Ephemeral Pleistocene woodlands connect the dots for highland rattlesnakes of the Crotalus intermedius group

Aim To test how Pleistocene climatic changes affected diversification of the Crotalus intermedius species complex. Location Highlands of Mexico and the south‐western United States (Arizona). Methods We synthesize the matrilineal genealogy based on 2406 base pairs of mitochondrial DNA sequences, fossil‐calibrated molecular dating, reconstruction of ancestral geographic ranges, and climate‐based modelling of species distributions to evaluate the history of female dispersion. Results The presently fragmented distribution of the C. intermedius group is the result of both Neogene vicariance and Pleistocene pine–oak habitat fragmentation. Most lineages appear to have a Quaternary origin. The Sierra Madre del Sur and northern Sierra Madre Oriental are likely to have been colonized during this time. Species distribution models for the Last Glacial Maximum predict expansions of suitable habitat for taxa in the southern Sierra Madre Occidental and northern Sierra Madre Oriental. Main conclusions Lineage diversification in the C. intermedius group is a consequence of Pleistocene climate cycling. Distribution models for two sister taxa in the northern and southern Sierra Madre Occidental and northern Sierra Madre Oriental during the Last Glacial Maximum provide evidence for the expansion of pine–oak habitat across the Central Mexican Plateau. Downward displacement and subsequent expansions of highland vegetation across Mexico during cooler glacial cycles may have allowed dispersal between highlands, which resulted in contact between previously isolated taxa and the colonization of new habitats.     

Areography of the genus Dendroctonus (Coleoptera: Curculionidae: Scolytinae) in Mexico

Objective To analyse whether the geographical ranges of Dendroctonus species are (1) associated with factors such as host species or elevation, and (2) in agreement with Halffter's Nearctic distribution pattern. (3) To identify and discuss the factors that are likely to act as barriers to the genus’ geographical distribution. (4) To explore whether there is an association between the size of the geographical ranges of Dendroctonus species and the number of Pinus host species used by each of them, and (5) to assess if these host species are most common at the elevations preferred by the individual Dendroctonus species. Site Mexico. Methods Records of 12 species of Dendroctonus were gathered from entomological collections in Mexico. Distribution ranges were defined by using the propinquity method ( Rapoport, 1975a ). Analysed parameters were: (1) geographical distribution of single species, (2) overlapping of species ranges, (3) disjunction patterns and barriers by means of isoprobabilistic lines, based on the morphotectonic subdivision of Mexico ( Ferrusquía‐Villafranca, 1998 ), (4) spatial variation in species richness with respect to latitude and altitude, (5) size of geographical ranges, and (6) host species for each Dendroctonus species. A correlation was determined between area size and number of pine host species. Results The species ranges varied in shape and size. Geographical ranges tend to be discontinuous in shape. Composite patterns showed that disjunctions among ranges do not closely follow Mexico's morphotectonic subdivision. There are repeated discontinuities among individual distributions, which define five areas: (1) Baja California Peninsula, (2) Sierra Madre Occidental (SMOC), (3) northern Sierra Madre Oriental (SMOR), (4) Sierra Madre de Chiapas, and (5) SMOR + Faja Volcanica Transmexicana (FVT) + Sierra Madre del Sur. The isoprobabilistic lines confirm that the inner part of SMOC provides an optimal environment for the genus, and the FVT province constitutes the broader corridor for it in the country. Richness does not directly decrease or increase with latitude. Richness behaviour of the insect is not associated with that of its host. Elevation distributions showed that most Dendroctonus species move within broad margins of tolerance and species richness is concentrated in the montane interval. Dendroctonus attack 24 of the 47 Pinus species distributed in Mexico. Preferred pine species belong predominantly to Leiophyllae, Ponderosae and Oocarpae subsections. The Spearman rank correlation between area size and number of pine host species was not significant. Dendroctonus clearly belongs to a Nearctic distribution pattern (sensu Halffter, 1987 ). Main conclusions Dendroctonus is present in all montane systems of Mexico and its species coexist within a high geographical sympatry. Overlapping of species distribution appears to be the result of two elements – generalized polyphagy inside Pinus and a wide elevation tolerance within mountainous environments. This behaviour, linked to a high vagility, has allowed the genus Dendroctonus to expand its distribution across Mexico and to employ mountainous systems as corridors separated by barriers that exert a low selective filter effect.     

