Historical biogeography and distribution of the freshwater calanoid copepods (Crustacea: Copepoda) of the Yucatan Peninsula, Mexico

Aim To determine and analyse the distribution of the freshwater calanoid copepod (Diaptomidae and Pseudodiaptomidae) fauna of the Yucatan Peninsula (YP) and its relation to the geological history of this Neotropical karstic plain. Location The Yucatan Peninsula, Mexico. Methods Plotting of geo‐referenced sites, analysis of local and regional geological history, analysis and comparison of regional and local records. Results The current composition and distribution of Diaptomidae and Pseudodiaptomidae in the YP mainly reflects recent, post‐Pliocene colonization events. This invasion did not reach America, but only parts of Central America (CA). The presence of diaptomids in the continent since the pre‐Cretaceous and the presumed post‐Cretaceous (Palaeocene–Oligocene) radiation of Diaptomidae in Middle America suggest earlier colonizations of the YP. The marine transgressions kept most of the YP submerged in different geological periods, thus eliminating any original primary freshwater colonizers, such as the diaptomids. The periods of marine regression probably represented opportunities for new waves of diaptomid colonization of the YP. The latest dispersal of diaptomids in the YP during the Holocene (8000 yr bp ) was probably an intermittent process because of the alternative dry and wet periods and interglacial transgressions. The presence of the Nearctic Leptodiaptomus and Arctodiaptomus in the YP and the current distribution of Mastigodiaptomus might represent remnants of earlier invasions of Diaptomidae in Middle America. The Neotropical Mastigodiaptomus probably originated in the Late Cretacic CA/proto‐Antilles complex. Forms derived from a M. albuquerquensis type ancestor radiated into the YP leaving relatively isolated populations of three species in the northern half of the peninsula. The distribution of the brackish water Pseudodiaptomus marshi well inside the coastline might have resulted from stranding and subsequent adaptation of this species during a marine regression in the Bacalar formation; this agrees with the vision of this taxon as being in process of invasion of freshwater environments. Main conclusions It is not probable that the South American (SA) diaptomid fauna originated from an invasion of upper Neotropical/Nearctic forms. The current distribution of freshwater calanoid copepods reflects relatively recent, post‐Pliocene biogeographical patterns, but probably older patterns are involved as well. The northern and eastern coasts of the Yucatan are the most recently colonized by diaptomids. Differing from other freshwater groups surveyed in the Yucatan that have marine relatives (i.e. fishes, amphipods, isopods, mysids, macrocrustaceans), there is no evidence of local vicariant events involving cave‐dwelling forms or marine relicts in the diaptomid fauna of the YP.     

Biogeographic regions of North American mammals based on endemism

Since the 19th Century, two regions have been recognized for North American mammals, which overlap in Mexico. The Nearctic region corresponds to the northern areas and the Neotropical region corresponds to the southern ones. There are no recent regionalizations for these regions under the criterion of endemism. In the present study, we integrate two methods to regionalize North America, using species distribution models of mammals: endemicity analysis (EA) and parsimony analysis of endemicity (PAE). EA was used to obtain areas of endemism and PAE was used to hierarchize them. We found 76 consensus areas from 329 sets classified in 146 cladograms, and the strict consensus cladogram shows a basal polytomy with 14 areas and 16 clades. The final regionalization recognizes two regions (Nearctic and Neotropical) and a transition zone (Mexican Transition Zone), six subregions (Canadian, Alleghanian, Californian‐Rocky Mountain, Pacific Central America, Mexican Gulf‐Central America, and Central America), two dominions (Californian and Rocky Mountain), and 23 provinces. Our analysis show that North America is probably more complex than previously assumed. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 485–499.     

Halffter's Mexican transition zone (1962–2014), cenocrons and evolutionary biogeography

