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The phylogenetic position of "Acomyinae" (Rodentia, Mammalia) as sister group of a Murinae + Gerbillinae clade: evidence from the nuclear ribonuclease gene.

The phylogenetic relationships of Acomys and Uranomys within Muridae were investigated using nuclear pancreatic ribonuclease A gene sequences. The various kinds of substitutions in the data matrix (15 taxa x 375 nucleotides) were examined for saturation, in order to apply a weighted parsimony approach. Phylogenies were derived by maximum parsimony (weighted and unweighted) and maximum likelihood procedures, using a dormouse (Gliridae) as outgroup. Maximum likelihood gave the most robust results. All analyses cluster some traditional taxa with a strong robustness, such as three species of the genus Mus, two South-East Asian rats, and two genera in each of the gerbil and vole families. When analyzed with those of other murid rodents representing Murinae, Gerbillinae, Arvicolinae, Cricetinae, and Sigmodontinae, sequences of the ribonuclease gene suggest that Acomys and Uranomys constitute a monophyletic clade at the subfamily level, denoted "Acomyinae." The relationships between the six subfamilies of Muridae appear poorly resolved, except for a clade uniting Murinae, Acomyinae, and Gerbillinae. Within this clade, the sister group of Acomyinae could not be identified, as the branch length defining a Gerbillinae + Murinae cluster is extremely short. The poor resolution of our phylogenetic inferences is probably the result of two confounding factors, namely the limited size of the pancreatic ribonuclease sequence and the probable short time intervals during the radiation of the six murid subfamilies involved in this study.

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Molecular evidence that the spiny mouse (Acomys) is more closely related to gerbils (Gerbillinae) than to true mice (Murinae).

Spiny mice of the genus Acomys traditionally have been classified as members of the Murinae, a subfamily of rodents that also includes rats and mice with which spiny mice share a complex set of morphological characters, including a unique molar pattern. The origin and evolution of this molar pattern, documented by many fossils from Southern Asia, support the hypothesis of the monophyly of Acomys and all other Murinae. This view has been challenged by immunological studies that have suggested that Acomys is as distantly related to mice (Mus) as are other subfamilies (e.g., hamsters: Cricetinae) of the muroid rodents. We present molecular evidence derived from DNA.DNA hybridization data that indicate that the spiny mouse Acomys and two African genera of Murinae, Uranomys and Lophuromys, constitute a monophyletic clade, a view that was recently suggested on the basis of dental characters. However, our DNA.DNA hybridization data also indicate that the spiny mice (Acomys) are more closely related to gerbils (Gerbillinae) than to the true mice and rats (Murinae) with which they have been classified. Because Acomys and the brush-furred mice Uranomys and Lophuromys share no derived morphological characters with the Gerbillinae, their murine morphology must have evolved by convergence, including the molar pattern previously considered to support the monophyly of the Murinae.

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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.

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Evolution of rRNA gene clusters and telomeric repeats during explosive genome repatterning in TATERILLUS X (Rodentia, Gerbillinae).

A survey of 28S and 5S rRNA gene clusters, and telomeric repeats was performed using single and double FISH in the Taterillus genus (Rodentia, Muridae, Gerbillinae). Taterillus was previously demonstrated to have undergone a very recent and extensive chromosomal evolution. Our FISH results demonstrate that rRNA genes can vary in location and number irrespective of the phylogenetic relationships. Telomeric repeats were detected in pericentromeric and interstitial regions of several chromosomes, thus providing nonambiguous evolutionary footprints of Robertsonian and tandem translocation events. These footprints are discussed in reference to the molecular process of these karyotypical changes. Also, examples of colocation of rDNA clusters and telomeric repeats lend support to their possible involvement in nucleolus formation. Finally, the presence of rRNA genes, and the extensive amplification of telomeric repeats at specific loci within a double X-autosome translocated element which were not observed on the homologous Y1 and Y2, served as basis for an epigenomic hypothesis on X-autosome translocation viability in mammals.

