Systematics and evolution of the subfamily Gerbillinae (Mammalia, Rodentia, Muridae)

Although they represent a quarter of the mammalian species, the evolutionary relationships among as well as within, the main murid lineages are still controversial. The subfamily Gerbillinae is no exception as previous studies based on morphological, karyotypical, and allozyme characters are highly incongruent. Here, we present the first molecular phylogeny for gerbils based on cytochrome b and 12S rRNA mitochondrial genes. Results are largely congruent between the two genes as well as with the concatenated data set with most of the nodes being well-supported. Based on the topologies retrieved here, we (1) propose the identification of three main clades, (2) support the split of Tatera genus into an Asian and an African group, the latter including Gerbillurus species, and (3) provide some evidence towards the inaccuracy of subgeneric divisions within both Gerbillus and Meriones. In addition, the sharp contrast between the genetic characters and morphological data sets suggest high levels of convergence, probably as a result of strong environmental constraints imposed on these rodents adapted to arid and semi-arid regions. Finally, molecular datings for the various cladogenetic events are in good agreement with the known gerbilline fossil record and support an African origin with subsequent migrations to Asia.     

Systematics,biogeography and evolution of the endemic Hemidactylus geckos (Reptilia,Squamata, Gekkonidae) of the Cape Verde Islands: based on morphology and mitochondrial and nuclear DNA sequences

A total of 1854 bp of mitochondrial DNA (669 bp of cytochrome b (cyt b) and 386 bp of 12S rRNA), and 804 bp of a nuclear gene (RAG2) were investigated in endemic Hemidactylus from eight Cape Verde Islands, and used to explore their phylogeny, biogeography and evolution. Maximum‐likelihood, maximum‐parsimony and Bayesian analyses based on mtDNA revealed four well‐supported clades with uncorrected genetic divergences of 7.8–12.4% in the cyt b plus 12S rRNA genes, which were also supported by nuclear DNA. A population from the southern island of Fogo is the most divergent in both molecules and morphology and is described as Hemidactylus lopezjuradoi sp. n., and the populations on Sal and Boavista are also assigned species status as H. boavistensis. Although divergent in their DNA, the clade on S. Nicolau and that in the north‐western islands are morphologically similar and both are assigned to H. bouvieri for the present. Hemidactylus b. razoensis from Raso is genetically similar to H. b. bouvieri and differs only in its smaller body size. A molecular clock suggests that the ancestor of the endemic Hemidactylus of the Cape Verde Islands colonized the archipelago approximately 10 ± 2.48 Mya, perhaps reaching the north‐eastern islands first. The H. lopezjuradoi lineage separated soon after, and the north‐western islands were colonized progressively but slowly, S. Nicolau probably being reached first, then S. Vicente and islands on the same bank, and finally Sto. Antão, which is likely to have been colonized less than 1 Mya. Hemidactylus boavistensis is abundant on the arid islands where it occurs, but H. bouvieri appears to have been uncommon at least since it was described 130 years ago, and the same may be true of H. lopezjuradoi sp. n. The impact of introduced H. angulatus and H. mabouia on the endemic Hemidactylus of the Cape Verde Islands is not clear, but the discovery of substantial genetic diversity in endemic Cape Verde Hemidactylus means that the conservation requirements of the group should be reassessed.     

Systematics and functional morphology of Molarochoerus yuanmouensis (Suidae, Mammalia) from the Late Miocene of Yunnan, China

The discovery of a propotamochoerine suid with unique features of the dentition and skull led to the naming of Molarochoerus yuanmouensis Liu & Pan [6]. Further study of the material reveals that this species was better adapted than Sus scrofa for the ‘rooting’ behaviour and that its dentition has morphological features indicating that it was eating hard food items. The posterior premolars are completely molarised, hence the generic name, the first time that any suid has been shown to possess almost perfectly molariform P4/s and p/4s. In lateral view the curvature of the occlusal surface of the upper cheek teeth is convex ventrally, the opposite of the usual situation in suids. The origins of the rostral musculature are exceptionally well developed, and the nasal bones are much more robust than is usually the case in Suinae, indicating powerful musculature and fortified osseous structures presumably for ‘rooting’ in hard ground or for extended periods of time. The masseteric musculature appears to have been more massive than it is in Sus, Microstonyx and Propotamochoerus. To cite this article: M. Pickford et al., C. R. Palevol 3 (2004).