The Mexican transition zone is the complex and varied area in which the Neotropical and Nearctic biotas overlap. In a series of contributions, Gonzalo Halffter provided a coherent theory that explains how sets of taxa that evolved in different geographical areas assembled in this transition zone. Halffter's theory developed gradually, being refined and clarified in successive contributions from him and other authors. After a review of the historical development of the Mexican transition zone, including the characterization of the dispersal or distributional patterns recognized by Halffter, its relevance for evolutionary biogeography is discussed briefly. The Mexican transition zone in the strict sense includes the highlands of Mexico and Guatemala (Sierra Madre Occidental, Sierra Madre Oriental, Transmexican Volcanic Belt, Sierra Madre del Sur and Chiapas Highlands provinces), whereas northern Mexico and the southern United States are clearly Nearctic, and the lowlands of southern Mexico and Central America are clearly Neotropical. The distributional patterns recognized by Halffter are considered to represent cenocrons (sets of taxa that share the same biogeographical history, constituting identifiable subsets within a biota by their common biotic origin and evolutionary history). The development of the Mexican transition zone is summarized into the following stages: (1) Jurassic–Cretaceous: the four Paleoamerican cenocrons extend in Mexico; (2) Late Cretaceous–Palaeocene: dispersal from South America of the Plateau cenocron; (3) Oligocene–Miocene: dispersal from the Central American Nucleus of the Mountain Mesoamerican cenocron; (4) Miocene–Pliocene: dispersal from North America of the Nearctic cenocron; and (5) Pleistocene: dispersal from South America of the Typical Neotropical cenocron.     

A preliminary survey of the Chironomidae (Diptera) of Costa Rica,with emphasis on the lotic fauna

Extensive collections of Chironomidae were made in Costa Rica, Central America, during 1986 and 1987. Fifty-five genera and at least 148 species belonging to the subfamilies Podonominae, Tanypodinae, Orthocladiinae and Chironomidae were found. Chironominae and Orthocladiinae predominated. Only one species of Podonominae was collected. Tanypodinae was represented by many genera, but species richness was low.Cricotopus was the most widespread and diverse genus of Orthocladiinae. Among the Chironominae the generaPolypedilum, Pseudochironomus, Tanytarsus andRheotanytarsus showed high species richness. Several species were collected that could not be assigned to genus. A number of range extensions were recorded for taxa found in the Neotropical region for the first time and for Neotropical taxa recorded outside of South America for the first time. The Costa Rican chironomid fauna consists of cosmopolitan, holarctic and neotropical components. There is probably an endemic Central American chironomid fauna at the species level.     

A new species of Leptodiaptomus (Copepoda, Diaptomidae) from Northwestern Mexico with comments on the distribution of the genus

A new species of the freshwater planktonic copepod genus Leptodiaptomus is described for a small pond in Northwestern Mexico. Leptodiaptomus dodsoni n. sp. can be easily distinguished mainly by the presence of an unusually large sinusoid spine on male antennular segment 13, and by the features of the fifth legs of both sexes. This genus is known to be distributed mainly in North America with 19 recognized species. Of these, six ocur in Mexico, and the new species seems to be closely related to most of them. It is probable that this group of species (including the new one) represents the southwards radiation of the genus from North America. Compared to the Caribbean and South American, the North American influence seems to be the most relevant for diaptomid copepods in Mexico. At least two Mexican species of Leptodiaptomus, including L.dodsoni, are restricted in distributional range to high-altitude temporal ponds, and both could be considered endemics.      

Revision of the aberrant New World genus Coenosopsia Diptera: Anthomyiidae, with a discussion of anthomyiid relationships

Abstract. Morphological evidence from the adults males, females corroborates the recent transfer of the problematic New World genus Coenosopsia from the Muscidae to the Anthomyiidae. It is argued that the calyptrate families Anthomyiidae and Muscidae are sister­groups, and that a Neotropical anthomyiid clade, Coenosopsia Malloch + Phaonantho Albuquerque, constitutes the sister-group of the remaining Anthomyiidae. At present, this clade is largely confined to tropical forests of South and Central America, i.e. an environment avoided by practically all other members of this pronouncedly cold-adapted, boreal family. The first, earliest Coenozoic, biotic exchange between North and South America and subsequent biotic isolation are proposed to lie behind the primary dichotomy of the Anthomyiidae. Five species of Coenosopsia , with apparently perfectly allopatric ranges, are recog nized: the Neotropical C.prima Malloch from Ecuador to SE Mexico, C.peruviana sp.n. from Peru, C.brasiliensis sp.n. from Brazil, and C.mexicana sp.n. from SW Mexico, and the Nearctic C.floridensis sp.n. from N Mexico and SE United States.     

