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.     

Cytochrome b-based phylogeny of the Praomys group (Rodentia,Murinae): a new African radiation?Phylogénie du groupe Praomys (Rodentia,Murinae) basée sur le cytochrome b : une nouvelle radiation africaine ?

Complete cytochrome b gene sequences allows, for the first time, establishing a nearly complete phylogeny among the Praomys group sensu lato. The genera Praomys, Mastomys and Stenocephalemys appear paraphyletic. Myomys is polyphyletic and this genus name probably needs to be restricted to its type species, M. verreauxii. The genera Zelotomys and Colomys appear as sister groups. Mastomys pernanus and Malacomys verschureni nest within the Praomys group, but their generic assignation must be further clarified. The genus Heimyscus appears closest to Praomys than to Hylomyscus. The different lineages probably result from an adaptive radiation at the end of the Miocene.     

Molecular phylogeny of Myomys/Stenocephalemys complex and its relationships with related African genera

Partial 16S rRNA mitochondrial gene sequences were used to infer the phylogenetic relationships among Stenocephalemys albocaudata, S. griseicauda and Myomys albipes, three closely related Ethiopian endemic murines and a selection of related species (Myomys daltoni, Praomys delectorum, Mastomys natalensis, Hylomyscus kaimosae, and Colomys goslingi) from other African regions. The obtained phylogeny confirms the close relationship between M. albipes and the two Stenocephalemys species, but it also suggests that both the genera Myomys and Stenocephalemys are paraphyletic, as M. albipes is closer to Stenocephalemys than to M. daltoni and S. griseicauda is more closely related to M. albipes than to S. albocaudata. These data, if confirmed, would argue that M. albipes should be renamed S. albipes. In conclusion, our study suggests that morphological similarity is not always a reliable measure for close genetic relationship in murines. Morphological similarity among species that evolved under similar ecological conditions can be the result of convergent evolution rather than a consequence of recent common ancestry.     

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.     

Chromosomal and protein evolution in morphologically similar species of Praomys sensu lato (Rodentia, Muridae)

Evidence of extensive chromosomal evolution in a biologically and economically important group of African murids of the Praomys/Mastomys complex was provided by examination of G- and C-band chromosomal data on P. coucha (2n = 32), P. fumatus (2n = 38), P. hildebrandti (2n = 32), P. jacksoni (2n = 28), P. misonnei (2n = 36), and P. cf. tullbergi (2n = 35). A coding system was developed for the chromosomal characters, and analyses were performed by a computer program to find the shortest tree with a minimum of 35 autosomal rearrangements (pericentric inversions, complex translocations, centric fusions, centric fissions, tandem fusions, euchromatic additions, and heterochromatic additions). The resulting phylogenetic hypothesis differs from traditionally accepted hypotheses regarding this complex group of rodents. The cytogenetic data show that 1) there is no support for the dichotomy of Mastomys/Praomys previously based on morphology, 2) the 2n = 32 species from eastern Africa (P. hildebrandti) is distinct from the 2n = 32 species from southern Africa (P. natalensis), and 3) there is a close association between P. jacksoni and P. cf. tullbergi. Polyacrylamide gel electrophoresis of liver membrane proteins demonstrated few differences in protein mobilities between species and even fewer between individuals of the same species taken from different habitats and localities in Kenya. Monoclonal antibodies produced against liver proteins of one species and tested for reactivity to other species confirmed the evolutionary similarity of species of this complex. This immunologic approach may provide a robust data set for future phylogenetic studies of muroid rodents.(ABSTRACT TRUNCATED AT 250 WORDS)     

Systematics of Maxomys Sody, 1936. (Rodentia: Muridae: Murinae): DNA/DNA hybridization studies of some Borneo-Javan species and allied Sundaic and Australo-Papuan genera

