Patterns of morphological variation amongst semifossorial shrews in the highlands of Guatemala,with the description of a new species (Mammalia,Soricomorpha, Soricidae)

Members of the Cryptotis goldmani group of small‐eared shrews (Mammalia, Soricomorpha, Soricidae) represent a clade within the genus that is characterized by modifications of the forelimb that include broadened forefeet, elongated and broadened foreclaws, and massive humeri with enlarged processes. These modifications are consistent with greater adaptation to their semifossorial habits than other members of the genus. The species in this group occur discontinuously in temperate highlands from southern Tamaulipas, Mexico, to Honduras. In Guatemala, there are three species: the relatively widespread Cryptotis goodwini and two species (Cryptotis lacertosus, Cryptotis mam) endemic to highland forests in the Sierra de los Cuchumatanes of western Guatemala. Ongoing studies focusing on the relationships of variation in cranial and postcranial skeletal morphology have revealed a fourth species from remnant cloud forest in the Sierra de Yalijux, central Guatemala. In this paper, I describe this new species and characterize its morphology relative to other species in the C. goldmani group and to other species of Cryptotis in Guatemala. In addition, I summarize available details of its habitat and ecology. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 163 , 1267–1288.     

Molecular Phylogeny Supports Repeated Adaptation to Burrowing within Small-Eared Shrews Genus of Cryptotis (Eulipotyphla,Soricidae)

Small-eared shrews of the New World genus Cryptotis (Eulipotyphla, Soricidae) comprise at least 42 species that traditionally have been partitioned among four or more species groups based on morphological characters. The Cryptotis mexicana species group is of particular interest, because its member species inhibit a subtly graded series of forelimb adaptations that appear to correspond to locomotory behaviors that range from more ambulatory to more fossorial. Unfortunately, the evolutionary relationships both among species in the C. mexicana group and among the species groups remain unclear. To better understand the phylogeny of this group of shrews, we sequenced two mitochondrial and two nuclear genes. To help interpret the pattern and direction of morphological changes, we also generated a matrix of morphological characters focused on the evolutionarily plastic humerus. We found significant discordant between the resulting molecular and morphological trees, suggesting considerable convergence in the evolution of the humerus. Our results indicate that adaptations for increased burrowing ability evolved repeatedly within the genus Cryptotis.     

Molecular systematics and biodiversity of the Cryptotis mexicanus group (Eulipotyphla: Soricidae): two new species from Honduras supported

Small-eared shrews of the genus Cryptotis (Mammalia: Eulipotyphla: Soricidae) are widespread in the northern Neotropics. Systematic studies of these shrews over the past two decades have revealed previously undocumented morphological and species diversity, resulting in a quadrupling of the number of recognized species. Unfortunately, a small proportion of the species in the genus have been included in molecular phylogenetic studies, and evolutionary relationships within the genus are incompletely known. Traditionally, species have been assigned to four or five morphologically defined ‘species groups’, but tests of the monophyly of some of these groups show weak support and relationships amongst species groups remain somewhat speculative. The largest species group is the C. mexicanus group inhabiting Mexico and northern Central America. We studied sequences from mitochondrial cytochrome-b and 16S genes, as well as nuclear ApoB and BRCA1 genes from 22 species of Cryptotis, including 15 species in the C. mexicanus group. Our combined analysis shows that the C. goldmani subgroup is very weakly supported as monophyletic; however, the C. mexicanus group as a whole is not monophyletic. Our molecular phylogenetic analyses confirm the distinctiveness of two newly described species (C. celaque and C. mccarthyi) from isolated highlands of western Honduras and illustrate their relationship with other species formerly considered part of a widespread C. goodwini.     

