taxonID	type	description	language	source
6C5E879A3848FFAED9CBF89AFC3EFE07.taxon	description	The species diversity of the mormoopids has been underestimated. Two hypotheses underlie competing assessments of species diversity. The first hypothesis assumes populations of widespread species as part of a continuum of differentiation that appears great at the extremes, but is only slight between adjacent groups (Koopman, 1955). Although some populations are allopatric, it is assumed that gene flow among them exists or occurred until recently. Because morphological intergradation (used to infer gene flow) among insular and continental populations is not observed, the range of (non) resemblance permitted in a given species has been widened (Smith, 1972).. The assessment of sequence variation among putative units (subspecies) within the mormoopids (Table 2) revealed multiple instances of characters that appear to be fixed in cytochrome b. These molecular data support a second hypothesis: gene flow between insular and continental populations appears to have ceased even before fully recognized biological species (e. g. P. gymnonotus and davyi; Fig. 3 B) evolved into separate lineages. The subtle morphological differences dismissed under a presumption of gene flow provide evidence for the isolation and independent evolution of separate lineages in widespread species such as parnellii, davyi, and personatus. Because sampling sizes for molecular markers were small, these differences alone cannot provide species limits. In some instances, molecular character differences, high sequence divergence among presumed conspecifics (bottom two tiers of Fig. 2), distributional ranges that encompass broad areas separated by water and land barriers (Table 1, Fig. 1), and taxonomic limits based on morphological variation (Smith, 1972) coincide and strengthen the hypothesis of evolutionary independence. These criteria (Tables 1 and 2) apply to named island populations of P. parnellii: parnellii (Gray, 1843), pusillus (Allen, 1917), and portoricensis (Miller, 1902); the continental P. parnellii ranging from Mexico to Guyana currently classified in the subspecies mexicanus (Miller, 1902), mesoamericanus (Smith, 1972), and rubiginosus (Wagner, 1843); the currently recognized subspecies of P. davyi: davyi (Gray, 1838) and fulvus (Thomas, 1892); and subspecies of P. personatus: personatus (Wagner, 1843) and psilotis (Dobson, 1878). Each of these populations should be considered as a species, named using the subspecies taxonomy. The name Pteronotus rubiginosus (Wagner, 1843) precedes mexicanus and mesoamericanus, and applies to the continental bats in the P. parnellii lineage as described above (note that the status of fuscus and paraguanensis was not evaluated; Table 1). Both cytochrome b (Table 2) and Rag 2 (Lewis Oritt et al., 2001) showed differentiation in Mexican and Central American populations of P. psilotis. Further sampling is necessary to determine if characters are fixed because taxonomic conclusions derived from single molecular exemplars would be suspect. Mitochondrial cytochrome b from samples of Pteronotus rubiginosus and P. personatus from northern South America west of Guyana is distinct from that sampled east of Guyana (Suriname and / or French Guiana; Table 2, Fig. 1). These character differences and attendant levels of sequence divergence had not been anticipated in the morphological study of Smith (1972). French Guianan specimens of P. parnellii can also be distinguished from those from the remainder of the range by their larger size (Simmons & Voss, 1998). In Venezuela, P. paraguanensis appears to have become isolated as a result of breaks in the humid forest (Gutiérrez, 2004). This mechanism might explain the differentiation observed, but greater geographical and character sampling is needed to investigate these (possibly) cryptic species, and test the possibility that accelerated rates of sequence evolution have led to this pattern (although this is unlikely, see Table 4). In P. quadridens discontinuous variation in cytochrome b occurs between Cuba and Jamaica, and Hispaniola and Puerto Rico, rather than coinciding with the subspecies taxonomy that separates Cuban from other Greater Antillean bats (Tables 1, 2). These taxa are not elevated to species here, despite the possible geographical isolation by ocean barriers, because sampling was sparse, the molecular differentiation does not match subspecies boundaries based on morphology, and no differences were detected in Rag 2. For the purpose of estimating ancestral areas, each terminal that appears with a name in Figure 3 was treated as a separate taxon.	en	DÁVALOS, LILIANA M. (2006): The geography of diversification in the mormoopids (Chiroptera: Mormoopidae). Biological Journal of the Linnean Society 88 (1): 101-118, DOI: 10.1111/j.1095-8312.2006.00605.x, URL: https://academic.oup.com/biolinnean/article-lookup/doi/10.1111/j.1095-8312.2006.00605.x
6C5E879A384AFFAFD967FDBDFD34F9C9.taxon	description	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 3 B, 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 3 B, 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. 6 A). By contrast, sequence divergence within the widespread rubiginosus clade is significantly lower (Fig. 6 A), 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.	en	DÁVALOS, LILIANA M. (2006): The geography of diversification in the mormoopids (Chiroptera: Mormoopidae). Biological Journal of the Linnean Society 88 (1): 101-118, DOI: 10.1111/j.1095-8312.2006.00605.x, URL: https://academic.oup.com/biolinnean/article-lookup/doi/10.1111/j.1095-8312.2006.00605.x
6C5E879A384BFFA8DAE5F9A9FD7EF8A6.taxon	description	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).	en	DÁVALOS, LILIANA M. (2006): The geography of diversification in the mormoopids (Chiroptera: Mormoopidae). Biological Journal of the Linnean Society 88 (1): 101-118, DOI: 10.1111/j.1095-8312.2006.00605.x, URL: https://academic.oup.com/biolinnean/article-lookup/doi/10.1111/j.1095-8312.2006.00605.x