Taxonomy of <Emphasis Type="Italic">Koeberlinia</Emphasis> (Koeberliniaceae)

Koeberlinia has a natural amphitropical distribution that includes the deserts of central Bolivia, northern Mexico, and the southwestern United States. Despite the long recognition of only one species, K. spinosa, field, herbarium, and SEM studies support the recognition of two species. Koeberlinia spinosa of northern Mexico and adjacent United States is recognized to consist of three varieties: K. spinosa var. spinosa of northeastern Mexico and the adjacent United States, K. spinosa var. tenuispina of the Sonoran Desert of southwestern Arizona, adjacent California, and northwestern Mexico, and K. spinosa var. wivaggii from south central Texas and northern Mexico to Arizona, which is described as new. Koeberlina holacantha, endemic to the deserts of Bolivia, is proposed as new.     

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.     

Distribution and Broadscale Habitat Relations of the Wolverine in the Contiguous United States

ABSTRACT Conservation of the wolverine (Gulo gulo) at the southern extent of its North American range requires reliable understandings of past and present distribution patterns and broad-scale habitat relations. We compiled 820 verifiable and documented records of wolverine occurrence (specimens, DNA detections, photos, and accounts of wolverines being killed or captured) in the contiguous United States from museums, the literature, and institutional archives. We spatially referenced 729 records with areal precision ≤1 township (93.2 km²) and temporal precision ≤10 years. Historical records (1827–1960) were located primarily in the western mountains and Great Lakes region. However, our data suggest that the historical distribution of wolverines in the Cascade Range and Sierra Nevada was disjunct, contradicting previous interpretations. Our results indicate that wolverine range in the contiguous United States had contracted substantially by the mid-1900s. Current records (1995–2005) are limited to north-central Washington, northern and central Idaho, western Montana, and northwestern Wyoming. We investigated potential relations between wolverines and alpine vegetation, cold temperatures, and spring snow cover by comparing the distribution of historical wolverine records with Kuchler's potential natural vegetation types, Holdridge's climatic life zones, and EASE snow-cover maps during the latter portion of the wolverine denning period (15 Apr-14 May). In the western mountains, historical wolverine records generally occurred in or near alpine vegetation and climatic conditions, especially at the limits of their distribution in the Cascade Range, Sierra Nevada, and southern Rocky Mountains. However, the only habitat layer that fully accounted for historical distribution patterns was spring snow cover. Causal factors for the extirpation of wolverines from the southern portions of their range in the contiguous United States are unknown, but are likely related to high levels of human-caused mortality and low to nonexistent immigration rates.     

Phylogeography of the bark beetle Dendroctonus mexicanus Hopkins (Coleoptera: Curculionidae: Scolytinae)

Dendroctonus mexicanus is polyphagous within the Pinus genus and has a wide geographical distribution in Mexico and Guatemala. We examined the pattern of genetic variation across the range of this species to explore its demographic history and its phylogeographic pattern. Analysis of the mtDNA sequences of 173 individuals from 25 Mexican populations allowed to us identify 53 geographically structured haplotypes. High haplotype and low nucleotide diversities and Tajima’s D indicate that D. mexicanus experienced rapid population expansion during its dispersal across mountain systems within its current range. The nested clade phylogeographic analysis indicates that the phylogeographic pattern of D. mexicanus is explained by continuous dispersion among lineages from the Sierra Madre Occidental, the Sierra Madre Oriental and the Trans-Mexican Volcanic Belt. However, we also observed isolation events among haplotypes from the Cofre de Perote/Trans-Mexican Volcanic Belt/Sierra Madre Oriental and the Trans-Mexican Volcanic Belt/Sierra Madre del Sur, which is consistent with the present conformation of mountain systems in Mexico and the emergence of geographical barriers during the Pleistocene.     