Using 18S rDNA to resolve diaptomid copepod (Copepoda: Calanoida: Diaptomidae) phylogeny: an example with the North American genera

The phylogenetic relationships among the numerous genera of diaptomid copepods remain elusive due to difficulties in obtaining sufficient numbers of phylogenetically informative morphological characters for cladistic analysis. Molecular phylogenetic techniques offer high potential to resolve phylogenetic relationships in the absence of sufficient morphological characters because of the ease in which many characters can be unambiguously coded. I present the first molecular phylogeny for diaptomid copepod genera using 18S rDNA. Specifically, I test Light’s (1939) hypothesis regarding the interrelationships among the North American diaptomid genera. The 18S phylogeny is remarkably consistent with Light’s hypothesis. The endemic North American genera represent a monophyletic group exclusive of the non-endemic genera. Moreover, his hypothesized basal genus for the North America genera, Hesperodiaptomus, is the basal genus in this analysis. However, his Leptodiaptomus group is not reciprocally monophyletic with his Hesperodiaptomus group, but is rather a derived member of the latter group. Finally, the genus Mastigodiaptomus is found to be more closely allied with the non-endemic genera, as Light suggested. This phylogeny contributes heavily to the understanding of phylogenetic relationships among North American diaptomids and has large implications for the systematics of diaptomids in general. The use of 18S rDNA sequences in phylogenetic analyses of diaptomid copepods can be used to confirm the monophyly of recognized genera, the interrelationships among genera, and subsequent biogeographic interpretation of the family’s diversification. The use of molecular data, such as 18S rDNA sequences, to test phylogenetic hypotheses based on a very limited number of morphological characters will be a particularly useful approach to phylogenetic analysis in this system.     

Panbiogeographical analysis of the genus Bomarea (Alstroemeriaceae)

Aim  A panbiogeographical analysis of the genus Bomarea was undertaken in order to determine generalized tracks and biogeographical nodes, and to evaluate the current distribution of the genus based on the available tectonic information and the biogeographical regionalization of Latin America.
Location  The Neotropical region from northern Mexico to northern Argentina, and the Nearctic and Andean regions.
Method  A total of 2205 records of 101 species were analysed, representing 95% of the species assigned to Bomarea . Localities were represented on maps and their individual tracks were drawn. Based on their comparison, generalized tracks were detected and mapped. Nodes were identified in the areas where different generalized tracks were superimposed.
Results  Five generalized tracks were recovered. One is located in the Eastern Central America and Western Panamanian Isthmus provinces (Caribbean subregion, Neotropical region), which was supported by three species of Central American distribution. The four remaining generalized tracks were located in South America, in the North Andean Paramo, Cauca and Puna biogeographical provinces. These tracks were supported by species of Bomarea with an Andean distribution. Biogeographical nodes were established in the Central Andean region of Colombia, central Ecuador and central Peru.
Main conclusions  The nodes obtained for Bomarea support a hybrid origin for the Andean region, which presents diverse components from both northern and southern South America. Likewise, the track recovered between Colombia and Ecuador includes Andean and Neotropical areas, providing further support for this hypothesis. The nodes obtained are coherent with vicariant elements evident for Bomarea. Species of three clades proposed for Bomarea are distributed in specific generalized tracks.     

The New World oil‐collecting bees Centris and Epicharis (Hymenoptera,Apidae): molecular phylogeny and biogeographic history

We present a first comprehensive time‐calibrated phylogeny for two Neotropical genera of bees, Centris and Epicharis, whose females collect floral oil together with pollen for larval provisioning, and that traditionally have been grouped in the tribe Centridini. Our analyses rely on a matrix of 167 taxa and 4228 aligned nucleotides for the subfamily Apinae, with denser sample of Centris and Epicharis. Centris and Epicharis are strongly supported monophyletic groups, but Centridini is paraphyletic in relation to the corbiculate bees. The inner phylogenetic relationships of Epicharis agree with the current taxonomic classification. In Centris, three main clades were recovered, namely the Centris, Trachina and Melacentris groups. Inner relationships in Centris suggest the reinstatement of some subgenera and proposition of new ones. Early diversification of Centris and Epicharis took place at the tropical regions of South America. Epicharis and the Melacentris group in Centris mostly diversified in that region, expanding over Central America and tropical North America only recently in the last 3 My. The groups Trachina and Centris present a complex biogeographic history, with expansions to the Nearctic region, the Antilles, and temperate regions of South America in the late Oligocene and Miocene.     