Résumé

Systématique et morphologie fonctionnelle de Molarochoerus yuanmouensis (Suidae, Mammalia) du Miocène supérieur de Yunnan, Chine. La découverte d”un Suidae Propotamochoerini présentant des caractères dentaires et crâniens uniques a conduit à la création de Molarochoerus yuanmouensis Liu & Pan [6]. Une étude du matériel montre que cette espèce était bien mieux adaptée au fouissage que Sus scrofa. Par ailleurs, les caractères morphologiques de sa dentition indiquent qu'il se nourrissait d'aliments coriaces. Les prémolaires postérieures sont complètement molarisées, d'où le nom générique, et c'est la première fois qu'un Suidae présente des P4 supérieures et inférieures presque parfaitement molariformes. En vue latérale, la courbure de la surface occlusale des dents jugales supérieures est convexe ventralement, différant ainsi de la morphologie classique des Suidae. La musculature rostrale est exceptionnellement bien développée et les os nasaux sont beaucoup plus robustes que chez les autres Suinae, suggérant des structures musculaires et osseuses puissantes, probablement liées au fouissage, soit dans des sols, soit sur une longue durée. Les muscles massétériens sont plus puissants que dans les genres Sus, Microstonyx et Propotamochoerus. Pour citer cet article : M. Pickford et al., C. R. Palevol 3 (2004).     

Molecular phylogeny and biogeography of Oriental voles: genus Eothenomys (Muridae, Mammalia)

Oriental voles of the genus Eothenomys are predominantly distributed along the Southeastern shoulder of the Qinghai-Tibetan Plateau. Based on phylogenetic analyses of the mitochondrial cytochrome b gene (1143 bp) obtained from 23 specimens (eight species) of Oriental voles collected from this area, together with nucleotide sequences from six specimens (two species) of Japanese red-backed voles (Eothenomys andersoni and Eothenomys smithii) and five species of the closely related genus Clethrionomys, we revised the systematic status of Eothenomys. We also tested if vicariance could explain the observed high species diversity in this area by correlating estimated divergence times to species distribution patterns and corresponding paleo-geographic events. Our results suggest that: (1) the eight species of Oriental voles form a monophyletic group with two distinct clades, and that these two clades should be considered as valid subgenera--Eothenomys and Anteliomys; (2) Eothenomys eleusis and Eothenomys miletus are not independent species; (3) Japanese red-backed voles are more closely related to the genus Clethrionomys than to continental Asian Eothenomys taxa; and (4) the genus Clethrionomys, as presently defined, is paraphyletic. In addition, the process of speciation of Oriental voles appears to be related to the Trans-Himalayan formation via three recent uplift events of the Qinghai-Tibetan Plateau within the last 3.6 million years, as well as to the effects of the mid-Quaternary ice age.     

Systematics and macroevolution of extant and fossil scalopine moles (Mammalia,Talpidae)

Scalopini is one of the two fully fossorial mole tribes in the family Talpidae, with remarkable adaptations to subterranean lifestyles. Most living Scalopini species are distributed in North America while a sole species occurs in China. On the other hand, scalopine fossils are found in both Eurasia and North America from upper Oligocene strata onwards, implying a complex biogeographical history. The systematic relationships of both extant and fossil Scalopini across North America and Eurasia are revised by conducting phylogenetic analyses using a comprehensive morphological character matrix together with 2D geometric–morphometric analyses of the humeral shape, with a specific emphasis on Mioscalops, a genus commonly found in North America and formerly known as Scalopoides. Our phylogenetic analyses support the monophyly of the tribe Scalopini as well as a proposed two‐subtribe‐division scenario of Scalopini (i.e. Scalopina and Parascalopina), although Proscapanus could not be assigned to either subgenus. Our geometric–morphometric analyses indicate that the European Mioscalops from southern Germany should be allocated to Leptoscaptor, which in turn implies that Mioscalops may be endemic to North America and never arrived in Europe. Examination of biogeographical patterns does not unambiguously determine the geographical origin of Scalopini. Nevertheless, it does support multiple transcontinental colonization events across Asia, Europe and North America. Scapanulus oweni, distributed in central China, is the only remaining representative of one of those out‐of‐North‐America migrations, whereas scalopine moles are common in North America nowadays with up to five species.     

