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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 2.1. Vicariant events

and extinction.
Within the Neotropics, the majority of recent Vicariant species are closely related species that
biogeographical articles have focused on the Neo- occupy different geographical or ecological areas.
gene of Meso-America, and the consequences of an However, the term vicariant biogeography has de-
emergent Central American Isthmus (Savage, 1982; veloped a more specialised meaning than originally
Stehli and Webb, 1985; Woods, 1989; Jackson et intended by Croizat (1958). Accepting that dispersal
al., 1996). Many of the resulting theories of species can take place, vicariant biogeographers believe that
dispersal were based primarily upon the analysis the successful crossing of barriers is a rare event and
of terrestrial organisms (Raven and Axelrod, 1974; that most species have evolved in situ rather than as
Rosen, 1976; Gomez, 1982; Savage, 1982; Good- a consequence of dispersal (Rosen, 1976).
friend, 1989; Miller and Miller, 1989). Although Leon Croizat (1952) noted that whereas a disper-
exceptions exist (Petuch, 1988; Collins, 1996), many sal event would affect only a single species, vicariant
of the basic patterns in the marine realm have yet species should affect groups of different taxa which
to be described or discovered (Jackson and Budd, had a similar disjunct distribution pattern. The in-
1996). terpretations of vicariant biogeography have been
The central aim of this paper is to describe, in enormously strengthened by the widespread accep-
terms of dispersal events, key patterns in the distri- tance of continental drift and the theory of plate
bution of Oligocene–Recent South American marine tectonics.
Ostracoda. The disposition and quality of sample Throughout the Phanerozoic Eon, numerous vi-
sites are never perfect, however, the presently as- cariant and combinatory biotas have been recog-
sembled database of over sixty Recent and fossil nised using ostracods. In “Ostracoda and Continental
locations is unequalled. The identification of over Palaeogeography” Whatley (1988, p. 109) has intro-
140 ostracod genera has enabled the present authors duced a number of classic examples.
to identify dispersal events within two major palaeo-
migratory pathways. One path lies to the south be- 2.2. Dispersal events (sensu stricto)
tween Antarctica and southern Argentina, and the
second, the Central American Isthmus, to the north. The central hypothesis of dispersal biogeography
By measuring the level of generic communality and is that species originate in a particular area or cen-
endemicity across these pathways, we are able to tre and, if successful, spread out from that area to
measure changes in the spatio-temporal distribution colonise new habitats. Charles Darwin (1859) was
of ostracods, and allude to the (a)biotic mechanisms first to write about such centres which he called
that have modified them. A list of primary data “single centres of creation”. Many of the arguments
sources and genera presence–absence data can be associated with centres of origin have focused on
found in Appendixes A and B 1 . 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;
2. The conflicting philosophies of ‘dispersal’ Whatley, 1987; Witte, 1993).
biogeography

The study of species distribution patterns has 3. Tectonic history of the southern continents
led to the development of two schools of thought
which emphasise divergent processes and philoso- The tectonic history of South America and es-
phies (Nelson and Platnick, 1984; Hallam, 1988). pecially the Caribbean is extremely complex (Bar-
ron, 1987; Perfit and Williams, 1989; Coates and
Obando, 1996). Since the initial fragmentation of
1Appendix B can be found at http:==www.elsevier.nl=locate= the once unified super-continent of Gondwanaland in
marmicro or http:==www.elsevier.com=locate=marmicro. the Early Cretaceous, these southern continents have
 A.M. Wood et al. / Marine Micropaleontology 37 (1999) 345–364 347

