Systematics of African lowland rainforest Praomys (Rodentia, Muridae) based on molecular and craniometrical data

Representatives of the genus Praomys occur throughout the African intertropical zone. It is unclear how many species this genus contains, nor do we know the exact distribution ranges and phylogenetic relationships of these taxa. Using molecular (16S rRNA gene sequencing) and morphological (multivariate craniometry) analyses we clarify the taxonomy and phylogenetic relationships among the Praomys occurring in Africa's lowland tropical rainforests. We studied most species known from this area, based on specimens collected in seven countries (Guinea, Ivory Coast, Cameroon, Central African Republic, Gabon, Congo Republic and Democratic Republic of Congo). In our study, Praomys appears to be monophyletic. Our results identify two species complexes: the jacksoni complex includes at least two species ( P. jacksoni and P. mutoni ) and the tullbergi complex contains at least four species ( P. tullbergi , P. rostratus , P. misonnei , P. petteri ). Although the 16S rRNA gene appears insufficient to resolve the phylogenetic relationships among all the members of the tullbergi species complex, it is suitable for the identification of most of the studied species, and its use has allowed us to redefine the geographical limits of several species.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 145 , 539–553.     

Molecular systematics,taxonomy and biogeography of the genus Cavia (Rodentia: Caviidae)

Phylogenetic analyses were conducted on cytochrome b sequence data of the most geographically and taxonomically broad sampling of Cavia taxa to date. Primary objectives included providing the first extensive molecular phylogenetic framework for the genus, testing the taxonomic and systematic hypotheses of previous authors and providing insight into the evolutionary and biogeographic history of the genus. Support was found for the morphologically defined species C. aperea, C. tschudii, C. magna and C. fulgida and the taxonomic placement of taxa previously subject to conflicting taxonomic opinions (e.g. C. nana, C. anolaimae and C. guianae) was further resolved. Additionally, we elevate the Ecuadorian C. a. patzelti to species status, restrict the distributional limits and suggest taxonomic affiliations of some C. tschudii subspecies, and provide strong evidence for the geographic origin of guinea pig domestication. Finally, we provide an estimated evolutionary timeline for the genus Cavia, which appears to extend well into the late Miocene.     

Phylogeographic structure and regional history of Lemniscomys striatus (Rodentia: Muridae) in tropical Africa

Aim This study aims to elucidate the phylogeography of the murid rodent Lemniscomys striatus and to evaluate the relative roles of ecological change, habitat patchiness, rivers and geological barriers in structuring patterns of diversity. Location Sub‐Saharan Africa. Methods The extent of phylogeographic patterns and molecular genetic diversity (cytochrome b gene) were addressed in a survey of 128 individuals of L. striatus from 42 localities. Using maximum parsimony, maximum likelihood, Bayesian, network and genetic structure analyses, we inferred intraspecific relationships and tested hypotheses for historical patterns of gene flow within L. striatus. Results Our results identified four major geographical clades within L. striatus: a West African clade, a Benin‐Nigeria clade, a Central African clade, and an East African clade. Several subclades were identified within these four major clades. Restricted gene flow with isolation by distance was recorded, which is congruent with the low dispersal ability of such a small murid rodent. No clear signal of population expansion was detected within clades or subclades. Main conclusions The western rift system and the Volta and Niger rivers may have acted as long‐term extrinsic barriers to gene flow, resulting in the emergence of the four main clades of L. striatus. The observed pattern of mitochondrial variation observed within each clade probably results from late Pleistocene climatic and vegetation changes: during adverse conditions (forest expansion), L. striatus may have survived only in refugia, and then experienced range expansion under favourable conditions (savanna expansion).     

Patterns of speciation in two sibling species of Graomys (Rodentia,Cricetidae) based on mtDNA sequences

To increase our understanding of the speciation process occurred in the sibling species Graomys griseoflavus and Graomys centralis, a phylogeographic study was conducted based on sequences of a hypervariable segment of the mtDNA D‐loop region. The resulting haplotype phylogenetic network showed two well‐defined clusters, one for each species. The clusters were connected by two haplotypes from localities that are almost 300 km apart, one situated in the Monte eco‐region and the other, in the Chaco. This result is in agreement with a previous hypothesis about the geographical context in which the cladogenetic process occurred. A divergence time of 0.15–0.28 million years was estimated, which is consistent with a process of recent speciation. An amova test confirmed that at present gene flow between species does not exist. The mismatch distribution analyses suggest that the geographical and demographic expansion undergone by the species is related to the climatic events that occurred in the region during the Quaternary.     