We compared five species of the murine genus Maxomys and representatives of nine other murid genera in a complete 15 × 15 DNA-hybridization matrix. FITCH trees were calculated for the entire suite of taxa and for subsets including only the five Maxomys and these together with the four nearest outgroups. All trees were validated by 'bootstrapping' and by jackknifing, performing both single- and multiple-deletions of taxa. The full 15 times 15 data set indicated a sister-group relationship between Maxomys and two pairs of genera ( Sundamys-Rattus sensu stricto and Mviventer-Leopoldamys ) that are more closely related to each other than to Maxomys; addition of data on Bandicota and Berylmys from another recent DNA-hybridization study confirmed that these genera are successive sister-taxa to the Sundamys-Rattus pair. Mus-Myomys and Uromys-Melomys were each distinct lineages from the above grouping of Rattus sensu lata species, and from the putative outgroup sigmodontine Peromyscus, but the interrelations of the three murine clades were unresolved. Within Maxomys, M. surifer and M. bartelsii are a related pair, and M. ochraceiventer probably forms an unresolved trio with M. rajah and M. whiteheadi. Calibration of a tree generated from saturation-corrected distances against a likely divergence-date of 12.2 Mybp for the separation of Mus and Rattus confirms a high rate of single-copy DNA change in murids (2.1%/Myr); and suggests that Sigmodontinae and Murinae diverged at around 15.3 Mybp, that Maxomys and the group of six other Rattus sensu lato separated approximately 7.6 Myr ago, and that Maxomys began to diversify 4.8 Myr ago.     

On the morphology of spermatozoa in some African murines (Rodentia, Muridae): the taxonomic and phylogenetic aspects

A comparative analysis of sperm-head morphology and measurements in 17 species from nine genera of African Murinae: Rattus rattus, Mastomys coucha, M. huberti, M. erythroleucus, Mastomys sp. 2, Praomys albipes, P. fumatus, Mus mahomet, Arvicanthis somalicus, A. abyssinicus, A. dembeensis, Arvicanthis sp., Lemniscomys macculus, Pelomys harringtoni, Acomys cahirinus, Acomys sp., Uranomys ruddi, was carried out. Spermatozoa of all examined species are of the same basic type. They consist of an asymmetrical head, falciform or scythelike in shape, and a tail attached to the ventrocaudal surface of the head. There are great interspecific differences in sperm morphology and size. The significance of this variation for estimation of taxonomic aspects and phylogenetic relationships among the species, as well as between them and other groups, is discussed. The sperm morphology supports a close evolutionary relationship among the genera Lemniscomys and Arvicanthis. It also indicates that Pelomys is distinctive. The relationships between Acomys and Uranomys are discussed.     

Chromosomal phylogeny of Muridae: a study of 10 genera

The karyotypes of 10 different species of the family Muridae (Acomys airensis, Arvicanthis niloticus, Hylomyscus stella, Malacomys longipes, Mastomys huberti, Myomys daltoni, Mus musculus, Rattus norvegicus, Thamnomys gazellae, and Uranomys ruddi) are compared by different banding techniques. From a reconstruction of the presumed ancestral karyotype of the Muridae the sequence of the various rearrangements leading to the present karyotypes is proposed in order to determine their phylogenetic relationships. In particular, the present karyotypes of the mouse and rat differ from the ancestral one by at least 12 and 7 rearrangements, respectively. A clear tendency for accumulation of a specific type of rearrangement in a given branch of the cladogram is observed. In regard to the mouse, a large number of translocations, with break points situated in the proximal part of the long arms, have occurred, which conserved the acrocentric form of the ancestral chromosomes but led to multiple recombinations of the bands.     

Diversity and distribution of murid rodent populations between forest and derived savanna sites within south western Nigeria

Southward encroachment of the derived savanna zone with its attendant denudation of the rainforest in Nigeria adversely affects the diversity and distribution of mammals, particularly murid rodents. This study was carried out to establish the identities of murid rodent populations and to compare their diversity between forest and derived savanna sites within south western Nigeria. Identification of captured specimens was carried out by multivariate statistics of body and skull measurements, and also by analysis of certain discrete characters. Six species were identified: Praomys tullbergi, Hylomyscus stella, Mastomys natalensis, Arvicanthis rufinus, Lemniscomys striatus and Lophuromys sikapusi. Forest sites possessed a higher Shannon’s diversity index (1.685) than the derived savanna sites (0.978), containing a higher number of taxa and larger abundance of murid rodents. This underscores the negative impact that encroachment of the derived savanna has upon diversity of animals in areas that were once forest.     