Phylogenetics and taxonomy of the New World leafy spurges,Euphorbia section Tithymalus (Euphorbiaceae)

The 480 species of leafy spurges, Euphorbia subgenus Esula, represent the main temperate radiation in the large genus Euphorbia. This group is distributed primarily in temperate Eurasia, but with smaller, disjunct centres of diversity in the mountains of the Old World tropics, in temperate southern Africa and in the New World. The majority of New World diversity (32 species) occurs in a single section, section Tithymalus. We analysed sequences of the nrITS and plastid ndhF, trnH‐psbA, trnS‐trnG and trnD‐trnT regions to reconstruct the phylogeny of section Tithymalus and to examine the origins and diversification of the species native to the New World. Our results indicate that the New World species of section Tithymalus form a clade that is sister to the widespread, weedy E. peplus. The New World species fall into two primary groups: a ‘northern annual clade’ from eastern North America and a diverse clade of both annual and perennial species that is divided into three subgroups. Within the second group, there is a small ‘southern annual clade’ from Texas and northern Mexico, a perennial ‘Brachycera clade’ from the western United States and northern Mexico, and a perennial ‘Esuliformis clade’ from montane areas of Mexico, Guatemala, Honduras and the Caribbean island of Hispaniola. Ancestral state reconstructions indicate that the annual habit probably evolved in the ancestor of E. peplus and the New World clade, with a subsequent reversal to the perennial habit. In conjunction with this phylogenetic framework, the New World species of section Tithymalus are comprehensively reviewed. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175 , 191–228.     

Systematics of the genus Mayazomus (Arachnida: Schizomida): the relevance of using continuous characters and pedipalp setae patterns to schizomid phylogenetics

The schizomid genus Mayazomus Reddell & Cockendolpher, 1995, endemic to south‐eastern Mexico, currently comprises seven species. It was originally proposed to accommodate two species, from Chiapas and Tabasco. Recently, five additional species from Chiapas were described. The monophyly of the genus has never been tested using cladistic analysis. We undertook a phylogenetic analysis using the seven described species of Mayazomus as the ingroup, ten exemplar species representing the most similar New World hubbardiids as the outgroup, and one protoschizomid species to root the tree. The analysis was based on 130 morphological characters (continuous and discrete characters). The resulting topologies recovered Mayazomus as paraphyletic, with Heteronochrus estor Armas & Viquez, 2010, from Guatemala nested within the genus; therefore, we formally propose its synonymy herein. Mayazomus appears to be most closely related to Rowlandius Reddell & Cokendolpher, 1995, a South American genus. This contribution also provides new characters derived from the pedipalp setae with important phylogenetic information; as well as the implementation of morphometric ratios, as continuous characters, to partially codify the shape of the male flagellum. The relationships recovered amongst the outgroups used in this contribution are a reliable baseline for future analyses of the phylogeny of the New World schizomids.     

Reassessment of the cranial characters of Glossotherium and Paramylodon (Mammalia: Xenarthra: Mylodontidae)

Nomenclatural confusion has existed within the Mylodontinae for several genera, and has resulted in the supposition that Paramylodon of North America is synonymous with Glossotherium of South America. A taxonomic revision of crania for Glossotherium and Paramylodon upholds their separation as distinct genera and provides a list of diagnostic characters that have been lacking. Assessment was made using principal components analysis for suites of cranial and mandibular measurements, evaluation of ratios and measurement distribution, and by examining qualitative characters. Results show the greatest characterization for the skull comes from differences relating to cranial length versus width, whereas the mandible is predominantly distinguished by qualitative characters of the predental spout. Examination of the Pliocene species Glossotherium chapadmalense from South America shows a combination of characters indicative of each genus, but exhibits more with Glossotherium and is tentatively retained under that genus. The mix of characters indicates that G. chapadmalense is the likely ancestor to Paramylodon, although when and where the transition took place is still unclear. During the evolutionary transition, Paramylodon crania emphasized an increase in length of the palate, whereas those of Glossotherium emphasized an increase in cranial width. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155 , 885–903.     