Potential distribution of Ursus americanus in Mexico and its persistence: Implications for conservation

The black bear Ursus americanus is an endangered species in Mexico. Its historical distribution has decreased by approximately 80% although its current distribution is not known with precision; it is only reported to be present in the mountains of Northern Mexico. This study proposes two ensemble models: Mexicós black bear (a) potential distribution compared with Natural Protected Areas (NPAs); and, (b) persistence areas for 2024. The current distribution variables are coniferous forest, elevation and dry forest. Suitable habitat for black bear (354,047 km², 18.07% of the country) was found mainly in the north of the Sonoran biogeographical zone, along the Sierra Madre Occidental, the center and south of the Sierra Madre Oriental and some northern regions of the Altiplano Norte. Comparing these areas with NPAs documented that only 12.41% of potential distribution coincided with current suitable habitat. There are unprotected areas in Sierra Madre Occidental center and central and southern of Sierra Madre Oriental. The model for 2024 indicates a reduction of suitable habitat of 64.5%, mainly in the northern Sonoran zone and the center Sierra Madre Occidental. On the other hand, areas that will persist (125,673 km²) are located along the two main mountain ranges of Mexico. Identification of these sites will allow strengthening of long-term conservation strategies.     

Range-wide chloroplast and mitochondrial DNA imprints reveal multiple lineages and complex biogeographic history for Douglas-fir

The contemporary genetic structure of species offers key imprints of how organisms responded to past geological and climatic events, which have played a crucial role in shaping the current geographical distribution of north-temperate organisms. In this study, range-wide patterns of genetic variation were examined in Douglas-fir (Pseudotsuga menziesii), a dominant forest tree species distributed from Mexico to British Columbia in western North America. Two organelle DNA markers with contrasting modes of inheritance were genotyped for 613 individuals from 44 populations. Two mitotypes and 42 chlorotypes were recovered in this survey. Both genomes showed significant population subdivision, indicative of limited gene flow through seeds and pollen. Three distinct cpDNA lineages corresponding to the Pacific Coast, the Rocky Mountains, and Mexico were observed. The split time of the two lineages from the Rockies lineage was dated back to 8.5 million years (Ma). The most recent common ancestors of Mexican and coastal populations were estimated at 3.2 and 4.8 Ma, respectively. The northern populations of once glaciated regions were characterized by a high level of genetic diversity, indicating a large zone of contact between ancestral lineages. A possible northern refugium was also inferred. The Mexican lineage, which appeared established by southward migration from the Rockies lineage, was characterized by the lowest genetic diversity but highest population differentiation. These results suggest that the effects of Quaternary climatic oscillations on the population dynamics and genetic diversity of Douglas-fir varied substantially across the latitudinal section. The results emphasize the pressing need for the conservation of Mexican Douglas-fir.     