DNA barcoding of Murinae (Rodentia: Muridae) and Arvicolinae (Rodentia: Cricetidae) distributed in China

Identification of rodents is very difficult mainly due to high similarities in morphology and controversial taxonomy. In this study, mitochondrial cytochrome oxidase subunit I (COI) was used as DNA barcode to identify the Murinae and Arvicolinae species distributed in China and to facilitate the systematics studies of Rodentia. In total, 242 sequences (31 species, 11 genera) from Murinae and 130 sequences (23 species, 6 genera) from Arvicolinae were investigated, of which 90 individuals were novel. Genetic distance, threshold method, tree‐based method, online BLAST and BLOG were employed to analyse the data sets. There was no obvious barcode gap. The average K2P distance within species and genera was 2.10% and 12.61% in Murinae, and 2.86% and 11.80% in Arvicolinae, respectively. The optimal threshold was 5.62% for Murinae and 3.34% for Arvicolinae. All phylogenetic trees exhibited similar topology and could distinguish 90.32% of surveyed species in Murinae and 82.60% in Arvicolinae with high support values. BLAST analyses yielded similar results with identification success rates of 92.15% and 93.85% for Murinae and Arvicolinae, respectively. BLOG successfully authenticated 100% of detected species except Leopoldamys edwardsi based on the latest taxonomic revision. Our results support the species status of recently recognized Micromys erythrotis, Eothenomys tarquinius and E. hintoni and confirm the important roles of comprehensive taxonomy and accurate morphological identification in DNA barcoding studies. We believe that, when proper analytic methods are applied or combined, DNA barcoding could serve as an accurate and effective species identification approach for Murinae and Arvicolinae based on a proper taxonomic framework.     

Phylogeny of the genus Apodemus with a special emphasis on the subgenus Sylvaemus using the nuclear IRBP gene and two mitochondrial markers: cytochrome b and 12S rRNA

Phylogenetic relationships among 17 extant species of Murinae, with special reference to the genus Apodemus, were investigated using sequence data from the nuclear protein-coding gene IRBP (15 species) and the two mitochondrial genes cytochrome b and 12S rRNA (17 species). The analysis of the three genes does not resolve the relationships between Mus, Apodemus, and Rattus but separates Micromys from these three genera. The analysis of the two mitochondrial regions supported an association between Apodemus and Tokudaia and indicated that these two genera are more closely related to Mus than to Rattus or Micromys. Within Apodemus, the mitochondrial data sets indicated that 8 of the 9 species analyzed can be sorted into two main groups: an Apodemus group, with A. agrarius, semotus, and peninsulae, and a Sylvaemus group, with uralensis, flavicollis, alpicola, sylvaticus, and hermonensis. The position of Apodemus mystacinus is ambiguous and might be either included in Sylvaemus or considered a distinct subgenus, Karstomys, more closely related to Sylvaemus than to Apodemus. Estimation of the divergence time for these taxa suggests a separation between 7 and 8 My ago for the three groups (mystacinus and the two subgenera Apodemus and Sylvaemus). Within each subgenus, divergence times are between 5.4 and 6 My for Apodemus and between 2.2 and 3.5 My for Sylvaemus and mystacinus.     

Confrontation of morphological and molecular data: the Praomys group (Rodentia, Murinae) as a case of adaptive convergences and morphological stasis

Phylogenetic relationships in a group of 21 African rodent species designated as the Praomys group (Murinae) were investigated using morphological characters and sequence data from the complete mitochondrial cytochrome b gene and nuclear IRBP gene fragment (840bp). The molecular results confirm the monophyly of the Praomys group, including the species Malacomys verschureni, while the other Malacomys species appear very divergent. The basal relationships within the Praomys group are poorly resolved, suggesting a rapid radiation at about 7-9 million years ago based on genetic divergence rates calibrated from the fossil record. Discrepancies between molecular and morphological results probably reflect of numerous convergences as well as variations in the rates of morphological evolution among lineages. Reconstructions of the ancestral character states suggest a savannah origin for the Praomys group, along with some morphological traits conserved by stasis in savannah taxa. At the same time, forest taxa seem to be characterized by an accelerated morphological evolution, with acquisition of convergent adaptive characters.     