Gone to Texas: phylogeography of two Trachymyrmex (Hymenoptera: Formicidae) species along the southeastern coastal plain of North America

Ants are widely recognized as ecologically important members of many low‐ to mid‐latitude ecosystems. Surprisingly, there is very little phylogeographical information on ants at regional scales. We examine here the phylogeography of two partially sympatric species of Trachymyrmex (Trachymyrmex septentrionalis and Trachymyrmex turrifex) ants in southeastern North America. We test the hypothesis that all Trachymyrmex species found in the USA expanded into North America from refugial populations located in northern Mexico as the post‐Pleistocene climate warmed. Phylogeographical theory predicts that these northward‐expanding species should exhibit higher genetic diversity in regions closer to Mexico and less diversity in more northern regions. We also examine, in the widely distributed T. septentrionalis, the hypothesis of vicariance that occurred at the formation of the Mississippi Embayment. Phylogeographical patterns indicate that T. septentrionalis has an eastern origin because diversity was highest east of the Mississippi, whereas T. turrifex probably has a Mexican origin because it lacked mitochondrial DNA (mtDNA) variation throughout its range and is currently absent from eastern North America. Both species are characterized by reduced haplotypic variation in the western coastal plain of the Gulf of Mexico (Texas and Louisiana), which may indicate recent expansion and/or bottlenecks associated with increased aridity and drought in these western regions. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 114 , 689–698.     

Distribution,host plant affiliation,phenology, and phylogeny of walnut‐infesting Rhagoletis flies (Diptera: Tephritidae) in Mexico

Many taxa of Nearctic origin have diversified in the subtropical highlands of Mexico. In particular, flies in the genus Rhagoletis have undergone episodes of isolation and gene flow during Pleistocene glaciations and post‐glacial times that have produced lineage differentiation and reproductive isolation. To reach a better understanding of the phylogeography of the genus Rhagoletis, a host plant survey of the walnut‐infesting Rhagoletis suavis species group was conducted across sixteen states comprising 34 different collecting sites in Mexico over a 9‐year period. Five species of Juglans were found to be infested by three species of walnut‐attacking Rhagoletis flies. Several species of parasitoids were also recovered from collections, but in contrast to their walnut fly hosts, they revealed little evidence for host or geographic subdivision. There was no consistent difference in mean eclosion time between walnut fly species or populations associated with different host walnuts in Mexico, unlike the case for other Rhagoletis species, in which allochronic isolation arising from variation in diapause timing is a major ecological adaptation, reproductively isolating flies. We compare the distribution of R. suavis flies in Mexico with those of other Rhagoletis species attacking hawthorns and cherries, and discuss its implications for population divergence and speciation. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 765–779.     

A tale of North and South America: time and mode of dispersal of the amphitropical genus Munroa (Poaceae,Chloridoideae)

Plant disjunctions have provided some of the most intriguing distribution patterns historically addressed by biogeographers. We evaluated the three hypotheses that have been postulated to explain these patterns [vicariance, stepping‐stone dispersal and long‐distance dispersal (LDD)] using Munroa, an American genus of grasses with six species and a disjunct distribution between the desert regions of North and South America. The ages of clades, cytology, ancestral characters and areas of distribution were investigated in order to establish relationships among species, to determine the time of divergence of the genus and its main lineages, and to understand further the biogeographical and evolutionary history of this genus. Bayesian inference recovered the North American M. pulchella as sister species to the rest. Molecular dating and ancestral area analyses suggest that Munroa originated in North America in the late Miocene–Pliocene (7.2 Mya; 8.2–6.5 Mya). Based on these results, we postulate that two dispersal events modelled the current distribution patterns of Munroa: the first from North to South America (7.2 Mya; 8.2–6.5 Mya) and the second (1.8 Mya; 2–0.8 Mya) from South to North America. Arid conditions of the late Miocene–Pliocene in the Neogene and Quaternary climatic oscillations in North America and South America were probably advantageous for the establishment of populations of Munroa. We did not find any relationship between ploidy and dispersal events, and our ancestral character analyses suggest that shifts associated with dispersal and seedling establishment, such as habit, reproductive system, disarticulation of rachilla, and shape and texture of the glume, have been important in these species reaching new areas. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 179 , 110–125.     

