den Bold, 1966b Venezuela
15 Van den Bold, 1963a,b Venezuela
the Caribbean and the southern part of the Pacific 16 Van den Bold, 1966a,b,c,d,e Trinidad
coast of North America. Although data are scarce 17 Van den Bold, 1972a Venezuela
for Antarctica it would appear that a new suite of 18 Van den Bold, 1972b Panama
neo-endemic, cryophilic genera, including a number 19 Sanguinetti, 1979 Brazil
of extant species, emerged on this continent during 20 Echevarrı́a, 1987 Argentina
21 Van den Bold, 1968a,b, 1969, 1970, Dominican Rep.
the ?Mio-Pliocene (Szczechura and Blaszyk, 1996). 1972a,b, 1973, 1974, 1975, 1976,
However, an expanding Drake Passage may not have 1977, 1983, 1985, 1988
been a major barrier to the dispersal of ostracods as a 22 Van den Bold, 1969 Puerto Rico
number of conspecific taxa has been described from 23 Van den Bold, 1966e Venezuela
both the Recent Subantarctic and Antarctic provinces 24 Van den Bold, 1964 Venezuela
25 Van den Bold, 1972a Venezuela
(Whatley et al., 1996, 1998c). 26 Van den Bold, 1963b Trinidad
To the north, the emergence of the Central Amer- 27 Van den Bold, 1957 Trinidad
ican Isthmus was preceded in the Late Miocene by 28 Van den Bold, 1958 Trinidad
the closure of deep water portals and the disruption 29 Sanguinetti et al., 1991, 1992 Brazil
of intermediate oceanic circulation (Keller and Bar- 30 Van den Bold, 1966a Colombia
31 Zabert, 1978 Argentina
ron, 1983). The generic associations from the Lower 32 Zabert and Herbst, 1977 Argentina
Pliocene of the Caribbean and Mexico are analogous 33 Van den Bold, 1968a,b, 1969, 1970, Dominican Rep.
(Carreño, 1985), however, the species assemblages 1972a,b, 1973, 1974, 1975, 1976,
are not. The alteration in oceanic circulation in the 1977, 1983, 1985, 1988
 A.M. Wood et al. / Marine Micropaleontology 37 (1999) 345–364 359

Appendix A (continued) Appendix B. Presence and absence data for

Oligocene to Recent genera from the Neotropics
Author Region
to Antarctica
34 Van den Bold, 1975 Cuba
35 Van den Bold, 1975 Cuba Primary data sources are supplied in Appendix A.
36 Van den Bold, 1972a Venezuela
37 Van den Bold, 1963a,b Venezuela
38 Carreño, 1985 Mexico
39 Echevarrı́a, 1988 Argentina Appendix C. Jaccard Coefficient similarity
40 Szczechura and Blaszyk, 1996 Antarctica matrices for the Oligocene, Early Miocene, Late
41 Van den Bold, 1957, 1963b Trinidad Miocene and Pliocene
42 Van den Bold, 1972a Venezuela
43 Bertels et al., 1982 Brazil Primary data sources are supplied in Appendix A.
44 Bertels, 1975 Argentina
45 Teeter, 1975 Belize
46 Van den Bold, 1975 Cuba
47 Van den Bold, 1964 Venezuela References
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