undergone numerous, and geologically rapid vicari- width, ostracods were still able to migrate between
ant and convergent events. There are many problems South America and Africa (Reyment and Aranki,
concerning the mutual positions of Africa, South 1991). As a consequence, the Recent Ostracoda of
America and Antarctica during the Tertiary. How- West Africa and South America still retain many
ever, based upon palaeontological and structural evi- generic legacies of this early Tertairy union (Witte,
dence the most reliable reconstruction of continental 1993; Wood and Whatley, 1994).
outlines for the Cretaceous and Tertiary periods can Much of the research into mid-Tertiary to Re-
be found in Tarling (1972, 1980), Smith and Briden cent Meso-American, South American and Antarc-
(1977), and Golonka et al. (1995). tic shelf Ostracoda have tended to be descriptive
A detailed review of continental migration in the in nature, as workers strive to fill the taxonomic
Southern Hemisphere is beyond the scope of this void. Subsequent ostracod research is many tiered
paper. However, a short résumé of major tectonic with several operational scales of inquiry (Sepkoski,
episodes, and a simplistic reconstruction of palaeo- 1988): alpha, the analysis of taxa at a single lo-
continental movements, based upon geological evi- cality; beta, differentiation of taxa between sites;
dence have been included (Fig. 1): and gamma, the taxonomic differentiation between
geographical regions. In these terms the majority
Mesozoic Andean Orogeny and the development of work has been undertaken at the alpha scale,
of subduction zones to the southwest focusing on species (palaeo)ecology, bathymetry,
and west of South America. intra-regional biostratigraphy or localised eustasy.
Eocene Subduction to the west and within the Very little has been attempted at the larger beta or
Caribbean region itself, the latter lead- gamma scales, although exceptions do exist for the
ing to the formation of the Lesser and Caribbean (Van den Bold, 1977), Argentina (What-
Greater Antilles.
ley et al., 1998a,b), Chile (Hartmann-Schröder and
?Oligo-Miocene Separation of Antarctica from South
America, and the subsequent develop-
Hartmann, 1962; Hartmann, 1966), Brazil (Pinto and
ment of the Drake Passage. Ornellas, 1970, 1978) and Antarctica (summarised in
Mio-Pliocene Northward migration of South America, Hartmann, 1997).
localised shoaling of the Caribbean Sea.
Formation of an extended Central Amer- 4.1. Oligocene
ican archipelago then Isthmus.
The most note-worthy palaeobiogeographical
study of Cainozoic ostracods of Central–South
4. Previous ostracod research America was made by Van den Bold (1977), which
described faunal provinces and dispersal mecha-
Ostracod research of the South Atlantic Ocean nisms for a number of selected genera (Van den
was initiated by Brady (1870, 1880, 1907) in the lat- Bold, 1970, 1974). Similarly, the distribution of
ter half of the Nineteenth Century, but only recently hemicytherid genera in the southern hemisphere has
have these founding taxonomic masterpieces been been outlined by Valicenti (1977).
augmented. A preponderance of ostracodological re- A register of Argentinean Palaeogene ostracods
search has focused on older Tethyan migration routes has recently been compiled by Echevarrı́a (1995),
(Dingle, 1988; Whatley, 1988; Whatley and Ballent, while additional references can be found in Ber-
1994; Boomer and Ballent, 1996), the juxtaposition tels (1975) and Kielbowicz (1988). Ostracods of
of southern continents (Whatley, 1988; Reyment and Oligocene age have only been described from one
Aranki, 1991), and mid-Tertiary faunas of Meso- Antarctic site on King George Island (Blaszyk,
America (Van den Bold, 1957, 1958, 1963a,b, 1964, 1987). This fauna was thought by Blaszyk to re-
1965, 1966a,b,c,d,e, 1968a,b, 1969, 1970, 1972a,b, semble (in part) assemblages obtained from Isla
1973, 1974, 1975, 1976, 1977, 1983, 1985, 1988). Grande de Tierra del Fuego, in southern Argentina
It has also been established that during the early (Echevarrı́a, 1982, 1987).
Tertiary, while the South Atlantic was half its present
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) 345–364 349