Distribution of small rodents and shrews in a lowland rain forest zone of Nigeria, with observations on their reproductive biology

This paper describes the habitat preferences of small rodents and shrews from April 1984 to March 1985 in four types of vegetation in a lowland rain forest zone of Nigeria. The vegetation types were high forest, scrub, farmlands and teak plantations in Ogba Forest Reserve, Nigeria. A collection of 359 small mammals, consisting of six species of small rodents and four species of shrews, was made in the study area. The small rodents and their percentage composition were Mus musculoides (26.2%), Praomys tullbergi (16.2%), Mastomys natalensis (7.0%), Lophuromys sikapusi (4.5%), Lemniscomys striatus (2.5%), and Arvicanthis niloticus (1.1%). The shrews were Crocidura nigeriae (19.5%), C. crossei (12.3%), C.grandiceps (7.8%) and C. ftavescens manni (3.1%). The vegetational distribution of the small rodents and shrews is discussed. In the high forest, seven species of small mammals were caught; these were Crocidura nigeriae, C. crossei, C.grandiceps, C.flavescens manni, Mus musculoides, Praomys tullbergi and Lophuromys sikapusi. Mus musculoides, Crocidura nigeriae and Praomys tullbergi were more abundant in the high forest. All the species of small mammals recorded in the study area, except Arvicanthis niloticus, were found in the scrub. In the farmlands, all ten species of small mammals were trapped. Mus musculoides was the commonest species in the farmlands. Three species of shrews, Crocidura crossei, C. grandiceps and C.flavescens manni, and also two species of small rodents, Mus musculoides and Mastomys natalensis were caught in the teak plantation. The reproductive data for Mus musculoides, Crocidura nigeriae and Praomys tullbergi show that breeding in these species occurs throughout most months of the year. Fecundity appears to be maximal in the wet season in Mus musculoides and Crocidura nigeriae and in the dry season in the case of Praomys tullbergi.     

Mitochondrial DNA phylogenetic structuring suggests similarity between two morphologically plastic genera of Australian freshwater mussels (Unionoida: Hyriidae)

In the Lake Eyre Basin, the Australian hyriid genus Velesunio is represented by three undescribed species, each of which are highly divergent genetically, but morphologically similar to Velesunio wilsonii (Lea 1859). A fourth species, Velesunio ambiguus (Philippi 1847), occurs not only in the Lake Eyre Basin but throughout much of eastern Australia, including the Murray-Darling Basin. In this study, we show that another hyriid, Alathyria jacksoni (Iredale 1934), which is sympatric with V. ambiguus, is genetically deeply nested within the Velesunio species complex, such that the genus Velesunio is paraphyletic with respect to A. jacksoni. Moreover, our mitochondrial phylogenetic reconstructions indicate that A. jacksoni is closely allied to one of the cryptic Velesunio species occurring in the Lake Eyre Basin, but distinct from V. ambiguus and the two other Velesunio species. These data suggest that the genera Alathyria and Velesunio are in need of revision. The shells of A. jacksoni and Velesunio spp. vary with local conditions and sometimes are difficult to distinguish. Our analyses also show that shell characters of these taxa do not closely match the phylogenetic data, and it appears that the traditional taxonomic emphasis on these plastic characters has obscured evolutionary relationships between these, and possibly other, Hyriidae.     

On the Parasitic Protozoa of Liberia. V. Coccidia of Liberian Rodents.

SYNOPSIS. In a survey of coccidia of rodents from Harbel, Libera, the following species of Eimeria were found and described: E. musculoidei n. sp. from Mus (Leggada) musculoides; E. separata Becker & Hall, 1931 from Rattus (Dephomys) defua (new host record); E. praomysis n. sp. from Rattus (Praomys) tullbergi rostratus; E. dasymysis n. sp. from Dasymys incomptus rufulus; E. lemniscomysis n. sp. from Lemniscomys striatus striatus; and E. lophuromysis n. sp., E. sikapusii n. sp., E. liberiensis n. sp., E. harbelensis n. sp., E. africana n. sp., and E. kruidenieri n. sp. from Lophuromys sikapusi sikapusi.     

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.     