A Pictorial Key for the Symphylan Families and Genera of the Neotropical Region South of Central Mexico (Myriapoda,Symphyla)

A pictorial key in English and Portugese is given for 2 families and 6 genera of the class Symphyla currently known from the Neotropical region, viz. Central America south of central Mexico, the Caribbean and entire South America.     

Biogeography of mutualistic fungi cultivated by leafcutter ants

Leafcutter ants propagate co‐evolving fungi for food. The nearly 50 species of leafcutter ants (Atta, Acromyrmex) range from Argentina to the United States, with the greatest species diversity in southern South America. We elucidate the biogeography of fungi cultivated by leafcutter ants using DNA sequence and microsatellite‐marker analyses of 474 cultivars collected across the leafcutter range. Fungal cultivars belong to two clades (Clade‐A and Clade‐B). The dominant and widespread Clade‐A cultivars form three genotype clusters, with their relative prevalence corresponding to southern South America, northern South America, Central and North America. Admixture between Clade‐A populations supports genetic exchange within a single species, Leucocoprinus gongylophorus. Some leafcutter species that cut grass as fungicultural substrate are specialized to cultivate Clade‐B fungi, whereas leafcutters preferring dicot plants appear specialized on Clade‐A fungi. Cultivar sharing between sympatric leafcutter species occurs frequently such that cultivars of Atta are not distinct from those of Acromyrmex. Leafcutters specialized on Clade‐B fungi occur only in South America. Diversity of Clade‐A fungi is greatest in South America, but minimal in Central and North America. Maximum cultivar diversity in South America is predicted by the Kusnezov–Fowler hypothesis that leafcutter ants originated in subtropical South America and only dicot‐specialized leafcutter ants migrated out of South America, but the cultivar diversity becomes also compatible with a recently proposed hypothesis of a Central American origin by postulating that leafcutter ants acquired novel cultivars many times from other nonleafcutter fungus‐growing ants during their migrations from Central America across South America. We evaluate these biogeographic hypotheses in the light of estimated dates for the origins of leafcutter ants and their cultivars.     

North American Glyptodontines (Xenarthra, Mammalia) in the Upper Pleistocene of northern South America

The Glyptodontidae is one of the most conspicuous groups in the Pleistocene megafauna of the Americas. The Glyptodontinae were involved in the Great American Biotic Interchange (GABI) and their earliest records in North America are about 3.9 Ma, suggesting an earlier formation of the Panamanian landbridge. Taxonomically it is possible to recognize two Pleistocene genera of Glyptodontinae:Glyptodon (ca. 1.8 – 0.008 Ma), restricted to South America, andGlyptotherium (ca. 2.6 – 0.009 Ma), including records in both North and Central America. Here we present the first report of the genusGlyptotherium in South America, from the Late Pleistocene of several fossil localities in Falcón State, northwestern Venezuela. A comparative analysis of the material, represented by cranial and postcranial parts, including the dorsal carapace and caudal rings, suggests a close affinity withGlyptotherium cylindricum (Late Pleistocene of Central Mexico). This occurrence in the latest Pleistocene of the northernmost region of South America Supports the bidirectional faunal migration during the GABI and the repeated re-immigration from North America of South American clades, as has been reported in other members of the Cingulata (e.g., Pampatheriidae).      

Biogeographical affinities among Neotropical cloud forests

 Biogeographical affinities among cloud forests in the Neotropical region were studied through a track approach, by constructing generalised tracks based on the results of a parsimony analysis of endemicity (PAE). Distributional data on 946 genera and 1,266 species of vascular plants (Pteridophyta, angiosperms, and gymnosperms) from 26 cloud forest patches from Colombia, Costa Rica, Cuba, Honduras, Jamaica, Mexico, Peru, Puerto Rico, and Venezuela were analysed; and four localities from eastern and western United States were also included as outgroups. The track analysis identified six generalised tracks: a first one that includes the majority of the cloud forests of Mexico, Central America, the Antilles, and northern Colombia; a second one that includes southern Mexico and northern Central America; a third one that includes the mountains in northwestern South America; a fourth one that includes the mountains in southwestern South America; and two others in western and eastern United States. It is concluded that the Neotropical cloud forests are closely related and that those of the Caribbean subregion exhibit complex relationships, which could be due to the complex tectonic history of the area. Received February 22, 2001 Accepted May 1, 2001     

New record of the genus Ithytrichia (Trichoptera: Hydroptilidae) for South America,with descriptions of male,larva and pupa of a new species from northwestern Argentina

Ithytrichia is a small genus of Hydroptilidae, recorded from Europe, North America, and north Central Mexico. The first record of Ithytrichia for South America is presented here. A new species of this genus Ithytrichia ferni is described from northwestern Argentina. Diagnostic characters of the male imago, pupa and larva are described and illustrated. Biological notes are included.     