essentially descriptive and=or has focused on the result of a succession of publications by Whatley et
stratigraphical application of ostracods (Echevarrı́a, al. (1987, 1988, 1995, 1996, 1998a,b).
1987; Sanguinetti et al., 1991, 1992; Carreño et
al., 1999). Von Ohmert (1968, 1971) published two 4.4.3. Antarctica
important papers on the Pliocene to Recent radi- Before 1964 Antarctic ostracod research was
ation of the genus Caudites, and the endemicity dominated by taxonomy. A brief history of research
of the subfamily Coquimbinae in Chile. More re- has recently been presented by Hartmann (1997), and
cently, Cronin and Dowsett (1996) have used R- and only one publication (Whatley et al., 1999) postdates
Q-mode cluster analyses in order to identify changes this. In a short zoogeographical summary, Hartmann
in ostracod communities and palaeoceanography in (1997) suggested that the faunas from the Antarctic
the Caribbean. were littoral Tertiary relics. The indigenous charac-
ter, and Recent distribution pattern of southern ocean
4.3. Pleistocene species were thought by Benson (1964) and Hazel
(1967) to be a product of climate and topography.
The few papers on the subject of Pleistocene Os-
tracoda have dealt mainly with relationship between
faunal associations and sea-level (Bertels, 1975; Vi- 5. Methodology: binary similarity coefficients
calvi et al., 1977; Bertels et al., 1982; Bertels and and endemicity as aids in panbiogeographical
Martinez, 1990; Aguirre and Whatley, 1995). study