Multigene phylogeny of the Old World mice, Murinae, reveals distinct geographic lineages and the declining utility of mitochondrial genes compared to nuclear genes

Despite its great diversity and biomedical importance, the rodent subfamily Murinae is poorly resolved phylogenetically. We present the first cladistic analysis sampling multiple representatives of most major groups based on DNA sequence for three nuclear (GHR, RAG1, and AP5) and one mitochondrial (COII and parts of COI and ATPase 8) fragments. Analyzed separately, the four partitions agree broadly with each other and the combined analysis. The basal split is between a clade of Philippine Old Endemics and all remaining murines. Within the latter, rapid radiation led to at least seven geographically distinct lineages, including a Southeast Asian Rattus clade; a diverse Australo-Papuan and Philippine clade; an African arvicanthine group including the otomyines; an African Praomys group; and three independent genera from Africa and Asia, Mus, Apodemus, and Malacomys. The murines appear to have originated in Southeast Asia and then rapidly expanded across all of the Old World. Both nuclear exons provide robust support at all levels. In contrast, the bootstrap proportions from mitochondrial data decline rapidly with increasing depth in the tree, together suggesting that nuclear genes may be more useful even for relatively recent divergences (< 10MYA).     

A molecular phylogenetic framework for the Ryukyu endemic rodents Tokudaia osimensis and Diplothrix legata

Diplothrix legata and Tokudaia osimensis are rodent species endemic to the Ryukyu Islands (Okinawa area) of the southern part of Japan. To place them within a phylogenetic framework of Asian rodent species, we examined the molecular relationships among these species and Apodemus, Micromys, Mus, and Rattus, using an analysis of mitochondrial cytochrome b gene (1140 bp) and nuclear IRBP gene (782 bp) sequences. The results indicate that the four main murine genera and Tokudaia diverged at similar evolutionary times, namely at the time of the radiation of Murinae, 14 to 40 million years ago, depending on different concepts for the divergence times of Rattus and Mus. In contrast, Diplothrix separated from the Rattus group more recently. Our results indicate that D. legata and the Rattus group diverged at 20-30% of the divergence time of Rattus and Mus under the assumption of the molecular clock, suggesting that D. legata established its lineage at least by the beginning of the Pleistocene era. It is thus evident that the Okinawa area preserves rare indigenous species with various levels of genetic endemicity.     

Molecular evolutionary relationships of three genera of nesomyinae,endemic rodent taxa from madagascar

The relationships of Nesomyinae, a group of murid rodents endemic to the island of Madagascar, were investigated with two comparative molecular approaches. Compared to those of other muroid rodents representing Murinae, Cricetinae, Cricetomyinae. Arvicolinae, and Sigmodontinae, complete sequences of the 12S rRNA mitochondrial gene suggest that the Malagasy nesomyinesMacrotarsomys andNesomys are monophyletic and that their sister-group among the taxa analyzed isCricetomys. A limited series of DNA/DNA hybridization experiments extends these observations to a third nesomyine genus,Eliurus, and a second cricetomyine taxon,Saccostomus. By relating the amounts of overall genomic divergence with geological time as calibrated by theMus/Rattus dichotomy estimated at 12–14 My, the oldest within-Nesomyinae dichotomy is estimated to be 10.8 to 12.6 My. Thus, these three genera of Malagasy nesomyine rodents appear to be a rather ancient offshoot from African ancestors whose Recent relatives are Cricetomyinae. This preliminary observation should be confirmed by sampling additional genera of nesomyines and additional representatives for other subfamilies of African muroids.     

Satellite DNA as a phylogenetic marker: case study of three genera of the Murinae subfamily

Satellite DNA (satDNA) represent tens percent of any of the vertebrate genome. Still, a complete set of sat-DNA fragments is not determined for either species. It is known that some genus with species-specific modifications possess a satDNA characteristic for the genus. So, satDNA was used as a phylogenetic marker in some cases when precise satDNA fragment was cloned. We used the probe of the whole pericentromeric region and 4 cloned satDNA fragments of Mus musculus in order to consider probes value for phylogenesis of 3 Murinae genera. Fluorescent in situ hybridization (FISH) revealed similar pattern on metaphase spreads inside genus Mus, though some difference was noted. None of the satDNA fragment gave signal in the centromeric region on chromosomes from genera Sylvaemus and Apodemus. These data are in agreement with those on satDNA fragments in the genome determined by dot-blot hybridization: M musculus satDNA fragments are absent in the genomes of both remote genera while they are present in the genomes of the genera Mus, though in different amounts. SatDNA of each genera should be cloned for the phylogenetic purposes.