Elephant Shrews Research Articles

First evidence of a menstruating rodent: the spiny mouse (Acomys cahirinus)

Advances in research relating to menstruation and associated disorders (eg, endometriosis and premenstrual syndrome) have been hindered by the lack of an appropriate animal model. Menstruation, the cyclical shedding of the decidualized endometrium in the absence of pregnancy, is believed to be limited to 78 higher-order primates (human beings and Old World monkeys), 4 species of bat, and the elephant shrew. This represents only 1.5% of the known 5502 mammalian species and <0.09% of these are nonprimates. Thus, many aspects of menstruation remain poorly understood, limiting the development of effective treatments for women with menstrual disorders. Menstruation occurs as a consequence of progesterone priming of the endometrial stroma and a spontaneous decidual reaction. At the end of each infertile cycle as progesterone levels decline the uterus is unable to maintain this terminally differentiated stroma and the superficial endometrium is shed. True menstruation has never been reported in rodents. Here we describe the first observation of menstruation in a rodent, the spiny mouse (Acomys cahirinus). Virgin female spiny mice (n= 14) aged 12-16 weeks were sampled through daily vaginal lavage for 2 complete reproductive cycles. Stage-specific collection of reproductive tissue and plasma was used for histology, prolactin immunohistochemistry, and enzyme-linked immunosorbent assay of progesterone (n= 4-5/stage of the menstrual cycle). Normally distributed data are reported as the mean ± SE and significant differences calculated using a 1-way analysis of variance. Nonnormal data are displayed as the median values of replicates (with interquartile range) and significant differences calculated using Kruskal-Wallis test. Mean menstrual cycle length was 8.7 ± 0.4 days with red blood cells observed in the lavages over 3.0 ± 0.2 days. Cyclic endometrial shedding and blood in the vaginal canal concluding with each infertile cycle was confirmed in all virgin females. The endometrium was thickest during the luteal phase at 322.6 μm (254.8, 512.2), when plasma progesterone peaked at 102.1 ng/mL (70.1, 198.6) and the optical density for prolactin immunoreactivity was strongest (0.071± 0.01 arbitrary units). The spiny mouse undergoes spontaneous decidualization, demonstrating for the first time menstruation in a rodent. The spiny mouse provides a readily accessible nonprimate model to study the mechanisms of menstrual shedding and repair, and may therefore be useful in furthering studies of human menstrual and pregnancy-associated disorders.

The effect of land-use on small mammal diversity inside and outside the Great Fish River Nature Reserve, Eastern Cape, South Africa

This study investigated small mammal species diversity at 10 paired contrast sites along a fence line inside and outside the Great Fish River Nature Reserve (GFRNR), Eastern Cape, South Africa. The sites outside the GFRNR are used for subsistence land-based activities including livestock production and fuelwood harvesting. From 145 live captures, a total of 114 unique individuals of five small mammal species (four rodents and one elephant shrew) were recorded over 1170 trap nights. Average small mammal species diversity and abundance were significantly higher inside the reserve than outside. Human activities such as livestock grazing seemed to explain low levels of small mammal diversity and abundance at the communal sites. Vegetation variables showed a complex interplay with small mammal diversity. In general, high vegetation diversity had a positive influence on small mammal diversity though the influence of some environmental variables was species-dependent. We conclude that the GFRNR is effective in protecting small mammals but the findings raise questions around the influence of land use practices such as livestock grazing on biodiversity, especially given that local communities in South Africa are continuously seeking greater access to reserves for livestock grazing and other provisioning services.

Nonplantigrade Foot Posture: A Constraint on Dinosaur Body Size.

Dinosaurs had functionally digitigrade or sub-unguligrade foot postures. With their immediate ancestors, dinosaurs were the only terrestrial nonplantigrades during the Mesozoic. Extant terrestrial mammals have different optimal body sizes according to their foot posture (plantigrade, digitigrade, and unguligrade), yet the relationship of nonplantigrade foot posture with dinosaur body size has never been investigated, even though the body size of dinosaurs has been studied intensively. According to a large dataset presented in this study, the body sizes of all nonplantigrades (including nonvolant dinosaurs, nonvolant terrestrial birds, extant mammals, and extinct Nearctic mammals) are above 500 g, except for macroscelid mammals (i.e., elephant shrew), a few alvarezsauroid dinosaurs, and nondinosaur ornithodirans (i.e., the immediate ancestors of dinosaurs). When nonplantigrade tetrapods evolved from plantigrade ancestors, lineages with nonplantigrade foot posture exhibited a steady increase in body size following Cope’s rule. In contrast, contemporaneous plantigrade lineages exhibited no trend in body size evolution and were largely constrained to small body sizes. This evolutionary pattern of body size specific to foot posture occurred repeatedly during both the Mesozoic and the Cenozoic eras. Although disturbed by the end-Cretaceous extinction, species of mid to large body size have predominantly been nonplantigrade animals from the Jurassic until the present; conversely, species with small body size have been exclusively composed of plantigrades in the nonvolant terrestrial tetrapod fauna.

