Pteronotus, Gray, 1838

DÁVALOS, LILIANA M., 2006, The geography of diversification in the mormoopids (Chiroptera: Mormoopidae), Biological Journal of the Linnean Society 88 (1), pp. 101-118 : 112

Other Antillean Pteronotus

Phylogenetic analyses including morphological data support the hypothesis of Smith (1972), whereby the P. macleayii and quadridens clade is sister to P. personatus s.l. ( Table 3 ), while analyses of Rag 2 result in the resolution of Figure 4 . The difference between alternatives is not significant ( Table 5 ), despite the posterior probability of 1.0 obtained for the latter resolution ( Fig. 5 ). Regardless, the phylogenies optimize the distribution of the ancestral lineage to include the western Greater Antilles ( Fig. 5 ). If distance between areas were an indication, Mexico and/or Central America would be the likely continental source ( Fig. 1 ). Near-interconnections between the Antilles and Middle America during periods of low sea level might have facilitated dispersal ( Smith, 1972; Griffiths & Klingener, 1988).

The biogeographical analysis, however, inferred northern South America as part of the ancestral area ( Fig. 5 ). Each of the plausible sisters to P. macleayii and quadridens contains both Middle American [ fulvus and perhaps davyi and gymnonotus ( Table 1 ), or the two paraphyletic lineages in psilotis ( Fig. 3 )], and South American lineages ( davyi and gymnonotus , or personatus ). Divergences between these continental populations are often significantly smaller than between P. macleayii and quadridens and its continental sister ( Fig. 6 ), suggesting dispersal to the Caribbean preceded range expansion within the continent by far (Czaplewski & Morgan, 2003). The direction of this expansion from north to south is in agreement with a Mexican and/or Central American origin for personatus s.l., and for the macleayii and quadridens clade if the two lineages were sister. The lack of resolution among fulvus , davyi , and gymnonotus precludes a firm conclusion but, if these three species all range into Middle America, then north-to-south range expansion would become parsimonious for this clade and its sister, despite the polytomy. This last question remains to be resolved because neither the northernmost range of davyi and gymnonotus , nor P. davyi incae , was sampled in molecular analyses (cf. Table 1 , Appendix).

Figures

Figure 1. Map of the Caribbean and biogeographical hypotheses about the origin of mormoopids. According to Smith (1972), ancestral mormoopids dispersed from northern South America or southern Central America to Mexico/ Central America. From there, the ancestors of Greater Antillean mormoopids reached the West Indies through Cuba via Yucatán, or Jamaica via Honduras. Dispersal through these routes would explain the distribution of the single lineage comprising Pteronotus quadridens and macleayii (ancient), the species Mormoops blainvillei (less ancient), and the Caribbean populations of Pteronotus parnellii (most recent). Czaplewski & Morgan (2003) concur on the dispersal routes to the Caribbean, but propose that mormoopids colonized the islands early in their evolutionary history. From Mexico/Central America, mormoopids would have reached South America recently, after the closing of the Isthmus of Panama.

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 4. Cladogram resulting from maximum likelihood analysis of concatenated mitochondrial ribosomal DNA, cytochrome b, and Rag2 sequences (– lnL = 24736.88). Sequences for at least two of the molecular partitions were available for each terminals. Support values are shown for mtrDNA in the first column, Rag2 in the second column, concatenated molecular sequences in the third column, and total evidence in the fourth column. The top row shows Bremer support indices; second row is the percent of 1000 50% jackknife pseudoreplicates using parsimony; and the third row is the percent of 300 50% jackknife pseudoreplicates using maximum likelihood. Asterisks indicate jackknife support values of 100%. Dashes indicate that the data set does not resolve the branch, or does not support the resolution shown (Table 3). NA, not applicable; indicating the partition contains data for only one terminal and the branch could not be scored, or no maximum likelihood analysis including morphological data was performed. Numbers in grey indicate that the data partition does not include all terminals in the branch. Models of sequence evolution used to analyse each partition and resulting loglikelihood values are shown in Table 4. Names of outgroups are in bold; for sequence data, see Appendix.

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.