Phylogenetic affinities and species limits within the genus Megadontomys (Rodentia: Cricetidae) based on mitochondrial sequence data

The evolutionary history of the genus Megadontomys, a group of mice allopatrically distributed along the cool‐humid forest in the highlands of México, is controversial. In this study, we examined phylogenetic relationships within the genus using sequences data from the complete mitochondrial cytochrome b gene. This information also allowed us to corroborate species limits, geographic boundaries of taxonomic entities and assess genetic variation within each taxon. The results of the phylogenetic analyses based on maximum likelihood, Bayesian inference and maximum parsimony were largely congruent in that M. nelsoni and M. thomasi were more closely related relative to M. cryophilus. These results are concordant with previous studies based on morphology and allozyme variation. However, testing of the alternative hypothesis of a closer evolutionary affinity between M. nelsoni–M. cryophilus did not produce a significantly less likely tree. The lack of unambiguous support towards one of these previously proposed contending hypotheses is congruent with the alternative scenario of an almost simultaneous diversification of the three species. Application of the phylogenetic species concept and the genetic species concept supports the recognition of three distinct taxonomic entities at the specific level. M. nelsoni inhabits the Sierra Madre Oriental (Hidalgo, Veracruz, and Puebla) including the Sierra Mazateca (Oaxaca); M. cryophilus is restricted to the Sierra de Juárez (Oaxaca); and M. thomasi occurs in portions of the Sierra Madre del Sur (Guerrero) and the Sierra Mixteca (Oaxaca). Our data show that M. thomasi is formed by two genetically distinct lineages that potentially may represent distinct Evolutionary Significant Units.     

The phylogeny of the Praomys complex (Rodentia: Muridae) and its phylogeographic implications

Among the African Murinae (Rodentia, Muridae), the Praomys complex, whose systematics has been studied by different approaches, has raised numerous taxonomic problems. Different taxa, namely Praomys, Mastomys , Myomys and Hylomyscus have been considered either as separate genera or subgenera of Praomys . In order to understand the relationships within the Praomys complex and to test the monophyly of the genus Praomys , a cladistic analysis was conducted, based on morpho-anatomical factors involving different species of Praomys , Mastomys , Myomys and Hylomyscus. The results indicate that the Praomys complex is monophyletic, as are the genera Hylomyscus , Mastomys and Myomys , whereas the genus Praomys appears paraphyletic. Indeed, a group of species including Praomys jacksoni was found to be more closely related to the genera Mastomys and Myomys than to a Praomys tullbergi -group. The biotopes and the distribution areas of the species were mapped on the phylogeny. It appears that the different clades each present a relative ecological cohesion and are arranged according to a gradient from closed to open habitats. From there, an evolutionary scenario is proposed for the emergence of the different clades and species of the genus Praomys sensu stricto.     

Mitochondrial phylogeny of African wood mice, genus Hylomyscus (Rodentia, Muridae): implications for their taxonomy and biogeography

This paper investigates the usefulness of two mitochondrial genes (16S rRNA and cytochrome b) to solve taxonomical difficulties within the genus Hylomyscus and to infer its evolutionary history. Both genes proved to be suitable molecular markers for diagnosis of Hylomyscus species. Nevertheless the resolving powers of these two genes differ, and with both markers (either analyzed singly or in combination), some nodes remain unresolved. This is probably related to the fact that the species emerged during a rapid diversification event that occurred 2-6 Myr ago (4-5 Myr ago for most divergence events). Our molecular data support the recognition of an "aeta" group, while the "alleni" and "parvus" groups are not fully supported. Based on tree topology and genetic divergence, two taxa generally recognized as subspecies should be elevated at the species level (H. simus and H. cf kaimosae). H. stella populations exhibit ancient haplotype segregation that may represent currently unrecognized allopatric species. The existence of cryptic species within H. parvus is questioned. Finally, three potentially new species may occur in West Central Africa. The Congo and Oubangui Rivers, as well as the Volta and Niger Rivers and/or the Dahomey gap could have formed effective barriers to Hylomyscus species dispersal, favoring their speciation in allopatry. The pronounced shifts in African climate during the late Pliocene and Miocene, which resulted in major changes in the distribution and composition of the vegetation, could have promoted speciation within the genus (refuge theory). Future reports should focus on the geographic distribution of Hylomyscus species in order to get a better understanding of the evolutionary history of the genus.     

