Tropical Plant Disjunctions: A Personal Reflection Author(s): Robert Thorne Reviewed work(s): Source: International Journal of Plant Sciences,

Vol. 165, No. S4, Tropical Intercontinental Disjunctions (July 2004), pp. S137-S138 Published by: The University of Chicago Press Stable URL: http://www.jstor.org/stable/10.1086/424023 . Accessed: 27/11/2011 14:51
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Int. J. Plant Sci. 165(4 Suppl.):S137S138. 2004. 2004 by The University of Chicago. All rights reserved. 1058-5893/2004/1650S4-0010$15.00

TROPICAL PLANT DISJUNCTIONS: A PERSONAL REFLECTION

Robert Thorne

It is most gratifying to this biogeographer to see in the contributions to this volume that the zealotry of the recent vicariance biogeographers has returned to normalcy with the current acceptance of vicariance, long-distance dispersal, and immigration from the boreotropics as possible explanations of intercontinental disjunctions. The refusal of the vicariantists to accept long-distance dispersal and oceanic islands was only a passing phase due to their limited knowledge of vagility in groups with which they were unfamiliar and failure to consider a time frame (Thorne 1996). Increasing knowledge of fossil distribution plus the use of molecular data, including molecular clocks (S. Magallon, in this issue), has added greatly to our ability to understand intercontinental disjunctions. Vicariance explanations have long been accepted by biogeographers, more especially since the acceptance of tectonic plate movement, i.e., continental drift (Thorne 1978). Good examples of such vicariance can be found among the subcosmopolitan orders and families like the Myrtales and Myrtaceae (Conti et al. 2002; Schonenberger and Conti 2003; F. Rutschmann et al.; K. J. Sytsma et al.). The largely South African Penaeaceae, Oliniaceae, and Rhynchocalycaceae and the currently tropical American Alzateaceae are surely closely related, with the intercontinental disjunction presumably dating back before the breakup of West Gondwana or before the gap between the two continents became too great (Schonenberger and Conti 2003). The southern and southeastern Asian Crypteroniaceae (F. Rutschmann et al.), sister to the Penaeaceae, Oliniaceae, Rhynchocalycaceae, and Alzateaceae, are suggested through molecular dating as having split from their West Gondwanan relatives in Early to Middle Cretaceous time and having reached Asia by rafting on the Indian Deccan Plate. Memecylon of the Memecylaceae, also of the Myrtales, likewise may have traveled from Africa to southeastern Asia via the Indian Ark, and possibly too the Kibessieae of the sister group Melastomataceae (Morley and Dick 2003). The more distantly related Myrtaceae of the same order appear to have an Australasian origin and probably reached South America before the link between Australasia-Antarctica-South America was broken (K. J. Sytsma et al.). The pantropical magnolialian Annonaceae and Myristicaceae (J. A. Doyle et al.) may also have a vicariant origin. The Annonaceae may have radiated in South America and Africa in Late Cretaceous time when the South Atlantic was a narrower ocean. Doyle et al. also suggest that several annonaceous lines dispersed from Africa-Madagascar into Laurasia when the Tethys Sea closed in the Tertiary. The history of the Myristicaceae appears more recent, but the family is relatively primitive, and the large, animal-dispersed seeds are not adapted to transoceanic dispersal. Other magnolialian famiS137

lies with widely disjunct ranges like the Winteraceae, Canellaceae, Illiciaceae, Calycanthaceae, and Lauraceae (Thorne 2000) have ancient histories and probably have vicariant origins, as do other of the pantropical angiosperm families. On the other hand, numerous genera and species with intercontinental disjunctions are best explained by longdistance dispersal over land and over sea (Thorne 1963, 1972, 1973, 1996; Les et al. 2003; T. J. Givnish and S. S. Renner; S. S. Renner). Renner, in her chapter on plant dispersal across the tropical Atlantic by wind and sea currents, presents an updated list of 110 disjunct tropical Atlantic genera. Almost all of their disjunctions are presumed to be too recent to be explained by tectonic plate displacement. TransAtlantic dispersal by water in both directions appears more likely than dispersal on birds or by wind. T. J. Givnish et al. studied two tropical American commelinid monocot families, Bromeliaceae and Rapateaceae, that have single species in a sandstone area in West Africa. Although that sandstone area abutted the Guayana Shield of northern South America before the two continents rifted, molecular dating suggests that the rapatead Maschalocephalus dinklagei Gilg. & K. Schumann reached West Africa by long-distance dispersal 7 m.yr. ago and the bromeliad Pitcairnia feliciana (A. Chevalier) Harms & Milbraed may have reached West Africa similarly 15 m.yr. ago. Another American family Cactaceae has a single genus, Rhipsalis, possibly only the single species, R. baccifera (J. Miller) W. T. Stearn, that has reached Africa and beyond to Madagascar and north to Sri Lanka. Its succulent white berries with viscous pulp are obviously specialized for bird dispersal (Thorne 1973, 1996). Aquatic owering plants are notorious for their often subcosmopolitan distributions. In a recent paper, Les et al. (2003), using molecular estimates of divergence time, found in most instances times far too recent to implicate tectonic plate displacement as the explanation for the disjunctions. Les et al. suggest long-distance dispersal by birds, involving both overland and transoceanic migratory routes. Similarly, the rather frequent western North Americantemperate South American disjunctions can be best explained by longdistance dispersals by water- or shorebirds (Raven 1963; Thorne 1970, 1978, 1996). In studying bicontinental disjunctions in the woody Boraginales, M. Gottschling et al. found from calculated ages that plate tectonics seem to play a minor role in the distribution patterns of the Cordiaceae, Ehretiaceae, and Heliotropiaceae, often included as subfamilies in the Boraginaceae. In most of the wide disjunctions of these woody borages, from their initial diversication in South America, rare events of longdistance dispersal seem the best explanation. Endozoochorous dispersal of drupes by birds and small mammals and hydrochoria via corky endocarps or air-lled sterile chambers in

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INTERNATIONAL JOURNAL OF PLANT SCIENCES latitude land connections when northern forests supported tropical vegetation. Increasing knowledge of the vagility of groups, of their fossil distributions, and of their temporal unfolding based on molecular clocks over the next years should further contribute to a balanced view of the causes of intercontinental disjunctions.

fruits of coastal species presumably are effective in sporadic events of long-distance dispersal. C. C. Davis et al., nally, discuss the third possible explanation for tropical south-Atlantic intercontinental disjunctions, the boreotropical hypothesis, which attributes some American-African tropical disjunctions to Tertiary high-

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