Cladistic biogeography of the Mexican transition zone

Biogeographic relationships among nine montane areas of endemism across the transition zone between North and South America are analysed cladistically based on phylogenetic hypotheses of thirty‐three resident monophyletic taxa of insects, fish, reptiles, and plants. Areas of endemism include the Arizona mountains (AZ), Sonoran Desert (SD), Sierra Madre Occidental (OCC), southern Sierra Madre Occidental (SOC), Sierra Madre Oriental (ORI), Sierra Transvolcanica (TRAN), Sierra Madre del Sur (SUR), Chiapan‐Guatemalan Highlands (CGH), and Talamancan Cordillera (TC). Area relationships are summarized using Brooks Parsimony Analysis and Assumption 0, with the former resulting in more defensible biogeographic hypotheses. Areas of endemism are dividable into two monophyletic groups; a northern group including AZ, SD, OCC, and ORI, and a southern group consisting of TC, CGH, TRAN, SUR, and the isolated southern regions of the Sierra Madre Occidental (SOC). The northern set of areas are characterized by recent, probably Pleistocene, isolation and prevalent widespread species, whereas the southerly areas probably diverged after Pliocene closure of the Panamanian isthmus. The southern areas are redundantly represented on many of the taxon‐area cladograms by endemic species, indicative of much higher levels of endemism in the Sierra Transvolcanica and further south. Use of a general area cladogram in such a transition zone permits explicit exploration of biogeographic patterns and establishes a predictive framework for taxonomy and conservation prioritization.     

Environmental suitability for Agrilus auroguttatus (Coleoptera: Buprestidae) in Mexico using MaxEnt and database records of four Quercus (Fagaceae) species

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Phylogeography of the mountain chickadee (Poecile gambeli): diversification, introgression, and expansion in response to Quaternary climate change

Since the late 1990s, molecular techniques have fuelled debate about the role of Pleistocene glacial cycles in structuring contemporary avian diversity in North America. The debate is still heated; however, there is widespread agreement that the Pleistocene glacial cycles forced the repeated contraction, fragmentation, and expansion of the North American biota. These demographic processes should leave genetic 'footprints' in modern descendants, suggesting that detailed population genetic studies of contemporary species provide the key to elucidating the impact of the late Quaternary (late Pleistocene-Holocene). We present an analysis of mitochondrial DNA (mtDNA) variation in the mountain chickadee (Poecile gambeli) in an attempt to examine the genetic evidence of the impact of the late Quaternary glacial cycles. Phylogenetic analyses reveal two strongly supported clades of P. gambeli: an Eastern Clade (Rocky Mountains and Great Basin) and a Western Clade (Sierra Nevada and Cascades). Post-glacial introgression is apparent between these two clades in the Mono Lake region of Central California. Within the Eastern Clade there is evidence of isolation-by-distance in the Rocky Mountain populations, and of limited gene flow into and around the Great Basin. Coalescent analysis of genetic variation in the Western Clade indicates that northern (Sierra Nevada/Cascades) and southern (Transverse/Peninsular Ranges) populations have been isolated and evolving independently for nearly 60,000 years.     

Molecular evidence for Pleistocene glacial cycles driving diversification of a North American desert spider, Agelenopsis aperta

The influence of historical climatic vs. geological changes on species diversification patterns was investigated in a widely distributed North American desert spider, Agelenopsis aperta (Araneae: Agelenidae), with particular reference to Pleistocene glacial cycles and earlier patterns of mountain building. Levels of sequence divergence obtained from the mitochondrial gene, cytochrome oxidase I, dated to the Pleistocene, eliminating Rocky Mountain orogeny as a cause of diversification, as orogeny ended 4 million years ago. The results of phylogenetic and network analyses showed the presence of three geographically defined clades, which were consistent with the presence of at least three glacial refugia: (i) east of the Rocky Mountains; (ii) between the Rocky Mountains and Sierra Nevadas; and (iii) west of the Sierra Nevadas. In addition, populations within the Rocky Mountains exhibited significantly lower genetic diversity than populations east of the Rocky Mountains and the haplotypes found within the Rockies were a subset of eastern haplotypes. These patterns suggest that a post-Pleistocene range expansion occurred out of an eastern glacial refugium into the Rocky Mountains. Examination of phylogeographical studies of other North American desert taxa indicated that mountain building explained diversification patterns more effectively for some taxa but Pleistocene climate change was more important for others, including A. aperta.     