Comparative phylogeography of two sibling species of forest‐dwelling rodent (Praomys rostratus and P. tullbergi) in West Africa: different reactions to past forest fragmentation

Two sibling species of the rodent genus Praomys occur in West African forests: P. tullbergi and P. rostratus. By sampling across their geographical ranges (459 individuals from 77 localities), we test the hypothesis that climatic oscillations during the Quaternary made an impact on the observed pattern of cytochrome b sequence variation. We show that, although these two species have parapatric geographical distributions, their phylogeographical histories are dissimilar, which could be related to their distinct ecological requirements. Since the arid phases of the Pleistocene were characterized by isolated forest patches, and intervening wetter periods by forest expansion, these changes in forest cover may be the common mechanism responsible for the observed phylogeographical patterns in both of these species. For example, in both species, most clades had either allopatric or parapatric geographical distributions; however, genetic diversity was much lower in P. tullbergi than in P. rostratus. The genetic pattern of P. tullbergi fits the refuge hypothesis, indicating that a very small number of populations survived in distinct forest blocks during the arid phases, then expanded again with forest recovery. In contrast, a number of populations of P. rostratus appear to have survived during the dry periods in more fragmented forest habitats, with varying levels of gene flow between these patches depending on climatic conditions and forest extent. In addition, historical variations of the West African hydrographic network could also have contributed to the pattern of genetic differentiation observed in both species.     

Genetic variation and evolution in the genus Apodemus (Muridae: Rodentia)

Genetic variation was studied using protein electrophoresis of 28–38 gene loci in 1347 specimens of Apodemus agrar-ius, A. peninsulae, A. flavicollis, A. sylvaticus, A. alpicola, A. uralensis, A . cf. hyrcanicus, A. hermonensis, A. m. mystacinus and A. m. epimelas , representing 121 populations from Europe, the Middle East, and North Africa. Mean values of heterozygosity per locus for each species ranged from 0.02 to 0.04. Mean values of Nel's genetic distance ( D ) between the taxa ranged from 0.06 (between A. flavicollis and A. alpicola) to 1.34 (between A. uralensis and A. agrarius ). The highest values of D were found between A. agrarius and other Apodemus species (0.62-1.34). These values correspond to those generally observed between genera in small mammals. Our data show that A. agrarius and A. peninsulae are sister species, well-differentiated from other taxa. High genetic distance between A. m. mystacinus and A. m. epimelas leads us to consider them distinct species and sister taxa to other Western Palaearctic species of the subgenus Sylvaemus . The data also suggest a recent separation of members of the latter group from a common ancestor, and subsequent rapid radiation, making it difficult to infer phylogenetic relationships. Some taxonomic implications of the results are discussed further.© 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 75 , 395–419.     

A species complex within the isopod genus Haploniscus (Crustacea: Malacostraca: Peracarida) from the Southern Ocean deep sea: a morphological and molecular approach

Seven new species of the genus Haploniscus from the deep Scotia and Weddell Seas are presented, combining morphological and molecular data (mitochondrial 16S rDNA and nuclear 18S rDNA). Haploniscus cassilatus sp. nov. , H. cucullus sp. nov. , H. weddellensis sp. nov. , H. procerus sp. nov. and H. kyrbasia sp. nov. are characterized by a prominent rostral process, the size and shape of which vary among species. The rostrum of H. microkorys sp. nov. is distinctly smaller than that of the former species, while H. nudifrons sp. nov. does not possess a rostrum. The status of the latter as separate species is obvious, owing to the stronger morphological differences. DNA was sequenced from three of the other five species. Genetic distances together with the more subtle morphological variation justify the erection of separate species. Overall morphological variations between these species are small yet noticeable and include, among others, the rostrum, the shape of the pleotelson and setation of pereopods. Our molecular data sets reveal detailed phylogenetic insights within the Haploniscus cucullus complex, supporting the monophyly of all species. We found p -distances of at least 0.0732 (16S rDNA) and 0.0140 (complete 18S rDNA) between pairs of species and show that both genes can be used as a marker for DNA taxonomy.  © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society , 2008, 152 , 655–706.     

