The palaeozoogeography of Oligocene to Recent marine Ostracoda from the Neotropics (mid- and South America) and Antarctica

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 Ineˆs 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 Geocie ˆ ncias, 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 The science of biogeography, and especially the related discipline of biodiversity, have gained global Ł Corresponding

author.

importance as an issue of both scientific and political concern (Wilson, 1997). It was appropriate that the first Environmental Summit Meeting was held in Rio de Janeiro, Brazil for here in South America an unparalleled and priceless biota now exists which 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

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biological processes such as speciation, dispersal and extinction. Within the Neotropics, the majority of recent biogeographical articles have focused on the Neogene 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; Goodfriend, 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 distribution of Oligocene–Recent South American marine Ostracoda. The disposition and quality of sample sites are never perfect, however, the presently assembled 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 palaeomigratory pathways. One path lies to the south between 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 philosophies (Nelson and Platnick, 1984; Hallam, 1988).

1

Appendix 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 developed 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 dispersal event would affect only a single species, vicariant species should affect groups of different taxa which had a similar disjunct distribution pattern. The interpretations of vicariant biogeography have been enormously strengthened by the widespread acceptance of continental drift and the theory of plate tectonics. Throughout the Phanerozoic Eon, numerous vicariant and combinatory biotas have been recognised using ostracods. In “Ostracoda and Continental Palaeogeography” Whatley (1988, p. 109) has introduced 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 centre 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 Triangle 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 especially the Caribbean is extremely complex (Barron, 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

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undergone numerous, and geologically rapid vicariant and convergent events. There are many problems concerning the mutual positions of Africa, South America and Antarctica during the Tertiary. However, based upon palaeontological and structural evidence 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 re´sume´ of major tectonic episodes, and a simplistic reconstruction of palaeocontinental movements, based upon geological evidence 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 leading to the formation of the Lesser and Greater Antilles. ?Oligo-Miocene Separation of Antarctica from South America, and the subsequent development of the Drake Passage. Mio-Pliocene Northward migration of South America, localised shoaling of the Caribbean Sea. Formation of an extended Central American archipelago then Isthmus.

4. Previous ostracod research Ostracod research of the South Atlantic Ocean was initiated by Brady (1870, 1880, 1907) in the latter half of the Nineteenth Century, but only recently have these founding taxonomic masterpieces been augmented. A preponderance of ostracodological research 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 MesoAmerica (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

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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 Recent Meso-American, South American and Antarctic 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 locality; 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 (Whatley et al., 1998a,b), Chile (Hartmann-Schro¨der 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 mechanisms 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 Echevarrı´a (1995), while additional references can be found in Bertels (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 resemble (in part) assemblages obtained from Isla Grande de Tierra del Fuego, in southern Argentina (Echevarrı´a, 1982, 1987).

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Pliocene (3mya)

Late Miocene (6mya)

Greater Antilles

Lesser Antilles

North America

South America

Greater Antilles

Lesser Antilles

North America

South America

Antarctica

Australia

Antarctica

Australia

CONVERGENCE Greater Antilles

Early Miocene (20mya)

Lesser Antilles

Antarctica North America

Australia

South America

VICARIANCE

Greater Antilles

Eocene/Oligocene (35mya)

North America

Lesser Antilles

South America Antarctica

Australia

VICARIANCE

Paleocene (60mya)

North America

South 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 Much of Van den Bold’s earliest work on Caribbean ostracods was taxonomic but also included a reli-

able post-Eocene biostratigraphical zonation scheme (Van den Bold, 1983) based upon rapid evolutionary (i.e., ‘Radimella’) events (Van den Bold, 1988). In Brazil and Argentina nearly all research has been

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essentially descriptive and=or has focused on the stratigraphical application of ostracods (Echevarrı´a, 1987; Sanguinetti et al., 1991, 1992; Carren˜o et al., 1999). Von Ohmert (1968, 1971) published two important papers on the Pliocene to Recent radiation of the genus Caudites, and the endemicity of the subfamily Coquimbinae in Chile. More recently, 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 Ostracoda have dealt mainly with relationship between faunal associations and sea-level (Bertels, 1975; Vicalvi 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) during his exploration of Neogene successions. These Recent data confirmed the presence of two zoogeographical (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 ostracod communities has been recognised in northeastern 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-Schro¨der 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 assemblages into zoogeographical provinces is as a

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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 character, 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 similarity) 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-temporal 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)

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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 number of palaeogeographical maps (Fig. 2a–d; Table 2). Complete similarity matrices for fossil sites are presented in Appendix C 2 .

