Karyotypie diversity and taxonomic problems in the genus Arvicanthis (Rodentia,Muridae)

The analyses of R- and C-banding patterns of chromosomes of Arvicanthis niloticus originating from two different localities (Egypt and Central African Republic) revealed karyotypic differences caused by one pericentric inversion and three translocations, one being reciprocal and the others Robertsonian. There were also some differences in centromeric heterochromatin patterns.The data indicate that these two forms are distinct species, cytogenetically isolated, and that a revision of the taxonomic status of the genus Arvicanthis is needed.     

Chromosomal diversity in the genus Arvicanthis (Rodentia, Muridae) from East Africa: a taxonomic and phylogenetic evaluation

In this paper we discuss the contribution of cytogenetics to the systematics of Arvicanthis in East Africa, by reviewing all the known chromosomal cytotypes of the genus in the area. We also provide G‐ and C‐banding comparisons for two recently described karyotypes, provisionally named ANI‐5 (2n = 56, NFa = 62) and ANI‐6 (2n = 60, NFa = 72). This, therefore, brings the total number of known cytotypes in this area to 10. Five of these correspond to the species recognized by the latest rodent checklist, i.e. A. nairobae (2n = 62, NFa = 78), A. neumanni (2n = 52–53, NFa = 62), A. blicki (2n = 48, NFa = 62), A. abyssinicus (2n = 62, NFa = 64) and A. niloticus (2n = 62, NFa = 60–62). The taxonomic status of the remaining five cytotypes (A. cf. somalicus, 2n = 62 NFa = 62–63; ANI‐5, 2n = 56, NFa = 62; ANI‐6/6a 2n = 60, NFa = 72/76; ANI‐7, 2n = 56, NFa = 78; and ANI‐8, 2n = 44, NF = 72) is discussed. Finally, we reconstruct the phylogenetic relationships among all the known karyotypes on the basis of banding data available for the genus in Africa and show the occurrence of two main clades, each characterized by different types of chromosomal rearrangements. The times of the cladogenetic events, inferred by a molecular clock, indicate that karyotype evolution has accomplished almost all the dichotomic events from the end of the Miocene to the present day. The discovery of a large chromosomal differentiation between populations showing low genetic distances and intrapopulation chromosomal polymorphism suggests that the process of chromosomal differentiation in Arvicanthis is still ongoing and may possibly be responsible for speciation.     

Chromosomal evidence for a polytypic structure of Arvicanthis niloticus (Rodentia, Muridae)

The analysis of R- and C-banding patterns of chromosomes of Arvicanthis niloticus from five localities of Africa (Egypt, Senegal, Burkina-Faso, Mali and Central African Republic) revealed the existence of three karyotypic forms labelled in the study as ANI-1, ANI-2 and ANI-3. These forms differ from each other by 6 to 8 chromosomal rearrangements such as reciprocal and Robertsonian translocations and pericentric inversions. Moreover, they possess different quantities of C-heterochromatin. The data indicate that these three forms are distinct species, cytogenetically isolated, and that a further taxonomic analysis of the genus Arvicanthis is needed.     

Cytogenetics of the genus Arvicanthis (Rodentia,Muridae). 3. Comparative cytogenetics of A. neumanni and A. nairobae from Tanzania

The African rats of the genus Arvicanthis have been widely studied during recent years to clarify species boundaries and phylogenetic relationships. The wide chromosomal variability of the genus has been highlighted in several studies, with each accepted species characterised by its individual karyotypes and others being revealed as cryptic species. In the present paper we report the karyotype and the C- and G-banding patterns of the two species A. nairobae and A. neumanni from seven localities of Tanzania, an area of the range poorly studied. The two karyotypes were compared to that of A. niloticus, which is considered to be primitive. The karyotype of A. neumanni is characterised by 2n = 53–54 and NFa = 62. This karyotypic variability depends on a widespread Robertsonian polymorphism. The karyotype of A. nairobae shows 2n = 62 and NFa = 78; it diverges from that of A. niloticus through one reciprocal translocation, five inversions and three heterochromatic additions. The comparison with the karyotypes of other species of the genus showed that A. neumanni belongs to the east African lineage (with A. abyssinicus, A. blicki, A. niloticus), while A. nairobae is closer to the central and the west African representatives which were all previously under the name of A. niloticus (ANI-2, ANI-3, ANI-4). The distribution of A. nairobae in east Africa opens new scenarios in the biogeographical pattern of evolution of the genus.     