Fomitiporia cupressicola sp. nov., a parasite on Cupressus arizonica, and additional unnamed clades in the southern USA and northern Mexico, determined by multilocus phylogenetic analyses

Fomitiporia cupressicola sp. nov., found in living Cupressus arizonica, is described on the basis of several collections originating from a high altitude forest in the northern Sierra Madre Occidental, Mexico. The species forms a monophyletic clade, basal to a larger lineage comprising species originating mainly from temperate to Mediterranean areas of the northern hemisphere. The phylogenetic approach in Fomitiporia also revealed multiple unnamed clades within the F. robusta complex in the southern USA and northern Mexico, representing potential species. The status of the F. robusta complex in North America is discussed briefly.     

Genetic variation coincides with geographic structure in the common bush-tanager (Chlorospingus ophthalmicus) complex from Mexico

Cloud forests are distributed in the Neotropics, from northern Mexico to Argentina, under very specific ecological conditions, namely slopes with high humidity input from clouds and mist. Its distribution in Mesoamerica is highly fragmented, similar to an archipelago, and taxa are thus frequently represented as sets of isolated populations, each restricted to particular mountain ranges and often showing a high degree of divergence, both morphologically and genetically. The common bush-tanager (Chlorospingus ophthalmicus, Aves: Thraupidae) inhabits cloud forests from eastern and southern Mexico south to northwestern Argentina. Here we use 676bp of mtDNA (around the ATPase 8 gene) to explore the genetic variation and phylogeographic structure of the Mexican populations of C. ophthalmicus. Phylogenetic analyses of mtDNA sequences indicate deep genetic structure. Five major clades, which segregate according to geographic breaks, are identified (starting from the deepest one in the phylogeny): (1) Southern Chiapas and Northern Central America, (2) Tuxtlas massif, (3) Sierra Madre del Sur, (4) Eastern Oaxaca and Northern Chiapas, and (5) Sierra Madre Oriental. The long history of isolation undergone by each clade, as suggested by the phylogeny, implies that the species status of each of them should be revised.     

Taxonomy and biogeography of the fiddler crabs (Ocypodidae: Genus Uca) of the Atlantic and Gulf coasts of eastern North America

Fifteen species of fiddler crabs are reported for eastern North America between Massachusetts and Quintana Roo, Mexico. Thirteen occur in the United States and 11 in Mexico, with eight in common to the two countries. Of 13 species in the Gulf of Mexico, five are endemic and a sixth is restricted largely to the peninsulas of Florida and Yucatan. The status of U. rapax in the northern Gulf remains to be resolved. Range limits of most species approximate one or the other of two sets of intersecting thermal and geological boundaries that subdivide the Gulf of Mexico along north-south and east-west axes. Species belonging to subgenus Minuca tend to replace one another at the thermally-controlled Carolinian-Caribbean marine biotic boundary across the Florida peninsula and northern Gulf. However, only U. minax of all the North American fiddler crabs exhibits the classical disjunct Carolinian distribution, and this appears basically to reflect the discontinuous distribution of temperate salt marshes that are the habitat of the species. Distributions of species belonging to subgenus Celuca adhere for the most part to the subdivision of the Gulf into western terrigenous and eastern carbonate sedimentary provinces. The northern transition occurs in the vicinity of Apalachee Bay and the southern at Laguna de Terminos. A third distributional pattern is shown by U. subcylindrica , a specialized endemic species of the hypersaline Laguna Madre system of the western Gulf. The level of endemism in the fiddler crabs is relatively high in comparison with that of other marine groups within the Gulf of Mexico. This may be a consequence of the adaptations of fiddler crabs as specialized deposit feeders to regional differences in climatic and edaphic characteristics of a marginally marine upper shore habitat. The distributional patterns of the endemics could prove useful in reconstructing palaeoecological events of evolutionary significance within the Gulf of Mexico.     