Phylogenetic interrelationships,taxonomy, and reductive evolution in the Neotropical electric fish genus Hypopygus (Teleostei,Ostariophysi, Gymnotiformes)

A phylogenetic reconstruction of the Neotropical electric fish genus Hypopygus based on 47 parsimony‐informative morphological characters is presented. A series of synapomorphies support the hypothesis of monophyly of Hypopygus, and partially resolve species‐level relationships within the genus. Hypopygus species are recognized here as miniaturized fishes based on two criteria; first, a derived condition of diminutive body size, and; second, the presence of a suite of reductive morphological characters, including partial or total losses, simplifications, and reductions of the anal‐fin rays, scales, cranial bones, and laterosensory canal system. Reductive characters associated with miniaturization comprise 45% of the total number of characters in the phylogenetic reconstruction of the genus. Miniaturization and reductive morphological evolution in Hypopygus are discussed here in the phylogenetic context. A taxonomic revision of Hypopygus is presented, in which five new species are described, two species previously assigned to the genus are redescribed, and a single known species of Stegostenopos is redescribed and included in Hypopygus as a junior synonym. Distribution maps and a key for all eight valid species of Hypopygus are provided, based on the examination of 5014 catalogued museum specimens. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 163 , 1096–1156.     

Phylogeny of the spider genus Ixchela Huber, 2000 (Araneae: Pholcidae) based on morphological and molecular evidence (CO1 and 16S), with a hypothesized diversification in the Pleistocene

The genus Ixchela Huber is composed of 20 species distributed from north‐eastern Mexico to Central America, including the five new species described here from Mexico: I xchela azteca sp. nov. , I xchela jalisco sp. nov. , I xchela mendozai sp. nov. , I xchela purepecha sp. nov. and I xchela tlayuda sp. nov. We test the monophyly and investigate the phylogenetic relationships among species of the genus Ixchela using morphological and molecular data. Parsimony (PA) analysis of 24 taxa and 40 morphological characters with equal and implied weights supported the monophyly of Ixchela with eight morphological synapomorphies. The PA analyses with equal and implied weights, and separate Bayesian inference (BI) analyses for the CO1 gene (506 characters), concatenated gene fragments CO1 + 16S (885 characters), morphology + CO1 (546 characters) and the combined evidence data set (morphology + CO1 + 16S) (925 characters) support the monophyly of Ixchela. Our preferred topology shows two large clades; clade 1 has a natural distribution in the Mesoamerican biotic component, whereas clade 2 predominates in the Mexican Montane biotic component. The genus Ixchela diverged in the late Miocene, and the divergence between the internal clades in the genus occurred in the late Pliocene; by contrast, most of the speciation events seem to have occurred mainly during the Pleistocene, where climatic changes brought on by repeated glaciations played an important role in the diversification of the genus. © 2015 The Linnean Society of London     

A new Stygonitocrella Petkovski (Copepoda: Harpacticoida) from a cave in Northern Mexico with comments on the taxonomy of the genus

The cave-dwelling harpacticoid copepod fauna of Mexico is still yet to be known. There are only a few records related to karstic caves, but most refer to epigean forms. The analysis of samples collected in a cave system in Northern Mexico yielded two female specimens of an unknown harpacticoid copepod species which is described herein. It was assigned to the genus Stygonitocrella Petkovski, 1976 mainly because of the presence of a single endopodal segment on legs 1–4; the new species, S. mexicana n. sp. differs from its congeners by a combination of characters related to the segmentation and armature of the swimming legs 1–4, the fifth leg of the female, and the strcture of the antennae. Based on the current limits of Stygonitocrella, two new subgenera are proposed to recognize different morphological patterns based on the presence or absence of an inner baseoendopodal lobe in the female fifth leg, a character not evaluated before. This lobe is absent in the new species and in other five species now contained in the subgenus Eustygonitocrella. A lobe is present in the new subgenus Fiersiella, also with six species. This is the second record of Stygonitocrella in North America, after S. (Fiersiella) sequoyahi Reid, Hunt & Stanley, 2003; it is the only North American cave-dwelling species of the genus. It is suggested that the influence of marine transgressions in Northern Mexico was related to the origin of the new species as part of a group of troglobitic forms originated from marine interstitial ancestors that colonized the freshwater caves. A key for the identification of all the known species of Stygonitocrellais also provided.     