Systematics, biogeography and conservation of Lactoridaceae

Lactoridaceae are a monotypic family confined to the Juan Fernandez (= Robinson Crusoe) Islands, Chile, an archipelago four million years old. Interest in the lone species, Lactoris fernandeziana, has increased in recent years due to its suspected role in early angiosperm evolution, as well as its endangered conservation status. Reports of fossil pollen of Lactoris (as Lactoripollenites) from deep sea sediments of Cretaceous age off the western coast of South Africa provide evidence for southern hemisphere occurrence of the family 70–90 million years ago. This age, plus trimerous symmetry and simple morphology of the flower with three tepals, six stamens and three nearly free carpels with laminar placentation, involve the family with hypotheses regarding evolution of early angiosperms, especially paleoherb and monocot divergences. Based on cladistic analyses with morphological and nucleotide data, recommended taxonomic placement of Lactoris is as a monotypic order, Lactoridales, allied most closely to Aristolochiales, and also near Piperales. Approximately 1000 individuals of Lactoris are believed to exist, and genetic variation within and among populations as measured by isozymes and RAPDs is low. This, in part, could be due to the self-compatibility and geitonogamous breeding system of this wind-pollinated species. More efforts at ex situ conservation should be attempted so that this important angiosperm family is maintained for future evolutionary studies.     

First identification of Decennatherium Crusafont, 1952 (Mammalia,Ruminantia, Pecora) in the Siwaliks of Pakistan

Previously undescribed remains of a new large giraffid have been identified from the late Miocene Siwaliks Hills of the Potwar Plateau in Pakistan. This taxon is very intimately related to the late Miocene giraffid genus Decennatherium, previously only certainly identified in the Iberian Peninsula, and with some possible remains assigned to Decennatherium crusafonti from Aliäbäd (Iran). The new material collected in the Siwaliks Hills shows a high morphological similarity with the Spanish remains of the genus, especially the early Vallesian Decennatherium pachecoi. Two previously undescribed ossicone fragments from the same area are also described and identified as cf. Decennatherium. These findings represent the easternmost occurrence of the genus Decennatherium and show a late Miocene migration of the genus throughout the Iberian Peninsula and Southern Asia.     

Systematics and biogeography of the jellyfish Aurelia labiata (Cnidaria: Scyphozoa)

The hypothesis that the common eastern North Pacific Aurelia is A. aurita is falsified with morphological analysis. The name Aurelia labiata is resurrected, and the species is redescribed, to refer to medusae differing from A. aurita by a suite of characters related to a broad and elongated manubrium. Specifically, the oral arms are short, separated by and arising from the base of the fleshy manubrium, and the planulae are brooded upon the manubrium itself, rather than on the oral arms. Aurelia aurita possesses no corresponding enlarged structure. Furthermore, the number of radial canals is typically much greater in A. labiata, and thus the canals often appear more anastomosed than in A. aurita. Finally, most A. labiata medusae possess a 16-scalloped bell margin, whereas the margin is 8-scalloped in most A. aurita. Separation of the two forms has previously been noted on the basis of allozyme and isozyme analyses and on the histology of the neuromuscular system. Partial 18S rDNA sequencing corroborates these findings. Three distinct morphotypes of A. labiata, corresponding to separate marine bioprovinces, have been identified among 17 populations from San Diego, California, to Prince William Sound, Alaska. The long-undisputed species A. limbata may be simply a color morph of A. labiata, or a species within a yet-unelaborated A. labiata species complex. The first known introduction of Aurelia cf. aurita into southern California waters is documented. Although traditional jellyfish taxonomy tends to recognize many species as cosmopolitan or nearly so, these results indicate that coastal species, such as A. labiata, may experience rapid divergence among isolated populations, and that the taxonomy of such species should therefore be scrutinized with special care.     

Systematics, biogeography, and evolution of the Neotropical peacock basses Cichla (Perciformes: Cichlidae)

To investigate forces influencing diversification in Neotropical fishes, the phylogenetic relationships among species and populations of the cichlid genus Cichla were examined. Mitochondrial DNA was sequenced for 454 individuals of the 5 nominal Cichla species and several putative undescribed species. Phylogenetic analyses support the distinction of two major clades of Cichla. Clade A includes C. temensis and two undescribed species from the lower Amazonas and Xingu Rivers. Clade B includes C. orinocensis, C. monoculus, C. ocellaris. C, intermedia, and an undescribed species from the upper Madeira River. Species boundaries were relatively well-circumscribed for clade B, while incomplete lineage sorting was inferred for clade A. Three probable instances of introgression were observed, including a regional population of C. orinocensis from the Negro River that shows a history of introgression. Biogeographic patterns from Cichla are partially congruent with those seen in several other Neotropical fish clades, and the diversification of Cichla species is inferred to result from both vicariance and sympatric divergence.     