Molecular phylogenetic analyses indicate paraphyly of the genus Hybomys (Rodentia: Muridae): Taxonomic implications

Widely distributed in Guineo‐Congolian forests, the genus Hybomys is represented by two species complexes (univittatus and trivirgatus), each restricted to one distinct forest block. In the last revision, these two species complexes were considered as distinct subgenera (Hybomys and Typomys). Previous morphological and karyological studies identified an important divergence between these two subgenera and raised the question of their taxonomic status (subgenus or genus). The number of species within this genus is also a matter of discussion: nine forms were described but only six (H. badius, H. basilii, H. lunaris, H. planifrons, H. trivirgatus, and H. univitttatus) are currently recognized as distinct species, the three others (H. pearcei, H. eisentrauti, and H. rufocanus) being considered as synonyms. The monophyly of the genus and its species have never been previously investigated with DNA sequence data. In this study, we combined mitochondrial and nuclear data (for a total of 3,264 nucleotide characters) to test the monophyly of Hybomys and to assess the specific status of H. eisentrauti and H. rufocanus. Our results highlight the paraphyly of the genus: members of the H. univittatus species complex appeared closely related to the genera Stochomys and Dephomys; representatives of H. trivirgatus are the sister clade of the node grouping Stochomys, Dephomys and member of the H. univittatus species complex. Combined with previous morphological findings, our results suggest that Typomys and Hybomys should be considered as two distinct genera. Based on tree topology and genetic distances, we propose to consider H. rufocanus as a valid species, distinct from H. univittaus, and to consider H. badius and H. eisentrauti as junior synonyms of H. rufocanus.     

Species limits within the Praomys delectorum group (Rodentia: Muridae: Murinae) of East Africa: a morphometric reassessment and biogeographical implications

We examined approximately 600 specimens that represent the Praomys delectorum species group (Muridae: Murinae: Praomyini), a rodent complex restricted to Afromontane landscapes in East Africa and currently viewed as a single species. Morphometric analyses of 21 population samples consistently disclosed cohesive patterns of craniodental differentiation that support the recognition of three species: Praomys delectorum Thomas, confined to extreme southern Malawi; P. melanotus Allen & Loveridge, found in highlands of south‐western Tanzania and contiguous northern Malawi; and P. taitae Heller (including octomastis Hatt), distributed in mountains and foothills of southern Kenya and northern and central Tanzania. Populations of the P. delectorum group are patchily distributed in moist montane forest, most collecting localities falling within 1000–2400 m, and their range collectively coincides with the Tanganyika–Nyasa Montane Forest Group sensu Moreau. Patterns of faunal similarity derived from distributions of 65 species of terrestrial small mammals recorded from Tanzania's highlands, including the Eastern Arc Mountains, demonstrated pronounced geographical discontinuities in montane associations but failed to uncover a prominent vicariant role for the Makambako Gap. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 165 , 420–469.     

Molecular identification of three sympatric species of wood mice (Apodemus sylvaticus,A. flavicollis,A. alpicola) in western Europe (Muridae: Rodentia)

The woodmouse (Apodemus sylvaticus) and yellow‐necked fieldmouse (Apodemus flavicollis) are sympatric and even syntopic in many regions throughout their European range. Their field discrimination on the basis of external characters is a real challenge for many fields of research. The problem is even more complicated in the Alpine chain where they live sympatrically with a third similar species: A. alpicola. A rapid and simple method is proposed to discriminate the three species in processing field‐collected biopsies as well as ethanol‐preserved museum samples.     

North-east Atlantic and Mediterranean species of the genus Buffonellaria (Bryozoa, Cheilostomata): implications for biodiversity and biogeography

In this study we revise the cheilostome bryozoan genus Buffonellaria Canu & Bassler, 1927 and its Mediterranean and north-east Atlantic species, thereby addressing several existing problems. First, a lectotype for the type species, Buffonellaria divergens (Smitt, 1873) from Florida, is chosen, which proves to be distinct from the European species. Second, the two hitherto established north-east Atlantic species [ Buffonellaria nebulosa ( Jullien & Calvet, 1903 ) and Buffonellaria porcellanum Arístegui Ruiz, 1987], are redescribed, which were poorly documented until now. Third, close inspection of material, collected from Spitsbergen to tropical West Africa, using scanning electron microscopy reveals that the actual number of species, all previously referred to either B. divergens or Stephanosella biaperta (Michelin, 1848), is distinctly greater in the north-east Atlantic than has been previously acknowledged. As a result, seven new species are introduced ( Buffonellaria acorensis sp. nov. , Buffonellaria antoniettae sp. nov. , Buffonellaria arctica sp. nov. , Buffonellaria harmelini sp. nov. , Buffonellaria jensi sp. nov. , Buffonellaria muriella sp. nov. , and Buffonellaria ritae sp. nov. ), whereas two are left in open nomenclature. With the increase in number of species, the extremely broad geographical range of distribution assumed for B. divergens breaks down to numerous restricted areas. However, although most species have only been reported from a single location, B. arctica sp. nov. seems to have a fairly wide distribution in the Arctic region.  © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society , 2008, 152 , 537–566.     