Mitochondrial and Nuclear Genes-Based Phylogeography of Arvicanthis niloticus (Murinae) and Sub-Saharan Open Habitats Pleistocene History

A phylogeographic study was conducted on the Nile grass rat, Arvicanthis niloticus, a rodent species that is tightly associated with open grasslands from the Sudano-Sahelian regions. Using one mitochondrial (cytochrome b) and one nuclear (intron 7 of Beta Fibrinogen) gene, robust patterns were retrieved that clearly show that (i) the species originated in East Africa concomitantly with expanding grasslands some 2 Ma, and (ii) four parapatric and genetically well-defined lineages differentiated essentially from East to West following Pleistocene bioclimatic cycles. This strongly points towards allopatric genetic divergence within savannah refuges during humid episodes, then dispersal during arid ones; secondary contact zones would have then stabilized around geographic barriers, namely, Niger River and Lake Chad basins. Our results pertinently add to those obtained for several other African rodent as well as non-rodent species that inhabit forests, humid zones, savannahs and deserts, all studies that now allow one to depict a more comprehensive picture of the Pleistocene history of the continent south of the Sahara. In particular, although their precise location remains to be determined, at least three Pleistocene refuges are identified within the West and Central African savannah biome.     

Circadian rhythm of photosensitivity and the adaptation of reproductive function to the environment in two populations of Arvicanthis niloticus from Mali and Burkina Faso.

Previous studies have shown that there is a circadian rhythm of photosensitivity in different rodent species of the Sahel (Burkina Faso) and that, despite the low amplitude of seasonal variations in daylength, the photoperiod may control reproductive function. The present investigation of Arvicanthis niloticus provides additional support for this hypothesis. Populations of Arvicanthis niloticus from two regions at the same latitude 1000 km apart but with different climates were studied. Oursi, Burkina Faso, has an arid climate (annual rainfall 315 mm) and Kamalé, Mali has a wetter climate (annual rainfall 1114 mm). The circadian rhythm of photosensitivity had the same features in both populations, involving inhibition of testicular activity, but the photosensitive phase began 11 h 30 min after dawn in the population from Burkina Faso and 45 min later in that from Mali. Comparison of these results with the annual variation of daylength showed that the photoperiod inhibits the reproductive activity of A. niloticus from April to December in Burkina Faso and only from mid-May to mid-August in Mali. The population of Arvicanthis niloticus living in an environment with a large and seasonally stable food supply (Mali) thus has a longer reproductive period. This corroborates results from field studies on annual variations of population density.     

Effects of photoperiod on the reproductive condition of Nile grass rats (Arvicanthis niloticus) from an equatorial population

We evaluated the effects of photoperiod on the reproductive condition of male and female Nile grass rats (Arvicanthis niloticus) descended from members of an equatorial population trapped 2°S of the equator. Study animals housed in 12:12 light:dark (LD) cycles were transferred either to short photoperiod (9:15) or long photoperiod (15:9) for 9 weeks (males) or 11 weeks (females), and various reproductive parameters were assessed. We observed no differences between short‐ and long‐day males with respect to plasma concentration of testosterone, testicular mass, seminal vesicle mass, or spermatogenesis. Similarly, we observed no differences between short‐ and long‐day females with respect to oestrous cycles, uterine mass, follicle size, or presence of corpora lutea. Reproductive parameters of male and female A. niloticus housed in short‐ and long photoperiods were similar to those typically observed among animals descended from the same equatorial population and housed in LD 12:12. Thus, photoperiod appears not to elicit changes in reproductive condition among A. niloticus from populations whose native habitat lies within 2° of the equator. These data contrast with the results of other studies indicating that photoperiod alters reproductive condition in A. niloticus populations living >10° from the equator.     

Spatial and temporal patterns in Arvicanthis niloticus (Desmarest, 1822) as revealed by radio-tracking

Arvicanthis niloticus was radio‐tracked in the grasslands of the Queen Elizabeth National Park in Uganda. Home range sizes calculated by Ranges V^® using the Minimum Convex Polygon Method (at 95%) were on average 5.5 times larger in the bushland–grassland mosaic than those in the Abutilon guineense–Ocimum suave bushland. An inverse relation between home range size and population density was found. In both habitats the species was highly active during daylight hours but differed in activity patterns.     