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The origin of multiple sex chromosomes in the gerbil Gerbillus gerbillus (Rodentia: Gerbillinae).

The sex chromosomes of the partly sympatric species of gerbils Gerbillus pyramidum and G. gerbillus (Mammalia: Gerbillinae) were investigated by a variety of light- and electron-microscope methods, including DNA replication banding and synaptonemal complex (SC) techniques. The sex-chromosome mechanism of G. pyramidum is of the maleXY:femaleXX type, whereas that of G. gerbillus is of the less common maleXY1Y2:femaleXX system. The results include the demonstration that the X chromosomes of both species are compound. One segment is added to the X chromosome of G. pyramidum, leading to an increase in length from the standard 5% to approximately 7.3%, whereas two different extra segments increase the length of the X chromosome of G. gerbillus to approximately 11% of the length of the haploid genome. In both cases the extra material is autosomal and is also represented in the respective Y chromosomes. Classifying heterochromatin by the variation in staining quality was helpful in elucidating the possible origin of the different chromosome segments, including the pericentromeric regions. Observations on meiotic chromosome pairing and chiasma formation have confirmed the homologies established by band comparisons. The occurrence of chiasmata between the sex chromosomes supports the autosomal origin of the pairing segments. These and other findings have been interpreted in the framework of a multistep evolutionary model. This sequence starts from a hypothetical pair of sex chromosomes, the X element of which amounts to 5% of the haploid genome, and leads through three translocations involving two pairs of autosomes and one pericentric inversion to the most complex situation of this series, manifested in G. gerbillus. The adaptive value, if any, of autosome incorporation into the sex chromosomes repeatedly occurring here is unknown. It is, however, a remarkable fact that in one species, G. gerbillus, the complex sex-chromosome constitution is conserved over vast geographic distances, and in the other, G. pyramidum, the compound X and Y chromosomes withstand change in the face of extreme autosome restructuring.

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A finding of the XX/XY1Y2 sex-chromosome system in Taterillus arenarius (Gerbillinae, Rodentia) and its phylogenetic implications.

A chromosome banding study (R- and C-bands) of a male Taterillus arenarius (Rodentia, Gerbillinae) specimen from Mauritania revealed the presence of an XX/XY1Y2 sex-chromosome system in the karyotype, as found previously in three other congeneric species. This finding allowed us to resolve the phylogenetic affinities of this species within the genus and to propose an evolutionary scenario leading to the formation of the species with an XX/XY1Y2 sex-chromosome system. A review of chromosome data in Taterillus suggests that there may be more species in the genus than hitherto recognized.

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Cyclic transmission of Sarcocystis gerbilliechis n. sp. by the Arabian saw-scaled viper, Echis coloratus, to rodents of the subfamily gerbillinae.

Infection experiments with rodents and snakes were performed to elucidate the life cycle of a Sarcocystis isolate found in an Arabian saw-scaled viper, Echis coloratus. Oocytes in feces of the naturally infected and of 2 experimentally infected Arabian saw-scaled vipers were already sporulated and contained 2 sporocysts each, measuring 12.7 (12.3-13.3) microns x 11.0 (10.7-11.4) microns. After oral inoculation of various rodent species with these sporocysts, sarcocysts developed in the esophagus and skeletal muscles of gerbils and related genera. Mature sarcocysts were filiform in shape and reached a maximum length of 11.7 mm after 5 mo postinoculation (PI), whereas the width did not exceed 190 microns. The primary cyst wall formed small, knoblike protrusions, which were up to 180 nm long and 120 nm wide. Mature schizonts were present in the liver and other organs of gerbils between 11 and 14 days PI. After inoculation of vipers of 3 different genera with mature sarcocysts from gerbils, oocysts developed in the intestine of Arabian saw-scaled vipers. A comparison of these data with those from previously described Sarcocystis species with snake-rodent life cycles suggests that Sarcocystis gerbilliechis is a new species.