A new genus and species of rodent from the Brazilian Atlantic Forest (Rodentia: Cricetidae: Sigmodontinae: Oryzomyini), with comments on oryzomyine biogeography

We describe in this paper a new genus and species of cricetid rodent from the Atlantic Forest of Brazil, one of the most endangered eco‐regions of the world. The new form displays some but not all synapomorphies of the tribe Oryzomyini, but a suite of unique characteristics is also observed. This new forest rat possesses anatomical characteristics of arboreal taxa, such as very developed plantar pads, but was collected almost exclusively in pitfall traps. Phylogenetic analyses of morphological (integument, soft tissue, cranial, and dental characters) and molecular [nuclear – Interphotoreceptor retinoid binding protein (Irbp) – and mitochondrial – cytochrome b– genes] datasets using maximum likelihood and cladistic parsimony approaches corroborate the inclusion of the new taxon within oryzomyines. The analyses also place the new form as sister species to Eremoryzomys polius, an Andean rat endemic to the Maranon valley. This biogeographical pattern is unusual amongst small terrestrial vertebrates, as a review of the literature points to few other similar examples of Andean–Atlantic Forest pairings, in hylid frogs, Pionus parrots, and other sigmodontine rodents. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 161 , 357–390.     

Taxonomic review of the genus Trisopterus(Teleostei: Gadidae) with recognition of the capelan Trisopterus capelanus as a valid species

Trisopterus is demonstrated to be monophyletic, including four distinct species: T. capelanus, T. esmarkii, T. luscus and T. minutus. The capelan T. capelanus is resurrected from the synonymy of poor cod T. minutus, and is shown to be morphologically more similar to T. luscus, to which species it is also more closely related, indicated by a phylogenetic analysis presented here. A lectotype is designated for T. luscus. Trisopterus fasciatus, the type species of Trisopterus, is a junior synonym of T. luscus, and the lectotype of T. luscus is designated as the neotype of T. fasciatus. The lectotype of T. luscus is also designated as the neotype of Gadus barbatus. Gadus barbatus has priority over T. luscus but the name is suppressed by prevailing usage of T. luscus. A neotype is designated also for T. minutus. A phylogenetic analysis using mitochondrial cytochrome b, and a fragment of the nuclear rhodopsin gene, shows that T. capelanus and T. luscus are sister species, and in turn sister to a clade formed by T. minutus and T. esmarkii.     

Genetic,ecological, and behavioural divergence between two sibling snapping shrimp species (Crustacea: Decapoda: Alpheus)

Examination of genetic and ecological relationships within sibling species complexes can provide insights into species diversity and speciation processes. Alpheus angulatus and A. armillatus, two snapping shrimp species with overlapping ranges in the north-western Atlantic, are similar in morphology, exploit similar ecological niches and appear to represent recently diverged sibling species. We examined phylogenetic and ecological relationships between these two species with: (i) sequence data from two mitochondrial genes (16S rRNA and COI); (ii) data on potential differences in microhabitat distribution for A. armillatus and A. angulatus; and (iii) data from laboratory experiments on the level of reproductive isolation between the two species. DNA sequence data suggest A. armillatus and A. angulatus are sister species that diverged subsequent to the close of the Isthmus of Panama, and that haplotype diversity is lower in A. armillatus than in A. angulatus. Both species are distantly related to A. heterochaelis and A. estuariensis, two species with which A. angulatus shares some similarities in coloration. Ecological data on the distribution of A. angulatus and A. armillatus from two locations revealed differences in distribution of the two species between habitat patches, with each patch dominated by one or the other species. However, there was no apparent difference in distribution of the two species within habitat patches with respect to microhabitat location. Ecological data also revealed that heterospecific individuals often occur in close proximity (i.e. within metres or centimetres) where sympatric. Behavioural data indicated that these species are reproductively isolated, which is consistent with speciation in transient allopatry followed by post-divergence secondary contact. Our data further resolve taxonomic confusion between the sibling species, A. armillatus and A. angulatus, and suggest that sympatry in areas of range overlap and exploitation of similar ecological niches by these two recently diverged species have selected for high levels of behavioural incompatibility.