4.4. Recent Panbiogeography is a philosophy that focuses

upon the measurement of communality (or similar-
Research into Recent ostracods has focused upon ity) between biotas of different geographical regions
three geographical regions: (Myers and Giller, 1994). Two contrasting measures,
binary similarity coefficients and percentage generic
4.4.1. Meso-America endemicity, have been used to study the spatio-tem-
Numerous Recent samples were collected and poral evolution of Meso-American, South American
analyzed by Van den Bold (1964, 1972b, 1975) dur- and Antarctic ostracods.
ing his exploration of Neogene successions. These
Recent data confirmed the presence of two zoogeo- 5.1. Binary similarity coefficients
graphical (sub)provinces, the Gulf of Mexico and
the Caribbean (Van den Bold, 1977). While working Binary similarity coefficients are essentially used
on sub-Recent samples from Belize, Teeter (1975) to measure beta diversity or the degree of association
recognised similarities in the generic associations of or similarity between paired sample sites (Whittaker,
the Recent and palaeo-Caribbean. 1977; Wilson and Shmida, 1984; Magurran, 1988;
Shi, 1993; Hallam, 1994; Rosenzweig, 1995).
4.4.2. Argentina, Brazil and Chile A large choice of binary measures are available,
A number of small-scale, sub-provincial ostra- although the Jaccard and Sorenson indices are most
cod communities has been recognised in northeast- commonly used (Hazel, 1970; Janson and Vegelius,
ern Brazil which contain species common to the 1981; Neil, 1995). In the field of ostracodology, the
Lesser Antilles and Central America (Coimbra et al., Simpson and Sanders coefficients were employed
1992). However, the Recent ostracod biotas along the by Van den Bold (1966a, 1972a) and Neil (1995)
Pacific (Hartmann-Schröder and Hartmann, 1962, respectively, as an aid to correlation. The simplicity
1965) and Atlantic (Whatley et al., 1998c) coasts of such coefficients are their greatest advantage,
of South America have been segregated into large however, one must be reminded that all groups count
geographically restricted areas or provinces. The equally irrespective of their abundance. The Jaccard
subdivision of southwestern Atlantic ostracod as- binary measure was chosen for the current study:
semblages into zoogeographical provinces is as a 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 The existence of amphi-atlantic taxa, our inade-
faunas; a D the number of species in fauna A, and b quate knowledge of late Tertiary African faunas, and
D the number of species in fauna B. dubious taxonomy has made the identification of a
The calculated levels of intra and inter-regional small number of endemic genera presently impossi-
generic similarity in the Oligocene, Early Miocene, ble.
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- 6. Results and discussion
sented in Appendix C 2 .
6.1. Oligocene
5.2. Endemicity
6.1.1. Palaeogeography and oceanography
Stenotopic or endemic biota can be broadly di- A detailed account of South American palaeo-
vided into two groups: neoendemics and palaeoen- (bio)geography can be found in Hallam (1994) and
demics (Engler, 1882). Neoendemics are confined Jackson et al. (1996). At the end of the Eocene the
in their distribution to the areas in which they Central American Isthmus had formed a continuous,
evolved, whereas palaeoendemics are relicts, iso- but submerged structural unit. A number of large sea-
lated geographically by extinction. However, both ways still existed between North and South America
types are influenced by contemporaneous ecologi- (Pindell et al., 1988). To the south, the development
cal factors and=or historical, and large-scale abiotic of the Drake Passage may well have been initiated
processes. The study of endemics has proved to be during the Oligocene (Hallam, 1994), although an
very useful, revolutionising biogeography with the epicontinental sea still existed between Patagonia
introduction of new methodologies (Humphries and and the West Antarctic Peninsula. The anticyclonic
Parenti, 1986) and measures (Bykov, 1979). Most re- gyre of surface currents in the South Atlantic was
cently, Boomer and Whatley (1996), Larwood et al. much larger allowing the warmer Brazilian current
(1996) and Larwood and Whatley (1993) have high- to extend farther south.
lighted that prolonged spatial isolation is essential to
the generation of endemic ostracods, therefore, by 6.1.2. Ostracod endemicity and communality
measuring the latter one can allude to the former. (Fig. 2a)
Endemicity is measured as a percentage, where gt is Due to incomplete Oligocene coverage, percent-
the total number of genera in a given region, and ge age generic endemicity has only been calculated for
the number of genera restricted to that region: three regions, the Austral Basin of South America
ge (18%), Western Antarctic Peninsula (0%) and Meso-
Percentage endemism D t ð 100
g America (3.4%). These regions were considered to
Where data allows, the percentage of generic be important migratory pathways during the Tertiary
endemicity has been calculated for Meso-America, (Simpson, 1940), however, the especially high levels
South America and Antarctica. Meso-America is of ostracod endemicity is indicative of low levels of
considered by the present authors to include the faunal interchange between the Austral Basin and the
Lesser Antilles, Greater Antilles, Central American Western Antarctic Peninsula. As a comparison, Re-
region between 23 and 8ºN, Colombia and Vene- cent ostracod endemicity values in equatorial West
zuela. As with communality, the regional endemicity Africa do not exceed 4.3%, while values for south-
of Neotropical to Antarctic ostracods is also pre- ern and northern Europe are almost zero (Wood and
sented on a series of reconstructed maps (Fig. 2a–d). Whatley, 1994).
A complete list of endemic genera is also given in A significant number of new ostracod genera first
Table 1. appeared in the Austral Basin of southern Argentina
during the Oligocene, making this an important cen-
2Appendix C can be found at http:==www.elsevier.nl=locate= tre of origin. Such levels of generic origination are
marmicro or http:==www.elsevier.com=locate=marmicro. 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 and South America (Bertels, 1969, 1975; Neufville,
level and habitat patchiness. The continental shelf 1979). However, since the Eocene its distribution
of southeastern South America covers a huge area has contracted southwestwards to southern Argentina
of some 1 million km2 . The greatest opportunity for (Bertels, 1975; Echevarrı́a, 1995) where it remained
speciation was probably associated with the subdi- as a palaeo-endemic until its final extinction in
vision of this epicontinental sea (and the nonplank- the Pliocene (Echevarrı́a, 1988). The accounts of
totrophic ostracod population) by changing sea levels Soudanella from the Neogene of West Africa are
during the Early Oligocene (Valentine and Jablonski, erroneous. This genus appears to have been confused
1983; Prothero and Berggren, 1992). with both Ruggieria and Keijella (Carbonnel, 1992).
Nine neo-endemic (evolved in situ) ostracod gen- Only one neo-endemic, Orionina has been de-
era and subgenera have been identified from south- scribed from the Oligocene of Meso-America (Van
ern Argentina. Only Soudanella Apostolescu can be den Bold, 1963a, 1965). Its status as an endemic was
considered a palaeo-endemic. A complete listing of short lived for rapid migration, via shoals in the An-
endemics is given in Table 1. Soudanella is an inter- tilles and along the coast of the emerging Central
esting genus for it remains the only palaeo-endemic American Isthmus, enabled it to colonise the southern
recorded from the upper Tertiary of South Amer- coasts of North America in the ?Early to mid-Miocene
ica. This genus was originally described from the (Swain, 1951). However, McKenzie (1987) suggests
Palaeocene of Senegal, however, it is now known that as few as 5% of species occurred in both the
to have occurred in the Palaeogene of the Middle Caribbean and Gulf Coast in the Early Oligocene.
East (Bassiouni, 1969), North Africa (Reyment and Similar filter routes were envisaged by Van den
Reyment, 1981), Caribbean (Van den Bold, 1975) Bold (1974) to account for the northward migration
 A.M. Wood et al. / Marine Micropaleontology 37 (1999) 345–364 353