Systematics and biogeography of the genus Actenodia (Coleoptera: Meloidae: Mylabrini)

Abstract.  The genus Actenodia Laporte de Castelnau (Meloidae: Mylabrini) is revised. It includes 18 species distributed in the Mediterranean and Saharo-Arabian regions, and in eastern and southern Africa; A. carpanetoi sp.n. from Mozambique is described. The bionomics of the genus is summarized, according to a collection of records on phenology, elevation, habitat preference and host plants. A classification of the genus is proposed after a morphological cladistic analysis: one Afrotropical lineage clearly emerged with three distinct groups of species, whereas other taxa, all Palaearctic except A. mirabilis , remained unresolved. The analysis of the nuclear internal transcribed spacer 2 rDNA sequences of some Palaearctic and Afrotropical species supports the distinction of the Afrotropical lineage. The first instar larvae of two species, A. chrysomelina from southern Africa and A. denticulata from Arabia, are described for the first time and compared with two other Palaearctic species described previously; inferences from larval morphology are discussed. The adult morphology of all species is described briefly and illustrated, and a key to the species is provided, as well as locality data. Three new synonymies are proposed: A. extera Dvořák, 1993 syn.n. = A. septempunctata Baudi di Selve, 1878; A. sexpunctata Pic, 1948 syn.n . = A. curtula Fähraeus, 1870 ; A. unimaculata lanzai Kaszab, 1973 syn.n . = A. unimaculata Pic, 1908. The biogeography of the genus is discussed within the framework of a more general analysis of the disjunct distribution models.     

A cranial morphometric study of deer (Mammalia, Cervidae) from Argentina using three-dimensional landmarks

This study examines morphological variation in the crania (n = 70) of eight cervid species from Argentina. Forty 3-dimensional landmarks were acquired on each adult cervid cranium. The data were analysed using Morphologika software. The co-ordinates were registered and scaled to remove size differences by Procrustes analysis, and then principal components analysis was applied to examine shape variation. Shape variation associated with each principal component can be visualised in the program. The first principal component correlates strongly with the centroid size of the crania and also with the body mass and height of each species. The larger species were distinguished by relatively longer snouts and relatively smaller brains. The smallerMazama andPudu species cluster closely on the first as well as the other principal components. Among the larger species, the twoHippocamelus species, which live at higher altitudes, were clearly distinguished from the lowland species,Ozotoceros bezoarticus andBlastocerus dichotomus, on the basis of cranial flexion and the orientation of the occipital region. FinallyO. bezoarticus andB. dichotomus were compared directly and small differences were noted in the orbital region. The shape data was used to produce a distance matrix and a phenogram, which we relate to some of currently accepted phylogenetic relationships of this group of cervids.     

Systematics, cladistics and biogeography of the American genus Farrodes (Ephemeroptera: Leptophlebiidae: Atalophlebiinae)

Cladistic and biogeographic analyses of the genus Farrodes are presented. Two species groups are delineated within Farrodes: F. caribbianus and F. bimaculatus. Three species formerly placed in other genera– Thraulus caribbianus Traver, Thraulus roundsi Traver and Homothraulus maculatus (Needham & Murphy)–are transferred to Farrodes. The species of the F. caribbianus species group are revised. Three new species are described: F. savagei from Venezuela, F. maya and F. mexicanus from Mexico. Keys to separate the two species groups of Farrodes and the species of the F caribbianus species group are provided. Successive cladistic analyses were carried out on both adult and nymphal characters using Hennig86 and CLADOS. The matrix was composed of all available data (nymphal characters were missing for some species), from nymphal and adult stages separately and on taxa represented by both adult and nymphal characters. Species of the genera Simothraulopsis and Homothraulus (components of the Farrodes lineage) were included in the analyses, and Ecuaphkbia was used as the outgroup. Results of the four analyses are compatible. The historical biogeography of Farrodes , with a distribution from nordiern Argentina to southern Texas, is analysed using the program COMPONENT. Areas of endemism are established, and some of their relationships compared with those of other groups available in the literature.     

Systematics and cladistics of Megalostomis Chevrolat,and the biogeography of Clytrini (Coleoptera: Cryptocephalinae)