6C5E879A384CFFA9D967FF4AFC68FDE3.taxon	description	The estimated ancestral area of Mormoops and the mormoopids (Fig. 5) encompasses both northern South America (Smith, 1972) and the Greater Antilles (Czaplewski & Morgan, 2003). The two biogeographical hypotheses are not mutually exclusive, and it is plausible that the most recent common ancestor of mormoopids was widespread from Mexico south to northern South America, and east to the Greater Antilles. Another interpretation of this result is that dispersal-vicariance analysis is inconclusive, and other sources of evidence are needed to clarify the geographical history of mormoopids. There are several reasons to doubt that the ancestor of Mormoops was as widespread as estimated in Figure 5. First, extant Mormoops species do not overlap on the continent, but both are known from the Greater Antilles (albeit, one only as fossil). Second, one additional species, Mormoops magna, is known from late Pleistocene remains on Cuba (Silva-Taboada, 1974), adding a third Mormoops lineage to the Greater Antilles. Third, it is parsimonious to postulate that the ancestor of Mormoops reached the Greater Antilles before splitting into the extant species but, even if it did not, the divergence between the Antillean blainvillei and its sister taxon is significantly greater than that between megalophylla populations (Fig. 6 A). The combination of species diversity and depth of divergence suggests Mormoops expanded its range from north to south. If Mormoops ranged into the Greater Antilles even before blainvillei and megalophylla differentiated, Caribbean colonization in this family can be traced back to the divergence between the mormoopid genera, and might be as ancient as the Oligocene or Miocene (Czaplewski & Morgan, 2003). A northern neotropical (and perhaps insular) origin for the genus can be overturned by the discovery of a basal Mormoops species in South America. An extensive fossil record shows that M. megalophylla ranged from Florida through the Greater Antilles to Bahia in Brazil during the Late Pleistocene (Ray, Olsen & Gut, 1963; Silva-Taboada, 1974; Czaplewski & Cartelle, 1998). Studies of morphological variation are necessary to determine the relationships among extant and fossil megalophylla populations and test the hypothesis presented here because more than one species might be involved (Morgan, 2001). One prediction following Czaplewski & Morgan’s (2003) biogeographical model is borne by the molecular data: divergences between Antillean and continental mormoopids are greater than those between Central American and northern South American populations (Fig. 6). There is only one exception in the P. parnellii lineage (subgenus Phyllodia), where two northern South American populations might not share a most recent common ancestor (Figs 3, 5). For every other mormoopid lineage, and even in one instance within Phyllodia, the divergence between Mexico / Central America and South America appears to be recent (Fig. 6), and might correspond to the completion of the Isthmus of Panama in the late Pliocene. Either Mexico / Central America or north-western South America was recently colonized by all mormoopid lineages. As discussed above, the direction of this expansion appears to be from north to south in Mormoops and P. personatus s. l., but the evidence is ambiguous for Phyllodia, as well as for P. davyi and gymnonotus. Because Mormoops is at the base of the mormoopid radiation, restricting its ancestral distribution to the northern Neotropics constrains the geographical origin of the family to that region. Other than differences in branch length (longer for northern neotropical splits, shorter for divergences between Mexico / Central America and South America), the fossil record also supports a north-to-south expansion. The oldest mormoopid diverged before the two extant genera (G. Morgan, pers. comm.), and ranged into Florida in the Oligocene (Czaplewski, Morgan & Naeher, 2003). In general, mormoopids appear to have reached South America late in their history, after diversifying in Mexico, Central America, and / or the Greater Antilles (Fig. 6). This finding is critical to the biogeographical history of noctilionoids. Both morphology (Simmons & Conway, 2001) and large concatenated molecular datasets (Teeling et al., 2005) indicate that mormoopids and phyllostomids are each other’s closest relative (this topology was not always recovered in this study, probably because taxon sampling among bat families was poor relative to the higher-level analyses cited above). Two phylogenetic hypotheses have been proposed to explain relationships among phyllostomids. One, based on analyses of mostly morphological data (Wetterer, Rockman & Simmons, 2000) identified the vampires (Desmodus, Diaemus, and Diphylla) as the oldest phyllostomid lineage. A second hypothesis based on mtrDNA and Rag 2 (Baker, Porter, Hoofer & Van Den Bussche, 2003) suggests that Macrotus diverged before any other phyllostomid. The geographical distribution of the basal lineage of the phyllostomids would have a disproportionate effect on ancestral area reconstructions for that family. Vampires range from Mexico to Chile and Uruguay, and fossils have been found on Cuba (Koopman, 1994). This lineage would not constrain the ancestral area of the phyllostomids because of its widespread distribution. Since the greatest diversity of phyllostomids is concentrated in northern South America and the vampires include it in their range, this would likely be the most parsimonious ancestral area for the family. By contrast, Macrotus is only known from the southwestern United States south to Guatemala, through the Greater Antilles and Bahamas (Koopman, 1994). If Macrotus is at the base of the phyllostomid radiation, then the ancestral distributions of mormoopids and phyllostomids were adjacent in the northernmost Neotropics. Phyllostomid fossils are known from the middle Miocene of La Venta (Czaplewski, 1997), indicating phyllostomids reached South America early in their history. The geographical distribution of these closely related families during their early history might help explain the remarkable differences in taxonomic and adaptive diversity between the two groups.	en	DÁVALOS, LILIANA M. (2006): The geography of diversification in the mormoopids (Chiroptera: Mormoopidae). Biological Journal of the Linnean Society 88 (1): 101-118, DOI: 10.1111/j.1095-8312.2006.00605.x, URL: https://academic.oup.com/biolinnean/article-lookup/doi/10.1111/j.1095-8312.2006.00605.x