of shallow water species of Cativella, Pellucistoma and East Antarctic Ice Sheet (Frankes et al., 1992).
and Costa from the Oligocene of Venezuela and The closure of the E–W-trending deep water con-
Colombia. Van den Bold also revealed that the dis- nection occurred in the Caribbean in association with
persal rates of ostracod genera were quite variable a regional shift from deep to mid bathyal-neritic con-
(Van den Bold, 1974, fig. 3). Puriana was considered ditions (Coates and Obando, 1996).
to be a neo-endemic of the Meso-American region,
however, it appears to have first appeared simultane- 6.2.2. Ostracod endemicity and communality
ously in the Oligocene of Puerto Rico (Van den Bold, (Fig. 2b)
1965), the Lesser Antilles (Van den Bold, 1966c) and Ostracod endemicity values for the Austral Basin
Southern Mississippi (Hazel et al., 1980). of South America are complicated by the presence
Mean intra-regional similarity values of between of Lazarus genera: taxa which seem to suffer ex-
0.28 and 0.41 for generic associations on island tinction but then reappear later in the stratigraphical
sites within the Greater and Lesser Antilles, would record (Jablonski, 1986). Neogene neo-endemics,
also support the idea of unrestrained interchange. including Papillosacythere, Australicythere, Benso-
However, these figures contrast considerably with nia, Australicytheridea, Brasilicythere and Argenti-
low inter-regional values between north and south. cytheretta Rose, 1975, appear to become extinct only
Although a passive transport agent existed in the to re-emerge from the dead (see Table 1). These
form of the warm Brazilian Current, the average genera undoubtedly existed within this region but are
similarity value of 0.14 would indicate restricted dis- as yet undiscovered. If Lazarus endemics are not in-
persal along the southwestern Atlantic shelf during cluded in the calculation, Early Miocene endemicity
the Oligocene. Paradoxically, the absence of a cold appears to decrease to 11%.
Falklands Current, and a less formidable temperature However, the eurythermal genus Coquimba does
gradient, should have encouraged ostracod dispersal appear to be an authentic escapee from the confine-
along the shelf of southern Brazil. ments of the Austral Basin, having being recorded
The King George Island assemblage of Antarctica from the Miocene of the Caribbean (Van den Bold,
(Blaszyk, 1987) has a low diversity and no endemics; 1972a, 1973). In order to eliminate the need for pos-
the assumption of remoteness is also supported by a tulating a single, epic, migratory event of some 3500
low communality score (0.15). miles, we speculate that intermediate stations may
have existed in Brazil.
6.2. Early Miocene No endemic ostracod genera were recorded from
Meso-America. The relative ease of genera redis-
6.2.1. Palaeogeography and oceanography tribution in the Caribbean appears to confirm a re-
The severing of Antarctica from Patagonia, and gional shift to shallower oceanic conditions and the
the establishment of a sea-way between Australia continued shoaling of the Antilles. Cronin (1987)
and Antarctica lead to the development of the cir- considered dispersal among Caribbean islands was
cum-Antarctic oceanic circulation system, and its passive, and not directly related to specific abiotic
northerly branch the Falklands (Malvinas) Current. events. Indeed, the colonisation of shallow water
The precise timing of this event is difficult to pin- habitats within the Caribbean could easily have been
point, however, Hallam (1994) suggests the Early achieved via natural rafts of drifting debris.
Oligocene. Additional evidence from planktonic fo- A number of low similarity values has been
raminiferal assemblages in the South Atlantic in- recorded between faunas of the Lesser and Greater
dicate ocean cooling across the Oligocene=Mio- Antilles (0.07 between Trinidad and Puerto Rico).
cene boundary (Spezzaferri, 1995), in the Middle These low figures have arisen because one is com-
Miocene (Flower and Kennett, 1994), and latest paring bathymetrically divergent faunas, bathyal ver-
Miocene (Boltovskoy, 1979, 1980). These Neo- sus neritic. The former depth zone is characterised
gene events may denote major changes in deep throughout the Caribbean by the genera Cardobair-
ocean circulation related to the gradual development dia, Krithe, Ambocythere, Bradleya and Henry-
of the circum-Antarctic oceanic circulation system howella (Van den Bold, 1963b, 1969).
354 A.M. Wood et al. / Marine Micropaleontology 37 (1999) 345–364