Diet and habitat aridity affect osmoregulatory physiology: An intraspecific field study along environmental gradients in the Rufous-collared sparrow

The urine field osmolality in Zonotrichia capensis along a latitudinal gradient in rainfall and temperature in Chile was examined. We also investigated latitudinal variation in the renal traits that mediate how these birds cope with dehydration. We used the δ¹⁵N of this species' tissue to investigate whether the reliance on animals and seeds varied among birds and if it had any effect on excretion and renal traits. We found a significant latitudinal variation in urine osmolality, a variable that was correlated with habitat aridity. We also found that the kidney size and proportion of kidney devoted to medullary tissue differed between birds from arid and mesic localities, but not in a lineal fashion with aridity. The increment in the position in the food web, as measured by δ¹⁵N, led to an increment in urine osmolality, without changes in kidney features. Our data suggested that differences in dietary habits in the field could be not extended enough to cause changes in the kidney structure in Rufous-collared sparrows.     

Electrophoretic studies on blood proteins of Arvicanthis niloticus

Electrophoretic patterns of blood proteins were compared among groups of Arvicanthis niloticus from widely distant origin: Senegal, Haute-Volta, Egypt. Four blood proteins/enzymes were studied: albumin, transferrin, esterase and 6-phosphogluconate dehydrogenase. The results demonstrate two levels of difference: (a) inter-population differences; and (b) intra-species genetic polymorphism. The latter was not observed for all populations nor for all loci. The differences between populations suggest that some process of specification could be under way among Arvicanthis.     

A Multivariate Analysis of Soil Yeasts Isolated from a Latitudinal Gradient

Yeast isolates from soil samples collected from a latitudinal gradient (>77°S to >64°N) were subjected to multivariate analysis to produce a statistical foundation for observed relationships between habitat characteristics and the distribution of yeast taxa (at various systematic levels) in soil microbial communities. Combinations of temperature, rainfall (highly correlated with net primary productivity), and electrical conductivity (EC) could explain up to ca. 44% of the distribution of the predominant yeast species, rainfall and pH could explain ca. 32% of the distribution of clades in the most common orders (Filobasidiales and Tremellales), whereas vegetation type (trees, forbs, and grass) played the same role for orders. Cryptococcus species with appropriate maximum temperatures for growth predominated in most soils. Cryptococcus species in the Albidus clade of the Filobasidiales predominated in desert soils; Cryptococcus species of other clades in the Filobasidiales and Tremellales predominated in wetter and more-vegetated soils, with Tremellalean species favored in soils of lower pH or higher EC. The predominance of Cryptococcus species in soils has been attributed to their polysaccharide capsules, particularly important when competing with bacteria in arid soils.     

Chromosomal characterization of Arvicanthis species (Rodentia,Murinae) from western and central Africa: implications for taxonomy

A chromosome study of unstriped grass rats of the genus Arvicanthis (Rodentia, Murinae) in western and central Africa is presented. The observations extend the data available to 242 specimens from 59 localities. All individuals karyotyped belong to four karyotypic forms, or cytotypes, earlier described as ANI-1, ANI-2, ANI-3, and ANI-4 and are presumed to correspond to four distinct species. In order to provide diagnostic characters for these western and one central African Arvicanthis species, we standardized the chromosomal data available and developed a G- and C-banded chromosome nomenclature that allows easy species identification. Each form is characterized by a distinct geographical distribution, roughly following the biogeographical domains of western Africa, although their precise limits remain to be assessed. The sole area of sympatry detected is the region of the inner delta of the Niger River, where both ANI-1 and ANI-3 can be found. It is proposed that the three western African species ANI-1, ANI-3, and ANI-4 be renamed as A. niloticus, A. ansorgei, and A. rufinus, respectively.     

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.     