We present a cladistic analysis of the subtribe Megalostomina, a Neotropical group of ‘case‐bearer’ leaf beetles. A comparative study of the external and internal adult morphology of Clytrini was undertaken. New characters are described for the subtribe Megalostomina, from the internal sac of aedeagus, which provide a useful phylogenetic signal. More than 180 photographs illustrating the most important characters (74 characters and their respective states) used in the cladistic analysis are provided. The cladistic analysis of 57 terminal taxa and 95 characters was undertaken, under equal weights, and also using implied weights as a means to down‐weight homoplasious characters. We test the monophyly and explore intergeneric relationships of the subtribe Megalostomina, and reconstruct the relationships among the species of Megalostomis Chevrolat. The 42 species recognized can be assigned either to a group mostly containing species of North and Central America, or to a larger one of mostly South American species. Support is low, and the formal naming of groups is deferred pending a revision of all Megalostomina. We confirm the subgenera of Megalostomis of previous classifications are unnatural, and the following changes in the generic classification of the subtribe Megalostomina are proposed: Coleorozena Moldenke syn.n. of Coscinoptera Lacordaire; Coleothorpa Moldenke syn.n. of Coscinoptera Lacordaire; and Euryscopa (Coleoguerina) Moldenke syn.n. of Coscinoptera Lacordaire. Furthermore, six formerly recognized subgenera of Megalostomis are considered junior synonyms of Megalostomis Chevrolat: Megalostomis (Minturnia) Lacordaire syn.n. ; Megalostomis (Heterostomis) Lacordaire syn.n. ; Megalostomis (Scaphigenia) Lacordaire syn.n. ; Megalostomis (Snellingia) Moldenke syn.n. ; Megalostomis (Coleobyersa) Moldenke syn.n. ; and Megalostomis (Pygidiocarina) Moldenke syn.n. Thus, no subgenera are recognized within Megalostomis. Previous hypotheses on Clytrini biogeography were revisited in the light of new biogeographic and phylogenetic knowledge. We hypothesize an origin of Clytrini in tropical/subtropical Gondwana, when South America, Africa, Madagascar and India were connected. Changes in the configuration of the tectonic plates in the Cenozoic allowed the dispersal of Clytrina to the Palaearctic and Nearctic regions, and dispersion of Babiina and Megalostomina through the Nearctic region.     

Mitochondrial and nuclear DNA phylogenies reveal a complex evolutionary history in the Australasian robins (Passeriformes: Petroicidae)

The Australasian robins (Petroicidae) comprise a relatively homogeneous group of small to medium-sized insectivorous birds. Their center of diversity is Australia and New Guinea (40 species) but seven species have managed to colonize geographically distant islands such as Tanimbar, New Britain, New Zealand, New Caledonia, Norfolk Island, Vanuatu, Solomon Islands, Fiji and Samoa. To resolve the evolutionary relationships within the Petroicidae, we here present the results of a phylogenetic analysis of sequence data from two mitochondrial genes (ND2, CO1) and one nuclear intron (β-Fibrinogen intron 5) for all 14 genera and 40 of the 46 currently recognized species. All phylogenetic analyses identified six primary lineages, treated here as subfamilies, within the Petroicidae: (1) Eopsaltriinae comprising Eopsaltria (excluding E. flaviventris), Tregellasia, Peneothello, Melanodryas, Poecilodryas and Heteromyias; (2) Drymodinae comprising Drymodes; (3) Microecinae comprising Microeca, Monachella and Eopsaltria flaviventris; (4) Petroicinae comprising Petroica and Eugerygone; (5) Pachycephalopsinae comprising Pachycephalopsis; and (6) Amalocichlinae comprising Amalocichla. The genera Eopsaltria, Microeca, Peneothello and Poecilodryas were found to be paraphyletic. Based on assessments of phylogenetic branching patterns and/or DNA divergence it also was apparent that Eopsaltriaaustralis, Tregellasialeucops, Melanodryascucullata, Heteromyiasalbispecularis, Drymodessupercilious and Microecaflavigaster may each comprise more than one species. The Petroicidae display a complex biogeographical history involving repeated radiations both within, and across Australia and New Guinea. It appears that dispersal into smaller islands such as New Britain, Tanimbar and the South Pacific has only been undertaken by species with a "flycatcher" body form.     

Systematics,evolution, and biogeography of the Pliocene stem meline badger Ferinestrix (Carnivora: Mustelidae)

Ferinestrix vorax is an extinct mustelid carnivoran of enigmatic relationships, known from a partial mandible and femur collected from the 3.2‐ and 3.6‐Myr‐old deposits of Hagerman Fossil Beds National Monument, Idaho, USA. Here, we report Ferinestrix rapax sp. nov. based on 80 remains of skull and dentition from a 3.1–3.6‐Myr‐old deposit of Udunga, Transbaikal, Russia. We demonstrate that Ferinestrix is a stem genus of the badger subfamily Melinae. This genus is distinctly larger and more carnivorous than any other total‐clade meline. We show that Ferinestrix originated in Asia and immigrated to North America no later than at the early (Zanclean) to late (Piacenzian) Pliocene transition, and that the North American F. vorax and Asian F. rapax underwent parallel evolution toward increased carnivory. © 2013 The Linnean Society of London, Zoological Journal of the Linnean Society, 2013, 167 , 208–226.