Pugs of the tribe Eupitheciini (Lepidoptera,Geometridae): the history of study and distribution over biogeographic regions

The World fauna of the tribe Eupitheciini is the most species-rich in the family Geometridae. This tribe includes about 1900 species (almost 3000 species-group names) from 47 genera; about one third of the genera (15) are monotypic. The generic diversity of Eupitheciini is the highest in the Australian (38 genera, 11 of them endemic) and Oriental regions (32 genera, 4 endemic) and the lowest in the Neotropical Region (possibly one genus only). The faunas of different biogeographic regions can be arranged in following order by their species richness: the Palaearctic (487 species), Oriental (397), Neotropical (346), Australian (251), Afrotropical (198), and Nearctic Regions (166 species). Eupithecia is the most species-rich genus in the family Geometridae and the entire order Lepidoptera, and one of the largest genera in the whole World fauna of insects. The greatest number of species of this genus is recorded in the Palaearctic Region (466 species), where Eupithecia accounts for about 95% of the tribe Eupitheciini. The mainland of the Oriental Region (especially the Himalayas) is also very species-rich; however the proportion of the Eupithecia representatives decreases towards Malaysia, Sundaland, and the Australian Region (about 2% of the tribe). The Eupitheciini faunas have the greatest similarity at the generic level between the Oriental and Australian Regions (the Jaccard and Sørensen coefficient values being 0.62 and 0.77, respectively). The Palaearctic fauna is more similar to the Afrotropical and Oriental faunas at the genus-group level. On the whole, the fauna of the Nearctic Region is similar to that the West Palaearctic, with the exception of the fact that representatives of the genera Gymnoscelis and Chloroclystis are absent in North America, although two endemic genera Nasusina and Prorella are present. At the genus-group level, the Nearctic fauna of Eupitheciini is more similar to the Neotropical (the Jaccard and Sørensen coefficients 0.20 and 0.33, respectively) than to the Palaearctic fauna (0.17 and 0.29). The number of synonymies is very high in the tribe Eupitheciini because of the homogeneity of this group, whose species are difficult to identify without the use of elaborate anatomical techniques. Modern revisions, catalogues, surveys, and atlases on Eupitheciini are absent for many countries and large geographic regions. Revisions of pugs of the tribe Eupitheciini for some biogeographic regions are extremely difficult because of fragmentation of entomological collections including the type specimens of many species-group taxa. A large fraction of synonyms is characteristic of parts of the World with the best known faunas: Europe (64% of synonyms) and North America (39%). On the contrary, the lowest levels of synonymy are typical of the less known faunas of the regions situated at the equatorial latitudes, namely the Neotropical (9%) and Afrotropical (8%) ones.     

Historical biogeography and diversification within the Neotropical parrot genus Pionopsitta (Aves: Psittacidae)

Aim We investigate spatial and temporal patterns of diversification within the Neotropical avifauna using the phylogenetic history of parrots traditionally belonging to the genus Pionopsitta Bonaparte 1854. This genus has long been of interest for those studying Neotropical biogeography and diversity, as it encompasses species that occur in most Neotropical forest areas of endemism. Location The Neotropical lowland forests in South and Central America. Methods Phylogenetic relationships were investigated for all species of the genus Pionopsitta and five other short‐tailed parrot genera using complete sequences of the mitochondrial genes cyt b and ND2 as well as 26 plumage characters. The resulting phylogeny was used to test the monophyly of the genus, investigate species limits, and as a framework for reconstructing their historical biogeography and patterns of diversification. Results We found that the genus Pionopsitta, as previously defined, is not monophyletic and thus the Chocó, Central American and Amazonian species will now have to be placed in the genus Gypopsitta. The molecular and morphological phylogenies are largely congruent, but disagree on the position of one of the Amazon basin taxa. Using molecular sequence data, we estimate that species within Gypopsitta diversified between 8.7 and 0.6 Ma, with the main divergences occurring between 3.3 and 6.4 Ma. These temporal results are compared to other taxa showing similar vicariance patterns. Main conclusions The results suggest that diversification in Gypopsitta was influenced mainly by geotectonic events, marine transgressions and river dynamics, whereas Quaternary glacial cycles of forest change seem to have played a minor role in the origination of the currently recognized species.     