From daily behavior to hormonal and neurotransmitters rhythms: Comparison between diurnal and nocturnal rat species

Mammalian species can be defined as diurnal or nocturnal, depending on the temporal niche during which they are active. Even if general activity occurs during nighttime in nocturnal rodents, there is a patchwork of general activity patterns in diurnal rodents, including frequent bimodality (so-called crepuscular pattern, i.e., dawn and dusk peaks of activity) and a switch to a nocturnal pattern under certain circumstances. This raises the question of whether crepuscular species have a bimodal or diurnal - as opposed to nocturnal - physiology. To this end, we investigated several daily behavioral, hormonal and neurochemical rhythms in the diurnal Sudanian grass rat (Arvicanthis ansorgei) and the nocturnal Long-Evans rat (Rattus norvegicus). Daily rhythms of general activity, wheel-running activity and body temperature, with or without blocked wheel, were diurnal and bimodal for A. ansorgei, and nocturnal and unimodal for Long-Evans rats. Moreover, A. ansorgei and Long-Evans rats exposed to light-dark cycles were respectively more and less active, compared to conditions of constant darkness. In contrast to other diurnal rodents, wheel availability in A. ansorgei did not switch their general activity pattern. Daily, unimodal rhythm of plasma leptin was in phase-opposition between the two rodent species. In the hippocampus, a daily, unimodal rhythm of serotonin in A. ansorgei occurred 7 h earlier than that in Long-Evans rats, whereas a daily, unimodal rhythm of dopamine was unexpectedly concomitant in both species. Multiparameter analysis demonstrates that in spite of bimodal rhythms linked with locomotor activity, A. ansorgei have a diurnally oriented physiology.     

Correlated patterns of variation in phenology and seed production in populations of two annual grasses along an aridity gradient

I applied a comparative approach to reveal correlated patterns of variation in phenology and seed production in four populations of two annual grasses Hordeum spontaneum and Avena sterilis, sampled in the same environments distributed along an aridity gradient in Israel. The steep aridity gradient in Israel represents two parallel clines of environmental productivity (annual rainfall) and predictability (variation in amount and timing of annual rainfall) that is likely to induce similar responses in natural plant populations distributed along the gradient, if (1) selection is strong, (2) species share the same ecological niche, and (3) there is genetic variation for ecologically important traits. I found in plants of both species (1) ultimate advance in onset of flowering, and (2) more but smaller seeds, with increasing aridity. The broad sense heritabilities of onset of flowering, seed size and seed yield in both species were very high, moderate and low, respectively. It appears that the observed adaptive complex of traits have evolved in both species in response to this specific array of environments.     

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.     

Neotropical savanna ants show a reversed latitudinal gradient of species richness,with climatic drivers reflecting the forest origin of the fauna

Aim

To evaluate the extent to which ant species richness in Neotropical savannas varies with macrogeographic variables, and to identify the potential climatic drivers of such variation.

Location

The Cerrado savanna biome of central Brazil, in a region spanning ca. 20° of latitude and 18°of longitude.

Methods

Standardized sampling of the arboreal and ground‐dwelling faunas was performed in 29 well‐preserved savanna sites using pitfall traps. Species were classified according to their habitat affinities: open‐savanna specialists, forest‐associated species or habitat generalists. We used generalized linear models to evaluate the importance of geographic (latitude, longitude and elevation) and climatic (mean temperature and three metrics of rainfall) variables as predictors of species richness.

Results

The total number of species recorded at each site varied more than twofold (from 59 to 144), and latitude was the best geographic correlate of overall species richness. However, contrary to the expected pattern, more species were found at higher than lower latitudes. This reversed latitudinal pattern of diversity occurred for both the arboreal and ground‐dwelling faunas, and for the habitat generalists and forest specialists. The savanna specialists showed a mid‐latitudinal peak in diversity. Overall, there was a significant positive association between rainfall and species richness, but the strength of this relationship varied with ant habitat affinity.

Main conclusions

The Cerrado ant fauna shows a reverse latitudinal gradient in species diversity, and this can be explained by increasing rainfall during the warmest months of the year (and therefore in plant productivity) with increasing latitude. The sensitivity of Cerrado ant diversity to declining rainfall contrasts with the high resilience to aridity of the Australian savanna ant fauna, and this reflects the contrasting evolutionary histories of these faunas. Our findings highlight the importance of historical processes as drivers of intercontinental contrasts in macroecological patterns.     