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Evidence for the existence of two distinct species: Psammomys obesus and Psammomys vexillaris within the sand rats (Rodentia, Gerbillinae), reservoirs of cutaneous leishmaniasis in Tunisia.

A thorough taxonomic knowledge about putative animal reservoirs of transmissible diseases is an absolute prerequisite to any ecological investigation and epidemiological survey of zoonoses. Indeed, accurate identification of these reservoirs is essential for predicting species-specific population outbreaks and therefore to develop accurate ecological control strategies. The systematic status of sand rats (genus Psammomys) remains unclear despite the pivotal role of these rodents in the epidemiology of Zoonotic Cutaneous Leishmaniasis (ZCL) disease as sand rats are the main known reservoir hosts of the protozoan parasite Leishmania major. In the present work, we expose morphological, biochemical, genetic and cytogenetic evidence supporting the identification of at least two cryptic species within the genus Psammomys in Tunisia. First, significant morphometric differences were observed and were correlated associated with external features and biogeographic origins. Second, differences in patterns of two isoenzymic systems (Glutamate Oxaloacetate Transaminase (GOT) and 6-PhosphoGluconate Dehydrogenase (6PGD)) were found, which makes it possible to amount these isoenzyme characters to two diagnostic loci. Third, based on the mitochondrial cytochrome b (cyt b) gene, a high magnitude of genetic distance (13.89%) was also observed. Fourth, cytogenetic analysis showed that these two populations groups differ in their diploid chromosome numbers, i.e. 2N=46 versus 2N=48. We consider that all these variations are enough important to be considered as demonstrative and we propose that these two lineages should be considered as two distinct species that we refer to the fat sand rat Psammomys obesus Cretzschmar, 1828 and the thin sand rat Psammomys vexillaris Thomas, 1925. Implications of such results on the eco-epidemiology of ZCL in Tunisia are discussed.

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Mitochondrial phylogeny reveals differential modes of chromosomal evolution in the genus Tatera (Rodentia: Gerbillinae) in Africa.

The African gerbils of the genus Tatera are widespread and abundant throughout sub-Saharan Africa. There is still today a certain controversy concerning the taxonomy of these rodents and very few attempts have been made to assess their systematic relationships. The present paper introduces findings based on the partial sequences of cytochrome b (495 bp) and the 16S rRNA (469 bp) mitochondrial genes of six (T. robusta, T. nigricauda, T. vicina, T. leucogaster, T. valida, and T. kempi) species together with two additional taxa. We also report the karyotypes of T. vicina and T. leucogaster. We propose that T. vicina should be considered as a valid species and show the monophyly of the robusta species group, with the exclusion of T. leucogaster. Our results show there is a different chromosomal evolutionary pattern within the two major lineages, which is recognizable through molecular phylogenetics. One is characterized by karyotype stability and the other by a considerable number of chromosomal rearrangements. The lineage divergence coincides with the formation of the East African Rift. The processes that led to the origin of the East African species seem to be related to the subsequent climatic changes, which caused cyclic contraction and expansion of the savannah biomes. Furthermore, geological activities that characterized East Africa during Plio-Pleistocene may also have contributed to lineage divergence.

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Recent radiation in West African Taterillus (Rodentia, Gerbillinae): the concerted role of chromosome and climatic changes.

West African gerbils of the genus Taterillus constitute a complex of seven sibling species distributed from sudano-guinean to saharo-sahelian regions. They display radically rearranged karyotypes despite low genic divergence and a very recent differentiation, that is, within the last 0.4 Myr for the six most derived species. We here provide a comparison of the seven specific karyotypes and perform a cladistic analysis using chromosomal rearrangements character states. When a posteriori polarized mutations were mapped onto the phylogenetic tree, 38 rearrangements were identified as fixed during the evolution of these rodents. This makes Taterillus one of the most striking examples of accelerated chromosomal evolution within placental mammals. Taking into account the types of chromosomal changes involved, divergence times between lineages, genetic distances, as well as reassessed geographic distributions, we suggest that (1) speciation in West African Taterillus was driven by chromosomal changes, and (2) the paleoclimatic oscillations of the Sahara desert have played a major role in their evolution. In particular, elevated plasticity of the Taterillus genome, as suggested by the patterns observed for some repetitive elements, would have led to a higher probability of mutation. We hypothesize that the process underpinning cladogenesis most probably involved highly underdominant genomic rearrangements that were fixed following pronounced populational bottlenecks resulting from drastic climatic and subsequent environmental changes. Major African rivers formed significant barriers to dispersal, limiting expansion during the more moist and so favorable periods. This scenario would explain the current parapatric species distributions and their relationship to the West African hydrographic features.