Phylogeography of Middle American gophersnakes: mixed responses to biogeographical barriers across the Mexican Transition Zone

Aim We used inferences of phylogeographical structure and estimates of divergence times for three species of gophersnakes (Colubridae: Pituophis) distributed across the Mexican Transition Zone (MTZ) to evaluate the postulated association of three Neogene geological events (marine seaway inundation of the Isthmus of Tehuantepec, formation of the Transvolcanic Belt across central Mexico, and secondary uplifting of the Sierra Madre Occidental) and of Pleistocene climate change with inter‐ and intraspecific diversification. Location Mexico, Guatemala, and the western United States. Methods We combined range‐wide sampling (67 individuals representing three putative species distributed across northern Middle America and western North America) and phylogenetic analyses of 1637 base pairs of mitochondrial DNA to estimate genealogical relationships and divergence times. The hypothesized concordance of inferred gene trees with geological histories was assessed using topology tests. Results We identified three major lineages of Middle American gophersnakes, and strong phylogeographical structure within each lineage. Gene trees were statistically congruent with hypothesized geological histories for two of the three postulated geological events. Estimated divergence dates and the geographical distribution of genetic variation further support mixed responses to these geological events. Considerable phylogeographical structure appears to have been generated during the Pleistocene. Main conclusions Phylogenetic and phylogeographical structure in gophersnakes distributed across northern Middle America and western North America highlights the influence of both Neogene vicariance events and Pleistocene climate change in shaping genetic diversity in this region. Despite the presence of two major geographical barriers in southern Mexico, extreme geological and environmental heterogeneity in this area may have differentially structured genetic diversity in highland taxa. To the north, co‐distributed taxa may display a more predictable pattern of diversification across the warm desert regions. Future studies should incorporate nuclear data to disentangle inferred lineage boundaries and further elucidate patterns of mitochondrial introgression.     

Modeling distributions of disjunct populations of the Sierra Madre Sparrow

ABSTRACT Sierra Madre Sparrows (Xenospiza baileyi) are among the least known of all bird species in Mexico. Recent surveys have discovered previously unknown populations and the current known distribution of Sierra Madre Sparrows consists of two populations separated by >800 km. We used available distributional information to develop ecological niche models that (1) predict much of the distribution potential of the species, (2) establish that the broad disjunction separating the two populations has ecological correlates that appear to be important to the distributional of these sparrows, and (3) illustrate the extremely restricted ecological distribution of the species. We used two sets of climatic and topographic variables, with one including all 22 variables available and the second with only six variables that were positively related to quality of distributional models. Although indications of differences between the two sets of populations were found based on the full 22‐dimensional environmental dataset, such a highly dimensional analysis is vulnerable to over‐fitting; models based on the reduced dataset indicated that the two populations occur in areas with similar ecological conditions. Our models also suggest that southern population of Sierra Madre Sparrows covers most of their potential range in that region. The potential range of the northern population, however, extends beyond known points of occurrence. To clarify the distribution of Sierra Madre Sparrows and evaluate their status and conservation opportunities, detailed searches for additional populations in areas identified by the model are needed.     

Phylogeography of the Buarremon brush-finch complex (Aves, Emberizidae) in Mesoamerica

The Buarremon brush-finches represent a complex suite of populations distributed in the montane New World Tropics from Mexico south to South America. Traditional taxonomic arrangements have separated populations of this genus into three species, based on plumage variation, although plumage patterns are well known to exhibit homoplasy. We present a first detailed phylogeographic and phylogenetic study, focused on Mesoamerican populations, and signal the existence of strong differentiation among populations with a clear geographic structure. We find well differentiated clades for (1) the Sierra Madre Oriental and Sierra Madre del Sur in Oaxaca, (2) western Mexican populations, including the B. brunneinucha populations in the Sierra Madre del Sur and B. virenticeps, (3) Sierra Madre Oriental and Sierra de los Tuxtlas, (4) northern Central America, (5) southern Central America, (6) middle Central America, and (7) South America. We demonstrate a lack of concordance with plumage patterns, and argue for several additional species to be recognized in the complex.