Diversity and evolution of African Grass Rats (Muridae: Arvicanthis)—From radiation in East Africa to repeated colonization of northwestern and southeastern savannas

African Grass Rats of the genus Arvicanthis Lesson, 1842, are one of the most important groups of rodents in sub‐Saharan Africa. They are abundant in a variety of open habitats, they are major agricultural pests, and they became a popular model in physiological research because of their diurnal activity. Despite this importance, information about their taxonomy and distribution is unsatisfactory, especially in eastern Africa. In this study, we collected the most comprehensive multilocus DNA dataset to date across the geographic and taxonomic range of the genus (229 genotyped specimens from 130 localities in 16 countries belonging to all currently recognized species). We reconstructed phylogenetic relationships, mapped the distribution of major genetic clades, and used the combination of cytogenetic, nuclear, and mitochondrial markers for species delimitations and taxonomic suggestions. The genus is composed of two major evolutionary groups, called here the ANSORGEI and NILOTICUS groups. The former contains four presumed species, while the latter is more diverse and we recognized nine species. Most relationships among species are not resolved, which suggests a rapid radiation (dated to early–middle Pleistocene). Further, there is an indication of reticulate evolution in Ethiopia, that is, the region of the highest Arvicanthis diversity. The distribution of genetic diversity suggests diversification in eastern Africa, followed by repeated dispersals to the west (Sudano‐Guinean savannas) and to the south (Masai steppe). We propose nomenclatural changes for Ethiopian taxa and provide suggestions for future steps toward solving remaining taxonomic questions in the genus.     

Species differences in carbon isotope ratios, specific leaf area and nitrogen concentrations in leaves of Eucalyptus growing in a common garden compared with along an aridity gradient

Leaves produced in 2004 of 422 species of Eucalyptus whose natural habitat is southern Australia were sampled at the Currency Creek Arboretum in South Australia where the annual (mainly winter) rainfall is about 400 mm. Tree height, leaf area, leaf dry weight, leaf nitrogen (N) concentration and leaf carbon isotope ratio (δ¹³C) were measured and the specific leaf area (SLA) calculated. Among the 422 species, the SLA varied from 1.5 to 8.8 m² kg^?1 and N concentration varied from 0.6 to 2.1%, much greater than in 64 species collected along an aridity transect from southwestern Western Australia to central Australia in 2003. Also, the range of leaf δ¹³C values was similar in the common garden to that across the aridity transect. For the 45 species present in both studies, the SLA and leaf N concentration in the common garden were similar to those measured in leaves along the aridity transect, indicating that these characteristics are inherent in the species and vary little with environment. The variation in leaf δ¹³C in the common garden was just as great as along the transect, but the values measured in the one location were poorly correlated with those along the transect. This was not expected, as the variation in δ¹³C at one common site in South Australia was anticipated to be less than along the aridity gradient where annual rainfall varied from 250 to 1200 mm. Path analysis on the 45 species common to both studies indicated that rainfall did not have a direct effect on δ¹³C, but the differences in δ¹³C resulted from indirect effects of rainfall on SLA and N concentration. δ¹³C was negatively correlated with SLA but positively correlated with N. Thus, both effects may compensate for each other so that no significant relationship between δ¹³C and rainfall was observable. However, there is a large degree of variation of δ¹³C at any level of rainfall. The origin and ecological implications of this observation are discussed.     

Temperature-related distributions of Metadiaptomus and Tropodiaptomus (Copepoda: Calanoida), particularly in southern Africa

Updated locality records of species of Metadiaptomus and Tropodiaptomus on the African continent confirm the generally disjunct distribution of these two taxa as recognised by Dumont (1980) in North Africa. Distributional data for southern Africa reveal little range overlap between these two genera. Apart from two south western Cape taxa, species of Metadiaptomus are largely confined to upland, higher latitude, semi-arid or arid warm subtemperate regions, while species of Tropodiaptomus generally occupy moist, lower-lying, lower latitude subtropical regions. Separation along latitudinal and/or altitudinal axes implicates temperature as a controlling factor, while separation on the precipitation axis suggests the importance of habitat permanence. Using a multiple regression equation derived for African waters to predict water temperature from latitude and altitude, it is shown that the two genera tend to separate around the 20 °C mean annual temperature isotherm. Additional factors influencing distribution (habitat permanence, water quality, competition and predation) are discussed.