Most significantly the communality between 6.4.2. Ostracod endemicity and communality
southern Brazil and the Caribbean reaches its acme (Fig. 2d)
within the Miocene with a similarity value of 0.21. As the Central American Isthmus emerged in the
An important fauna containing the genera Bair- Late Pliocene a ‘corridor’ type pathway was estab-
doppilata, Pellucistoma and the species Orionina lished which aided the migration of benthonic ostra-
vaughani (Ulrich and Bassler), were described from cods. The result was a further decline in Caribbean
the Pelotas Basin (32ºS) by Sanguinetti (1979). In endemism to only 1.2%. With mean similarity values
the Recent, Orionina vaughani occurs as far south of between 0.36 and 0.42, intra-regional commu-
as Espirito Santo State (20ºS). The southward dis- nality remained high within the Caribbean Sea, and
placement of ‘equatorial’ genera into southern Brazil between the Caribbean and the southern part of the
during the Early Miocene was probably linked to tec- Pacific coast of North America. Before the forma-
tonism in the south, and the gradual development of tion of the isthmus two Oligocene endemics of Ar-
the circum-Antarctic oceanic circulation system. The gentina, Ambostracon (Ambostracon) and Coquimba
subsequent decoupling of the South Atlantic gyre were able to enter the Gulf of California (Valentine,
from Antarctic waters resulted in the intensification 1976; Carreño, 1985), whether this was achieved
of the warm Brazilian Current (Kennett, 1980). via a Central American portal or the west coast of
South America remains unclear. One younger en-
6.3. Late Miocene demic, Pseudoceratina, which emerged in the Late
Miocene remains confined to the Caribbean.
6.3.1. Palaeogeography and oceanography It is likely that the distinctive species character
Partial uplift of the submerged Central Ameri- of the Pacific coast and Caribbean–Gulf Coast ostra-
can Isthmus occurred with a resultant disruption of cod communities was established before the Early–
the California Current (Duque-Caro, 1990; Coates mid-Pliocene (Carreño, 1985). However, the isola-
and Obando, 1996). Major tectonic activity in the tion of ostracod populations by the Central Amer-
Caribbean Sea also resulted in the rapid uplift and ican Isthmus appears not to have aided speciation
subsidence of parts of the ocean floor (Van den Bold, among pre-isthmus species of the genera Puriana
1968b, 1988; Steineck et al., 1984). (Cronin, 1987) or Orionina (Gunther and Swain,
1976; Cronin and Schmidt, 1988).
6.3.2. Ostracod endemicity and communality The percentage of endemics remained high in
(Fig. 2c) South America (11.7%) as the southward migration
Both endemicity and communality parallel the of Antarctica lead to the expansion of the Drake Pas-
Oligocene, although two neo-endemics, Caribella sage. The computed values for endemicity (9%) and
and Pseudoceratina are recorded from Meso-Amer- communality (0.08) of the Cockburn Island assem-
ica. High similarity values (0.5) were obtained blage in Antarctica (Szczechura and Blaszyk, 1996)
(as expected) between sites from Venezuela and affirm this progressive disconnection.
Trinidad, but as in the Oligocene, the similarity
between genera of the north and south remains low 6.5. Recent
(0.14). Only common cosmopolites such as Aurila,
Bradleya, Cytherella, Krithe and Xestoleberis occur 6.5.1. Ostracod endemicity and communality
in both regions. Sadly, no ostracods have yet been (Fig. 3; Table 2)
described from the Miocene of Antarctica. A recent succession of geographically diverse
publications has improved our basic knowledge of
6.4. Pliocene ostracod endemicity and communality in the south-
western Atlantic (Fig. 3; Table 2; Whatley et al.,
6.4.1. Palaeogeography and oceanography 1987, 1988, 1995, 1996, 1998a,b). The communal-
The Central American Isthmus emerged in the ity values are presented in a similarity matrix that
Late Pliocene (Jackson et al., 1996) in association is subdivided on the basis of mean regional sim-
with renewed uplift of the Andes. 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

