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Rafting Mammals Or Drifting Islands?: Biogeography of the Greater Antillean Insectivores Nesophontes and Solenodon

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Rafting Mammals Or Drifting Islands?: Biogeography of the Greater Antillean Insectivores Nesophontes and Solenodon Journalof Biogeography (1 980) 7, 11-22 Raftingmammals or driftingislands?: biogeographyof theGreater Antillean insectivores Nesophontesand Solenodon BRUCE J.MAC FADDEN FloridaState Museum, University of Florida, Gainesville,Florida 32611, U.S.A. ABSTRACT.During the late Mesozoicand earlyCenozoic, certain soricomorph insectivoreswere distributed throughout North America, Nuclear Central America, and the proto-Antilles.A regionaltectonic reorganization during this time resultedin the decouplingof Caribbeanlithosphere from surrounding plates and movementof the Greater(proto-) Antilles eastward relative to the Americas. This movementresulted in biogeographicsubdivision (vicariance sensu Rosen, 1975) of the ancestralsoricomorph distribution. As far as we know, the Americanpart of the distribution,represented by apternodontidsand possibly geolabidids,became extinct by thelate Oligocene.The GreaterAntillean part of the distribution,represented by Nesophontesand Solenodon,has persisted throughto the Recent.In short,these Caribbean insectivores are islandrelicts of a once morewidespread ancestral distribution of soricomorphinsectivores. Similardistributional evidence from numerous other elements of theCaribbean biota supportthe biogeographichypothesis proposed forNesophontes and Solenodon. 'I do not deny that there are many and grave the absence of 'top' carnivores.Phylogenetic- difficultiesin understandinghow several of ally primitiveforms, or 'relicts,' tend to be the inhabitants of the more remote islands, preserved in island biotas. In recent years whetherstill retainingthe same specific form much work has been done on quantitative or modified since their arrival, could have descriptionsof island biotas (e.g. MacArthur, reached theirpresent homes.' 1972; Simberloff, 1974). Hypotheses that Charles Darwin (1859, p. 396) explain the origin of these biotas from an external source have been assumed to be adequate when dealing with easily dispersed forms such as plants and mobile animals, but Introduction fall short when considering immobile and exclusivelyterrestrial animals. Biogeographers have long recognized the The Greater Antilles possess a peculiar peculiarities of island biotas. Darwin (1859) assemblage of relict insectivoresthat include discussed this problem as seen in the Gala- the genera Nesophontes and Solenodon. pagos. It is known that islandstend to exhibit Workers such as Matthew (1939), Simpson a high degree of endemism. Biotic diversity (1956) and Patterson (1962) have tradition- is lower on islands than on mainland regions ally explained the presence of these insecti- with comparable areas. Frequently islands vores in the Caribbean by sweepstakes, or show an apparent ecological imbalance of overwater dispersal from the Americas. trophic group structure,for example, with Recently, Rosen (1975) hypothesized that 0305-0270/80/0300-0011$02.00 ? 1980 BlackwellScientific Publications 11 12 Bruce J. MacFadden virtuallythe entireCaribbean biota, including However, when dealing with groups with Nesophontes and Solenodon, was biogeo- relatively poor fossil records, one is never graphicallyseparated (by vicariance) froman sure if the oldest representativesare in fact ancestral American biota as a result of plate closest to the centre of origin. Centres of tectonics in that region. The purpose of this origin and subsequent dispersal events are paper is to extend Rosen's study and focus hypothesized with some confidence in geo- on the biogeographic origin of Nesophontes chronologically controlled sequences for and Solenodon throughexamination of rele- certain abundant groups, such as marine vant phylogeneticand geologicalinformation. planktonic microfossils and late Cenozoic mammals. The problem remains as to when a fossil group is well enough sampled and Models of vertebratebiogeography supported by enough temporal data so that discussions of biogeography are meaningful. Matthew (1939) and Simpson (1940, 1952, The solution to this dilemma is certainly and numerous other references)were instru- elusive. mental in the formulationof a widely used Croizat, Nelson & Rosen (1974) summa- biogeographicmodel for terrestrialmammals. rized the 'vicariance' model by which syste- Simpson's model includes the concept of matists can reconstruct the biogeographic sweepstakes, filter, and corridor dispersal history of monophyleticgroups by analysing routes. The 'traditional (Matthew-Simpson) cladistic interrelationshipsof constituenttaxa dispersal' model for mammals was founded (also see more recent discussion by Rosen, on the assumption that the relativepositions 1978, and Platnick& Nelson, 