The existence of amphi-atlantic taxa, our inadequate knowledge of late Tertiary African faunas, and dubious taxonomy has made the identification of a small number of endemic genera presently impossible.

6. Results and discussion 6.1. Oligocene

5.2. Endemicity Stenotopic or endemic biota can be broadly divided into two groups: neoendemics and palaeoendemics (Engler, 1882). Neoendemics are confined in their distribution to the areas in which they evolved, whereas palaeoendemics are relicts, isolated geographically by extinction. However, both types are influenced by contemporaneous ecological 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 recently, Boomer and Whatley (1996), Larwood et al. (1996) and Larwood and Whatley (1993) have highlighted 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 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 Venezuela. As with communality, the regional endemicity of Neotropical to Antarctic ostracods is also presented on a series of reconstructed maps (Fig. 2a–d). A complete list of endemic genera is also given in Table 1. 2

Appendix C can be found at http:==www.elsevier.nl=locate= marmicro or http:==www.elsevier.com=locate=marmicro.

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 seaways 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, percentage generic endemicity has only been calculated for three regions, the Austral Basin of South America (18%), Western Antarctic Peninsula (0%) and MesoAmerica (3.4%). These regions were considered to 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, Recent ostracod endemicity values in equatorial West Africa do not exceed 4.3%, while values for southern 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 centre of origin. Such levels of generic origination are undoubtedly linked to a number of mutually in-

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351

no endemics

1.7%

0.28

0.33

0.36 0.41 0.34

0.07 0.54

0.44

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 0.28 0.57

0.36

1.2%

0.39

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.

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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

Meso-America South America

Orionina

A. (Ambostracon) no endemics A. (Patagonacythere) Argenticytheretta Australicythere Australicytheridea Bensonia Brasilicythere Coquimba Papillosacythere Soudanella

South America

A. (Patagonacythere) Caribella Argenticytheretta Pseudoceratina Brasilicythere Papillosacythere a Australicythere a Soudanella a Bensonia a Australicytheridea a

no data

Antarctica no data

Antarctica no endemics Pliocene endemics

Pleistocene–Recent endemics

Meso-America South America

Antarctica

Pseudoceratina Papillosacythere Soundanella Brasilicythere a Australicytheridea a Argenticytheretta a

Meridionalicythere Caribella Ruessicythere

a Lazarus

Argenticytheretta Australicythere Bensonia Brasilicythere Australicytheridea a Papillosacythere a Soudanella a

Antarctica

Meso-America

North Brazil

South Brazil=Argentina Antarctica

Tanella Whatleyella

Argenticytheretta Australicytheridea Austroaurila Brasilicythere Falklandia Papillosacythere new genus A

Antarcticythere Austrocythere Macroscapha Pelecocythere Pontocypria new genus B

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 subdivision of this epicontinental sea (and the nonplanktotrophic 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 genera and subgenera have been identified from southern Argentina. Only Soudanella Apostolescu can be considered a palaeo-endemic. A complete listing of endemics is given in Table 1. Soudanella is an interesting genus for it remains the only palaeo-endemic recorded from the upper Tertiary of South America. 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; Echevarrı´a, 1995) where it remained as a palaeo-endemic until its final extinction in the Pliocene (Echevarrı´a, 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 described 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 Antilles 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

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of shallow water species of Cativella, Pellucistoma and Costa from the Oligocene of Venezuela and Colombia. Van den Bold also revealed that the dispersal 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 simultaneously 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 dispersal 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 circum-Antarctic oceanic circulation system, and its northerly branch the Falklands (Malvinas) Current. The precise timing of this event is difficult to pinpoint, however, Hallam (1994) suggests the Early Oligocene. Additional evidence from planktonic foraminiferal assemblages in the South Atlantic indicate ocean cooling across the Oligocene=Miocene boundary (Spezzaferri, 1995), in the Middle Miocene (Flower and Kennett, 1994), and latest Miocene (Boltovskoy, 1979, 1980). These Neogene events may denote major changes in deep ocean circulation related to the gradual development of the circum-Antarctic oceanic circulation system

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and East Antarctic Ice Sheet (Frankes et al., 1992). The closure of the E–W-trending deep water connection occurred in the Caribbean in association with a regional shift from deep to mid bathyal-neritic conditions (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 extinction but then reappear later in the stratigraphical record (Jablonski, 1986). Neogene neo-endemics, including Papillosacythere, Australicythere, Bensonia, Australicytheridea, Brasilicythere and Argenticytheretta 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 included 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 confinements 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 postulating 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 redistribution in the Caribbean appears to confirm a regional 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 comparing bathymetrically divergent faunas, bathyal versus neritic. The former depth zone is characterised throughout the Caribbean by the genera Cardobairdia, Krithe, Ambocythere, Bradleya and Henryhowella (Van den Bold, 1963b, 1969).