Historical biogeography and distribution of the freshwater cyclopine copepods (Copepoda, Cyclopoida, Cyclopinae) of the Yucatan Peninsula, Mexico

Aim To determine and analyse the distribution of the freshwater cyclopine copepod fauna of the Yucatan Peninsula (YP) and its relationship with the geological and climatic history of this Neotropical karstic zone. Location The Yucatan Peninsula, Mexico. Methods Plotting of georeferenced sites, analysis of local and regional geological and climatic history, analysis and comparison of regional and local faunistic records. Results Distinct dispersal and/or vicariant processes seem to be linked to the current distributions of the seven genera known in the YP. In general, the endemic hypogean or benthic crevicular forms (i.e. Diacyclops chakan, D. puuc and Mesocyclops chaci), derived from epigean, tropical, widely distributed forms (some of them South American) may have been among the earliest colonizers of the subterranean habitats in the YP. The distribution of these and other endemic forms seem to be related to the Holocene dry periods that desiccated the largest bodies of water and isolated local populations of different species. These vicariant processes resulted in forms with restricted distributional areas; some of these formed sister species that speciated in geographically close localities but related to a common identifiable ancestor. Overall, the processes of cyclopine colonization of the YP show the influence of the South American fauna, as the closest relatives of some species endemic to the YP are South American forms; the Nearctic influence is low. The cyclopine fauna of the YP is formed by a mixture of Nearctic‐derived (species of Acanthocyclops), Neotropical (i.e. M. edax, M. longisetus, A. panamensis, Thermocyclops inversus and T. tenuis), and epigean and hypogean endemic forms. The highly dynamic geomorphology of the YP and the recent climatic changes in the Holocene define the YP as a peculiar subregion that harbours a diverse fauna of cyclopine copepods with a high endemism. Main conclusion The current distribution of cyclopine copepods reflects relatively recent, post‐Pliocene biogeographical patterns; probably older patterns are involved as well. The eastern coast of the Yucatan is the most recently colonized by cyclopine copepods. Most of the genera are linked with South American forms, and the Nearctic influence is weakly represented. This group has no marine relatives, but there is evidence of vicariant events involving cave‐dwelling forms.     

New World biogeography and the evolution of polychromatism: evidence from the bee assassin genus Apiomerus (Heteroptera: Reduviidae: Harpactorinae)

The crassipes and pictipes species groups of Apiomerus Hahn together contain 12 species, many with high intraspecific chromatic variability, which represent the majority of Nearctic species in this New World assassin bug genus. Because of their geographical distribution and their varying degrees of polychromatism, these two species groups provide a unique opportunity to study the evolution of polychromatism and analyse relationships among areas of endemism in the Nearctic, as well as determine the boundary between the Nearctic and Neotropical regions. The results of a morphology based phylogenetic analysis allowed investigation of these questions while also determining relationships among the 12 species in the two species groups. The crassipes and pictipes species groups were each supported as monophyletic and as sister taxa. Apiomerus rufipennis (Fallou) was not included in the original concept of the crassipes species group, but is shown here to be a member of the group. Apiomerus barrocoloradoi Forero, Berniker & Szerlip, which had been hypothesized previously to belong to the pictipes species group, is excluded from this group. Intraspecific polychromatism for each species was identified as being present in one of three states: no polychromatism; limited polychromatism; or polychromatism as discrete colour morphs. Limited polychromatism was here found to be the ancestral state for Apiomerus, and species with discrete colour morphs are restricted to the crassipes and pictipes species groups. Polychromatism appears to be a greatly homoplastic character within the genus. A Brooks parsimony analysis recovered distinct Nearctic and Neotropical clades. The Nearctic clade is divided between areas in the central and eastern U.S.A. and areas in the Western U.S.A. and Mexico. The Nearctic–Neotropical boundary for the taxa included in the analysis is along the Isthmus of Tehuantepec in southern Mexico.