1978). Ancestral of continents have remained stable through- geographic distributions of taxa may be out geological time. Recent discussions of separated by the formation of barriers vertebratebiogeography that modifythe ideas (vicariance). The separation of ancestral of Matthew and Simpson in the light of distributions results in evolutionary diver- plate tectonics have been presented by gence (e.g. allopatric speciation) of the taxa workers such as McKenna (1972, 1973), involved. This process of biogeographicsub- Hallam (1974), Cracraft (1974) and Cox division could have occurred numeroustimes, (1974). Many of these studies incorporate resulting in repeated events of speciation. the recognitionof centresof originand subse- Withinthe groups being studied, more recent quent direction of dispersal for faunas or biogeographic subdivisions (and allopatric particularfaunal constituents. speciations) would result in more closely Centres of origin can be recognized for related species (or sister) groups. Evidence taxa that have undergone dispersal during from other groups of organismswith similar historical times, e.g. the introduction of phylogeneticand biogeographichistories could starlings (Sturnus vulgaris) into North be used to support an hypothesis of the America (Krebs, 1972), rabbits (Oryetolagus biogeographic subdivision events. Vicariance cuniculus), into Australia (Ricklefs, 1974), has been used to explain the biogeography and the northward spread of the nine- of spiders (Platnick, 1976), corals, seagrasses banded armadillo (Dasypus novemcinctus) and mangroves(McCoy & Heck, 1976), and into southern North America (Humphrey, parts of the Caribbean biota, particularly 1974). In prehistoricaltimes, recognitionof fishes(Rosen, 1975). centres of origin becomes somewhat more The vicariance model is not a panacea for complex. Some biogeographersworking with interpretingall biotic distributions; but it wholly extant taxa state that, withina mono- does provide an elegant explanation for phyletic group, the most primitivetaxon will organismswhose biogeographyhas been pro- be closest to the centreof origin.Others state foundly influenced by barriers,such as those that the most derivedtaxon will be closest to produced by plate tectonics.Despite claims to the centreof origin.Dispersal hypotheses used the contrary, vicariance is not generally by palaeontologists usually assume that the applicable to many relativelyrecent biogeo- older fossil record of a group in one area graphic subdivisions.A complex biota might indicates a closeness to the centre of origin. have been affected by several major biogeo- Antillean insectivorebiogeography 13 graphicevents which producedits present- The insectivoresinclude the generaNeso- day taxonomic composition.As McDowall phontes and Solenodon. Nesophontes consists (1978) has shown,the biogeographichistory of at least six species foundpredominantly of a complex biota could combine both in Quaternary cave deposits on Cuba, vicarianceand traditionaldispersal models. Hispaniola, Puerto Rico, and smallersur- roundingislands (Allen, 1918; Koopman & Ruibal, 1955; McDowell, 1958).Nesophontes Antilleaninsectivores has beenreported to be extantuntil as recently as the 1930s. A possibilityexists that this Knowledgeof the Antilleanbiota has greatly genusstill may be extant.During the begin- increasedsince the turnof the centuryas a ningof thiscentury, some naturalistsclaimed result of numerousnatural history expedi- that Solenodon had become extincton His- tions to this region.Classic papers dealing paniola. Verrill(1907, p. 55) stated that: withthe fossiland extantmammals collected 'For manyyears it has been commonlycon- duringthe early expeditionswere published sidered extinct, and when, in December, by G. M. Allen (e.g. 1911, 1918) and 1906, I undertooka collectingtrip to San Anthony(e.g. 1918). Particularlycentral to Domingo with the avowed intention of the presentdiscussion, Matthew (1939), in obtainingSolenodon, prominentzoologists his Climate and evolutdon,devoted an entire stated that the quest was hopeless,one of chapter to the biogeographyof Antillean themsaying that I wouldbe as likelyto secure mammals.In recentyears numerous Antillean specimensof ghosts as of Solenodon para- studies have been presentedon Quaternary doxus.' Solenodon is representedby at least cave faunas(e.g. Koopman& Ruibal, 1955; two extantand severalextinct species from Varona & Garrido,1970), generalsurvey of Cuba and Hispaniola (McDowell, 1958). mammals(Varona, 1974), bats (e.g. Silva- Patterson(1962) named a new solenodon, Taboada & Koopman, 1964), primates Antillogale,from Quaternarycave deposits (Williams& Koopman, 1952; Hershkovitz, in Hispaniola.Van Valen (1967) questioned 1970), rodents(e.g. Varona,1970), edentates the genericvalidity of Patterson'sAntillogale; (e.g. Matthew& de PaulaCouto, 1959) insecti- Solendon is used here in a broad sense to vores(e.g.
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