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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 Bairdoppilata, 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 displacement of ‘equatorial’ genera into southern Brazil during the Early Miocene was probably linked to tectonism 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 American 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-America. 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 established which aided the migration of benthonic ostracods. 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 communality remained high within the Caribbean Sea, and between the Caribbean and the southern part of the Pacific coast of North America. Before the formation of the isthmus two Oligocene endemics of Argentina, Ambostracon (Ambostracon) and Coquimba were able to enter the Gulf of California (Valentine, 1976; Carren˜o, 1985), whether this was achieved via a Central American portal or the west coast of South America remains unclear. One younger endemic, 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 ostracod communities was established before the Early– mid-Pliocene (Carren˜o, 1985). However, the isolation of ostracod populations by the Central American 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 Passage. The computed values for endemicity (9%) and communality (0.08) of the Cockburn Island assemblage 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 southwestern Atlantic (Fig. 3; Table 2; Whatley et al., 1987, 1988, 1995, 1996, 1998a,b). The communality values are presented in a similarity matrix that is subdivided on the basis of mean regional similarities 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

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?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 transitional (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 isolation of Antarctica appears to have assisted endemicity where values rise to 9.6%. However, distance alone would not suffice in maintaining isolation and, therefore, endemicity. As within the northeastern Atlantic system (Wood and Whatley, 1994), oceanic structures such as the circum-Antarctic oceanic circulation system, Subantarctic and Antarctic fronts (Tomczak and Godfrey, 1994) are significant in the formation and maintenance of zoogeographical 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 Brazilian shelf currents. The levels of ostracod endemicity on the continental shelf decline northwards as oceanic conditions 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; Coimbra et al., 1995). Boltovskoy (1981) and Ramos (1996) have suggested that a combination of seasonal 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 exemplified 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 spatio-temporal distribution of ostracods, and therefore ascertain the mechanisms of change. The philosophies of dispersal and vicariant biogeography advocate divergent mechanisms of dispersal, 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 facilitated 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 America in the late Tertiary, few managed to disperse northwards into the Caribbean; the exceptions were

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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.

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Ambostracon (Ambostracon) and Coquimba. These two genera appear to possess some preadaptation or fortuitous co-option, possibly linked to thermal tolerance, which assisted their rapid dispersal into the Caribbean and western Pacific. The general pattern of late Tertiary ostracod distribution and dispersal is presented in Fig. 4a–d. The migratory success of benthonic ostracods is closely linked to extrinsic factors such as oceanography and climate, and the intrinsic physiology of the taxa, but not distance alone. Passive dispersal appears to be a significant phenomenon for relatively few taxa in the shallow marine realm (McKenzie, 1973; Witte, 1993). Alternative means were available to the Neotropical species Cyprideis salebrosa (Van den Bold) and Cyprideis beaconensis (Leroy). By the end of the Miocene both species existed throughout the Americas, their dispersal agent was undoubtedly dynamic and avian (Van den Bold, 1976). Within the Meso-American region the availability of both filter and corridor pathways enabled rapid dispersal of shallow water ostracods during the late Tertiary, thus reducing levels of generic endemism in the Caribbean to <2.7%. Two major vicariant events occurred in the Neotropics and Antarctica during the late Tertiary. Antarctica separated from South America in the ?Oligo-Miocene, and the Central American Isthmus emerged, separating ostracod populations in the Caribbean and the southern part of the Pacific coast of North America. Although data are scarce for Antarctica it would appear that a new suite of neo-endemic, cryophilic genera, including a number of extant species, emerged on this continent during the ?Mio-Pliocene (Szczechura and Blaszyk, 1996). However, an expanding Drake Passage may not have been a major barrier to the dispersal of ostracods as a number of conspecific taxa has been described from both the Recent Subantarctic and Antarctic provinces (Whatley et al., 1996, 1998c). To the north, the emergence of the Central American Isthmus was preceded in the Late Miocene by the closure of deep water portals and the disruption of intermediate oceanic circulation (Keller and Barron, 1983). The generic associations from the Lower Pliocene of the Caribbean and Mexico are analogous (Carren˜o, 1985), however, the species assemblages are not. The alteration in oceanic circulation in the

Miocene may have initiated the development of a proto-Panamanian Province (sensu Valentine, 1976) prior to the emergence of the Isthmus itself. Other than the constraints of physiology, it has been demonstrated that changing ocean currents and water-mass temperature have regulated the dispersal potential of Recent and fossil shelf genera in the southwestern Atlantic Ocean; the result is ostracod provinciality.

