Explanations for biogeographic disjunctions involving SouthAmerica and Africa typically invoke vicariance of western Gondwanan biotas or long distance dispersal. These hypotheses are problematical because many groups originated and diversified well after the last known connection between Africa and South America (Ϸ105 million years ago), and it is unlikely that ''sweepstakes'' dispersal accounts for many of these disjunctions. Fig. 1). Approximately 85% of the species occur in the New World where they are pollinated by specialized oil-collecting bees that are absent from the Old World. This distribution has invited the development of alternative theories for the origin and diversification of Malpighiaceae. Vogel (2) proposed the ''Gondwanian aborigine'' hypothesis, in which the current distribution resulted from the break-up of western Gondwana (the supercontinent comprising Africa and South America). This hypothesis implies that the Malpighiaceae originated before western Gondwana divided [i.e., Ϸ105 mya (3)], as did several included lineages with disjunct distributions between the New and Old Worlds. In contrast, Anderson (1) and others (4) have favored an ''American colonist'' hypothesis, according to which Malpighiaceae originated in northern South America, in isolation from Africa, and dispersed eastward across the Atlantic at least twice. This hypothesis predicts that the ages of Malpighiaceae and of divergences between New World and Old World lineages are younger than the last known land connection between South America and Africa.To elucidate the biogeographic history of Malpighiaceae, we used maximum likelihood to estimate the phylogeny for the group by using DNA nucleotide data from chloroplast ndhF and nuclear phytochrome C (PHYC) sequences (see Materials and Methods). This data set included sequences from one outgroup and 70 species of Malpighiaceae. To test these biogeographic hypotheses, it is necessary to attach a temporal dimension to the phylogeny to infer the timing (i) of the origin of Malpighiaceae and (ii) of the disjunctions between New and Old World lineages. Although molecular divergence estimates must be viewed with caution, these approaches may identify a window of time for branching events that will help select among competing hypotheses.
Materials and MethodsOur data set included 71 ndhF sequences and 71 PHYC sequences, which were readily aligned by eye. This data set included sequences from one outgroup and 70 species of Malpighiaceae representing the majority of genera, including multiple accessions from morphologically diverse or putatively non-monophyletic genera. Most ndhF sequences were previously obtained by using the methodology published in Davis, Anderson, and Donoghue (5) and in Davis (6). Two additional ndhF sequences were also generated for this study (GenBank accession nos. AF500495 and AF500496). The PHYC sequences are newly generated (GenBank accession nos. AF500522-AF500582), except for AF436794 -AF436804 from Davis (6), and were obtained by using previously detaile...