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Explosive chromosome evolution and speciation in the gerbil genus Taterillus (Rodentia, Gerbillinae): a case of two new cryptic species.

The five morphologically sibling gerbil species of the genus Taterillus in West Africa were first identified from karyotypes. These species possess an XX/XY(1)Y(2) sex-chromosome system and are characterized by significant karyotypic reorganization, thus making them a suitable model for studying the role of chromosomal rearrangements in the speciation process. We present here a description of two new cytotypes, Taterillus sp. 1 and Taterillus sp. 2, from the Lake Chad area, the former having a 2n = 22/23, NFa = 40, and the latter 2n = 24/25, NFa = 44. Comparison of their G- and C- banding patterns with those of T. pygargus (2n = 22/23, NFa = 38/40), examined in an earlier paper, revealed that all three species differ from each other by 7 to 11 chromosomal rearrangements, comprising tandem translocations, pericentric inversions, and Robertsonian metacentrics displaying monobrachial homology. Meiotic configurations formed in potential hybrids among any of these three forms would consist of complex rings and chains, alone or in combination, resulting, as expected, in a significant disruption of gametogenesis. These results provide support for assigning Taterillus sp. 1 and Taterillus sp. 2 to two different biological species, which, as demonstrated by our preliminary molecular studies, would have emerged recently. Possible factors responsible for the rapid karyotypic evolution and speciation in this West African gerbil complex are discussed.

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Unusually extensive karyotype reorganization in four congeneric Gerbillus species (Muridae: Gerbillinae).

Comparative analysis of the G- and C-banding patterns in four morphologically poorly differentiated Gerbillus species (G. pyramidum, G. perpallidus, G. tarabuli and G. occiduus) was carried out. These gerbils have similar karyotype morphology with 2n and NF equal to 38/76, 40/76, 40/78 and 40/80, respectively. Our study revealed that possibly 70 Robertsonian (Rb) fusions, two pericentric inversions, one tandem translocation and at least 13 non-identified rearrangements have occurred during the karyotypic evolution of these species. The number of chromosomal changes by which any of these species differ from each other ranges from 33 to 49. One Rb fusion was common to two of the species, with only a single autosome-gonosome translocation shared by all four, suggesting a monophyletic origin of these karyotypically highly divergent species. Based on the chromosomal data obtained here, the systematic and geographic implications for these North African species are also discussed.

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[Biotope distribution of two gerbil species (Meriones meridianus and M. tamariscinus) (Rodentia, Gerbillinae) in southern Kalmykia].

Our studies have been conducted in Southern Kalmykia (Black Lands) in 1994-1996. The data obtained on the same territories in 1981-1983, as well as in 1997-1998 were used for comparison. Over the studied period, the size of Meriones tamariscinus population underwent a significant increase. In the early 1980s, the colonies of this species were found only on sandy dunes covered with tamarix. At the present time, M. tamariscinus populations have spread over stabilized sands, abandoned shepherd camps, and riparian thickets of tamarix along the shores of lakes. Both gerbils species (M. meridianus and M. tamariscinus) establish joint colonies in all biotopes, but not in riparian tamarix thickets near lakes, where only Meriones tamariscinus have been found. The abundance of food and favorable conditions for burrowing in stable sands belong to factors, responsible for permanently high numbers of gerbils in these areas.

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