Pteronotus parnellii, (Gray, 1843)

DÁVALOS, LILIANA M., 2006, The geography of diversification in the mormoopids (Chiroptera: Mormoopidae), Biological Journal of the Linnean Society 88 (1), pp. 101-118 : 111

The P. parnellii lineage (subgenus Phyllodia)

The molecular phylogeny challenges the biogeographical hypothesis of Smith (1972) on the single, Middle American origin of Antillean populations. Lewis Oritt et al. (2001) first proposed northern South America as the ancestral area of Antillean Phyllodia , but this result is not significantly different from the traditional biogeographical explanation [except when using the Templeton (1983) test; Table 5 ]. The phylogeny of Figure 5 is the first to suggest that P. parnellii is not sister to a clade containing P. pusillus . Because both trees ( Figs 3B , 5 ) are equally good at explaining the data (P = 0.352, Shimodaira–Hasegawa test), Phyllodia might have reached the Caribbean or the continent more than once. Two Phyllodia species have been recorded as Quaternary fossils on Hispaniola ( Morgan, 2001; Table 1 ); the extant pusillus and sp. cf. rubiginosus ; perhaps corresponding to separate waves of colonization from the continent and/or adjacent islands.

Whether bats in this lineage first arose on the islands or the continent cannot be established because both regions are optimized in the ancestral area (not shown). Furthermore, the low support values ( Figs 3B , 5 ) mean that parnellii , pusillus and portoricensis , rubiginosus , or ‘ rubiginosus ’ from Suriname and French Guiana could each be the oldest branch within the lineage, adding uncertainty to the geographical origin of the subgenus. The Caribbean-continent divergence within this lineage is as great as that between Surinamese and Guianan ‘ rubiginosus ’ and sister clade ( Fig. 6A ). By contrast, sequence divergence within the widespread rubiginosus clade is significantly lower ( Fig. 6A ), as expected if the expansion to Middle America or north-western South America had happened recently.

Taken together, the results imply that the geographical history of these bats is more complex than proposed hitherto ( Smith, 1972), and suggest avenues for future research. First, more rapidly evolving characters are needed to resolve relationships among the species in this subgenus ( Figs 3 , 5 ). Second, geographical sampling must include the entire range of Phyllodia because apparently continuous populations show deep divergences that might represent additional independently evolving lineages (e.g. in northern South America). Third, such studies should include morphological characters because the Caribbean Pteronotus pristinus ( Silva-Taboada, 1974; Simmons & Conway, 2001) and Pteronotus sp. cf. rubiginosus ( Morgan, 2001) are only known as fossils and might provide fresh insights into the history of exchange between the continental and insular Neotropics.

Figures

Figure 3. A, strict consensus of eight most parsimonious cladograms resulting from analysis of cytochrome b (L = 1792 steps, consistency index = 0.439, retention index = 0.775). Numbers below branches are Bremer support values, above branches are percent of 1000 jackknife replicates. Names of outgroups are in bold; for sequence data, see Appendix. B, phylogram resulting from maximum likelihood analysis using a rate-constant GTR+I+Γ model of DNA evolution (–lnL = 9181.23). Numbers above or below branches are percent of 300 50% jackknife replicates, thicker lines indicate 100% jackknife support.

Figure 5. Majority rule (50%) consensus of 19 000 cladograms resulting from Bayesian analysis of concatenated molecular data for all diagnosable mormoopid taxa (– lnL = 24 910; 95% confidence interval = 24,890–24 920). Dashed branches had posterior probabilities between 0.50 and 0.95. All other branches had posterior probabilities between 0.95 and 1. Names of outgroups are in bold; for sequence data, see Appendix. The top panel shows the ancestral area inferred for branch 1, the bottom panel shows the ancestral area of branches 2 and 3. DIVA Optimizations were constrained to a maximum of two areas, and all solutions are shown. Three alternatives to the polytomy of Pteronotus davyi and Pteronotus gymnonotus, two alternatives to the sister of Pteronotus quadridens and macleayii (davyi and gymnonotus, or personatus), and two taxonomies (the traditional species taxonomy of Smith (1972), or that shown in Figure 3 were analysed, and all result in the same composite estimates. Geographic distributions are as shown in Table 1. Pteronotus pristinus and Mormoops magna were not analysed. FG, French Guiana; Hon., Honduras; Mex., Mexico; PR, Puerto Rico; Ven., Venezuela.

Figure 6. Confidence intervals around observed sequence divergence resulting from parametric bootstrapping of rate-constant mormoopid phylogenies. A, estimates of divergence for mitochondrial ribosomal DNA (black diamonds) and the cytochrome b gene (white diamonds). B, estimates of divergence for nuclear Rag2. CA, Central America; FG, French Guiana; Mex., Mexico.