transitional Bonaerensian Province which lies at the

convergence of the cold Falklands and warm Brazil-
ian shelf currents.
?2.4%
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
4% 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
8.8% (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
9.6% 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
RECENT (corridor and filter) migratory route ways.

Fig. 3. Recent regionalised endemicity values of ostracod shelf

genera from the southwestern Atlantic sea board.
7. Conclusion

Four major zoogeographical regions are recognised: The value of benthonic ostracods as tools in
the Caribbean, Brazilian, Subantarctic and Antarctic. palaeobiogeographical analysis have been exempli-
The spatial extent of these regions appear to parallel fied many times (reviewed in Whatley, 1988, p. 104).
those recently described by Whatley et al. (1998c). It has been demonstrated that by measuring certain
The only deviation from their scheme is the apparent biogeographical properties of faunal ‘nests’, one can
absence of their Bonaerensian Province (43–36ºS). confirm modifications to both the structure and spa-
It is certain that differences in the scale of sampling tio-temporal distribution of ostracods, and therefore
and taxonomic investigation has caused this transi- ascertain the mechanisms of change.
tional (ecotonal) province to be subsumed within the The philosophies of dispersal and vicariant bio-
Brazilian and Subantarctic provinces. geography advocate divergent mechanisms of dis-
At 8%, levels of endemicity remain high in persal, but neither can alone explain the distribution
Argentina=southern Brazil, while the continued iso- patterns of Oligocene to Recent Ostracoda, from the
lation of Antarctica appears to have assisted en- Neotropics to Antarctica. A compromise is required.
demicity where values rise to 9.6%. However, dis- As with species, new genera also have centres
tance alone would not suffice in maintaining iso- of origin, and it would appear that southern South
lation and, therefore, endemicity. As within the America represents such a region in the Oligocene.
northeastern Atlantic system (Wood and Whatley, Speciation in the Austral Basin was probably facil-
1994), oceanic structures such as the circum-Antarc- itated by the subdivision (vicariant event) of the
tic oceanic circulation system, Subantarctic and continental shelf by changing sea levels in the
Antarctic fronts (Tomczak and Godfrey, 1994) are Early Oligocene. Although a large number of genera
significant in the formation and maintenance of zoo- evolved on the southern peripheries of South Amer-
geographical domains. The capacity of water masses ica in the late Tertiary, few managed to disperse
to control ostracod distribution can be seen in the 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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