Leaf epidermal micromorphology of Cercis (Fabaceae: Caesalpinioideae)

The micromorphology of leaf epidermal cells and stomata of all eight species and one form (11 samples) of Cercis were observed by scanning electron microscopy and light microscopy. Both the adaxial and abaxial epidermal cells are polygonal or irregular in shape; the anticlinal walls are straight and arched or undulate. Two types of stomata, which occur only on the abaxial surface of the leaves, are found in the genus. The atypical paracytic type is present in only one species, Cercis chingii, and the anomocytic type is present in all other species. Interspecific differences are minor in the genus with regard to leaf epidermal characters, except for C. chingii, which is characterized by atypical paracytic‐type stomata, a two‐lipped outer stomatal rim, the highest stomatal density and undulate and densely pitted anticlinal walls in the adaxial epidermis. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 158 , 539–547.     

Evaluating hypotheses on the origin and diversification of the ringneck snake Diadophis punctatus (Colubridae: Dipsadinae)

Close affinities recognized between taxa in Mexico and the contiguous USA have led to a variety of biogeographical scenarios. One such hypothesis suggests that species that occur in both countries have an origin in central Mexico followed by dispersal into the USA. This study expands upon previous phylogeographical work of the ringneck snake Diadophis punctatus by incorporating new data from previously unsampled areas appropriate to critically assess hypotheses regarding a Mexican origin for this species. Maximum likelihood and maximum parsimony analyses inferred a derived position for the lineage from southern Mexico with constraint tests for alternate evolutionary hypotheses resulting in significantly worse likelihood values. Ancestral area reconstructions inferred an origin for D. punctatus in the south‐eastern USA followed by a south‐east to north‐east then westward directionality of historical migration. The position within the phylogeny and date estimate for the south‐western + Mexico clade suggests a recent invasion into central Mexico with expansion into the Nearctic/Neotropic transition zone. The extensive lineage diversity inferred from the mtDNA suggests that the genus is a complex of cryptic species whose conservational status should be re‐evaluated on both the national and regional levels. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 158 , 629–640.     

Systematics and cladistics of a new Naupactini genus (Coleoptera: Curculionidae: Entiminae) from the Andes of Colombia and Ecuador

A new genus of broad‐nosed weevils belonging to the tribe Naupactini (Coleoptera: Curculionidae) is described, based on six species from the Páramo–Puna subregion (Andean region) of South America. Three species are new and three have been transferred from the genus Asymmathetes (Asymmathetes nigrans, Asymmathetes rugicollis, and Asymmathetes vulcanorum). The new genus is recognized by the black, denuded, and shiny tegument, the well‐developed pre‐epistome, the elytral base curved backwards on middle, the reduction of the hindwings, and the widely separated procoxae, closer to the anterior than to the posterior margin of the prosternum. A cladistic analysis of the six species of the new genus plus five out‐groups (Amitrus, Amphideritus, Asymmathetes, Melanocyphus, and Trichocyphus), using 49 morphological characters, resulted in a single cladogram. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 166 , 54–71.     