Appendix A Primary data sources, author(s), date of publication and research region, used to calculate communality and endemicity values

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21

22 23 24 25 26 27 28 29 30 31 32 33

Author

Region

Van den Bold, 1963a Van den Bold, 1965 Van den Bold, 1966c Van den Bold, 1957 Van den Bold, 1958 Van den Bold, 1972a Valicenti, 1977 Bertels, 1975 Echevarrı´a, 1991 Echevarrı´a, 1995 Kielbowicz, 1988 Blaszyk, 1987 Van den Bold, 1973 Van den Bold, 1965 Van den Bold, 1966b Van den Bold, 1963a,b Van den Bold, 1966a,b,c,d,e Van den Bold, 1972a Van den Bold, 1972b Sanguinetti, 1979 Echevarrı´a, 1987 Van den Bold, 1968a,b, 1969, 1970, 1972a,b, 1973, 1974, 1975, 1976, 1977, 1983, 1985, 1988 Van den Bold, 1969 Van den Bold, 1966e Van den Bold, 1964 Van den Bold, 1972a Van den Bold, 1963b Van den Bold, 1957 Van den Bold, 1958 Sanguinetti et al., 1991, 1992 Van den Bold, 1966a Zabert, 1978 Zabert and Herbst, 1977 Van den Bold, 1968a,b, 1969, 1970, 1972a,b, 1973, 1974, 1975, 1976, 1977, 1983, 1985, 1988

Cuba Puerto Rico Lesser Antilles Trinidad Trinidad Venezuela Argentina Argentina Argentina Argentina Antarctica Cuba Porto Rico Venezuela Venezuela Trinidad Venezuela Panama Brazil Argentina Dominican Rep.

Puerto Rico Venezuela Venezuela Venezuela Trinidad Trinidad Trinidad Brazil Colombia Argentina Argentina Dominican Rep.

A.M. Wood et al. / Marine Micropaleontology 37 (1999) 345–364 Appendix A (continued) Author 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52

53

54 55 56

57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74

Van den Bold, 1975 Van den Bold, 1975 Van den Bold, 1972a Van den Bold, 1963a,b Carren˜o, 1985 Echevarrı´a, 1988 Szczechura and Blaszyk, 1996 Van den Bold, 1957, 1963b Van den Bold, 1972a Bertels et al., 1982 Bertels, 1975 Teeter, 1975 Van den Bold, 1975 Van den Bold, 1964 Van den Bold, 1972a Van den Bold, 1963b Van den Bold, 1966d Coimbra et al., 1992

Region

Cuba Cuba Venezuela Venezuela Mexico Argentina Antarctica Trinidad Venezuela Brazil Argentina Belize Cuba Venezuela Venezuela Trinidad Panama North Brazil– Pernambuco Chukewsky and Purper, 1985a,b North Brazil Coimbra and Ornellas, 1986, 1987, 1989 Coimbra et al., 1992 Medeiros and Coimbra, 1989 Ornellas and Coimbra, 1985 Purper and Ornellas, 1987a,b Coimbra et al., 1994 Brazil — Rio de Janeiro Dias-Brito et al., 1988 Ramos et al., 1999 Vicalvi et al., 1977 Brazil–Sao Paulo Kotzian and Eilert, 1985 Brazil Coimbra et al., 1995 South Brazil Purper and Ornellas, 1989 Ramos, 1994, 1996 Whatley et al., 1987, 1988, South Brazil 1998a,c Whatley et al., 1998a,b,c Brazil=Argentina Bertels and Martinez, 1990 Argentina Whatley et al., 1998b,c Argentina=42–40ºS Whatley et al., 1998b,c Argentina=52–47ºS Whatley et al., 1998b,c Argentina=Ushuaia Whatley et al., 1995 Falkland Island Whatley et al., 1996 Magellan Straits Benson, 1964 Antacrtica Mu¨ller, 1908 Antarctica Neale, 1967 Antarctica Hartmann, 1988 Antarc.=Coronation Island Hartmann, 1989 Antarc.=South Georgia Hartmann, 1992 Antarc.=Elephant Island Hartmann, 1993 Antarc.=Isla de los Estados Hartmann, 1992 Antarctica=Halbinsel Whatley et al., 1999 Antarctica Hartmann-Schro¨der and Chile Hartmann, 1962, 1965

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Appendix B. Presence and absence data for Oligocene to Recent genera from the Neotropics to Antarctica Primary data sources are supplied in Appendix A.

Appendix C. Jaccard Coefficient similarity matrices for the Oligocene, Early Miocene, Late Miocene and Pliocene Primary data sources are supplied in Appendix A.

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