Phylogenetic analysis of Micrathena and Chaetacis spiders (Araneae: Araneidae) reveals multiple origins of extreme sexual size dimorphism and long abdominal spines

The phylogenetic relationships amongst the New World spiny orb‐weaving spiders Micrathena and Chaetacis were assessed through parsimony and Bayesian analyses of morphological characters. A total of 146 characters was scored for ten outgroup taxa and 37 Micrathena and four Chaetacis species. The results indicate that Chaetacis nests within Micrathena and we propose Chaetacis as a junior synonym of Micrathena. Twelve subgeneric species groups of Micrathena are recognized and diagnosed. Species with extremely long spines evolved at least eight times in the genus and we suggest that this may be related to antipredator defences. Micrathena is primitively sexually monomorphic and extreme sexual size dimorphism has arisen at least six times in the genus. Most of these events are because of enlargement of the female in relation to the ancestral size, although in two cases sexual dimorphism was attained through male reduction, adding more data to the ‘giant females’ vs. ‘dwarf males’ controversy. The genus is probably of South American origin and has repeatedly invaded Central and North America. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 166 , 14–53.     

Biogeographical analysis of two Polypodium species complexes (Polypodiaceae) in Mexico and Central America

We analyzed the geographical and elevational distributions of two Polypodium complexes from Mexico and Central America. Distribution data of nine species of the Polypodium colpodes complex and the Polypodium plesiosorum complex were obtained from almost 1500 herbarium specimens, field collections in Mexico and Costa Rica, and literature studies. The presence of each species was recorded for each Mesoamerican country, in 1° × 1° grid‐cells and biogeographical provinces. The rarity of species was also evaluated. Although the two complexes show extensive overlap, the P. colpodes complex is distributed mainly along the Pacific versant of Mexico and Central America, whereas the P. plesiosorum complex occurs mainly along the Atlantic versant. Those biogeographical provinces with maximum species diversity are Chiapas (seven species), Sierra Madre del Sur (six species), and the Trans‐Mexican Volcanic belt (six species). Grid‐cells with more species are located mainly in the mountains of central‐southern Mexico and northern Central America. Richness does not decrease or increase with latitude. Elevation distributions showed that most Polypodium species are concentrated in the montane interval and three species groups were recognized based on elevational preferences. Polypodium colpodes and P. plesiosorum are the most widely distributed species, whereas Polypodium castaneum and Polypodium flagellare are the only two species that possess the three attributes of rarity (narrow geographical distribution, high habitat specificity, and scarce local populations). Polypodium species of both complexes are present mainly in the montane regions of the study area and show some degree of geographical sympatry, especially in southern Mexico and northern Central America. This overlapping is explained by the elevation tolerance within montane systems and because most species inhabit three or more vegetation types. The distributional patterns of these complexes coincided with the three regional highlands of Mesoamerica, which are separated from each other by the Isthmus of Tehuantepec and by the lowlands of Nicaragua. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ?? , ??–??.     

Multilocus species delimitation in Mesoamerican Scaptotrigona stingless bees (Apidae: Meliponini) supports the existence of cryptic species

Accelerating taxonomic knowledge and making accurate species identifications are critically important given the current biodiversity crisis, particularly in biodiversity hotspots such as Mesoamerica. Objective species delimitation that reduces investigator‐driven bias is fundamental to the establishment of appropriate conservation strategies, above all in managed species. Previous morphological and molecular studies on three managed stingless bee species of the genus Scaptotrigona distributed in Mexico (S. mexicana, S. pectoralis and S. hellwegeri) suggested that both S. mexicana and S. hellwegeri are cryptic species complexes. Herein we tested species delimitation by analysing sequence information of five markers (two mitochondrial: cox1 and 16S, and three nuclear: ITS1, EF1‐α, ArgK) within a Bayesian coalescent framework to test the putative species. We obtained two different hypotheses using a Generalized Mixed Yule Coalescent (GMYC) model: four (cox1) and six (16S) species. After the species validation step with the Bayesian species‐delimitation analysis (BPP), we suggest that only S. mexicana is a complex of two species with different distribution (along the Pacific and the Atlantic coasts, respectively). We highly recommend avoiding colony exchange between geographical regions in order to conserve the genetic integrity of both taxa.