Fossils and living taxa agree on patterns of body mass evolution: a case study with Afrotheria.

Most of life is extinct, so incorporating some fossil evidence into analyses of macroevolution is typically seen as necessary to understand the diversification of life and patterns of morphological evolution. Here we test the effects of inclusion of fossils in a study of the body size evolution of afrotherian mammals, a clade that includes the elephants, sea cows and elephant shrews. We find that the inclusion of fossil tips has little impact on analyses of body mass evolution; from a small ancestral size (approx. 100 g), there is a shift in rate and an increase in mass leading to the larger-bodied Paenungulata and Tubulidentata, regardless of whether fossils are included or excluded from analyses. For Afrotheria, the inclusion of fossils and morphological character data affect phylogenetic topology, but these differences have little impact upon patterns of body mass evolution and these body mass evolutionary patterns are consistent with the fossil record. The largest differences between our analyses result from the evolutionary model, not the addition of fossils. For some clades, extant-only analyses may be reliable to reconstruct body mass evolution, but the addition of fossils and careful model selection is likely to increase confidence and accuracy of reconstructed macroevolutionary patterns.

Solar Radiation during Rewarming from Torpor in Elephant Shrews: Supplementation or Substitution of Endogenous Heat Production?

Many small mammals bask in the sun during rewarming from heterothermy, but the implications of this behaviour for their energy balance remain little understood. Specifically, it remains unclear whether solar radiation supplements endogenous metabolic thermogenesis (i.e., rewarming occurs through the additive effects of internally-produced and external heat), or whether solar radiation reduces the energy required to rewarm by substituting (i.e, replacing) metabolic heat production. To address this question, we examined patterns of torpor and rewarming rates in eastern rock elephant shrews (Elephantulus myurus) housed in outdoor cages with access to either natural levels of solar radiation or levels that were experimentally reduced by means of shade cloth. We also tested whether acclimation to solar radiation availability was manifested via phenotypic flexibility in basal metabolic rate (BMR), non-shivering thermogenesis (NST) capacity and/or summit metabolism (Msum). Rewarming rates varied significantly among treatments, with elephant shrews experiencing natural solar radiation levels rewarming faster than conspecifics experiencing solar radiation levels equivalent to approximately 20% or 40% of natural levels. BMR differed significantly between individuals experiencing natural levels of solar radiation and conspecifics experiencing approximately 20% of natural levels, but no between-treatment difference was evident for NST capacity or Msum. The positive relationship between solar radiation availability and rewarming rate, together with the absence of acclimation in maximum non-shivering and total heat production capacities, suggests that under the conditions of this study solar radiation supplemented rather than substituted metabolic thermogenesis as a source of heat during rewarming from heterothermy.

Afromontane small mammals do not follow the hump-shaped rule: altitudinal variation in the Soutpansberg Mountains, South Africa

Abstract:Altitudinal transects of biodiversity are important to understanding macro-ecological patterns. Hump-shaped altitudinal profiles in species richness are a common pattern in terrestrial small-mammal communities studied previously mostly in New World mountain ranges. Based on capture–mark–recapture live-trapping conducted over four seasons (four nights per session) along an altitudinal transect from 1000 to 1747 m asl on the southern slopes of the Soutpansberg Mountains of northern South Africa, we recorded 12 species of small mammal, including nine rodents, two shrews and one elephant shrew (sengi). Three species of rodent and the sengi dominated the communities whilst six species were recorded with three or fewer captures. Although we found no effect of season, we found a significant linear increase in terrestrial small-mammal richness (but not Simpson's diversity, D) with altitude in the Soutpansberg Mountains located just north of the Tropic of Capricorn in South Africa. This trend could best be explained by an increase in the proportion of rocky cover and a change in vegetation from savanna to grassland. Peak densities of three common rodent species and one common insectivore varied from 10 ha−1 (Elephantulus myurus) to 54 ha−1 (Micaelamys namaquensis) and density showed no significant variation with season. However, density either declined (Aethomys ineptus) or increased (Rhabdomys dilectus and Elephantulus myurus) significantly with altitude; this variation was best explained by the proportion of rocky cover. Seasonal variation had a significant effect on body mass of two rodent species. A review of small-mammal studies from major mountain ranges in Africa and Madagascar found that the hump-shaped pattern is not common. Declines in richness with increasing altitude as well as cases of no significant altitudinal change were the most common patterns noted. Tropical African mountains (including the Soutpansberg with 27 species) have far higher small-mammal diversity than temperate African mountains.

Distribution, occupancy, and habitat associations of the gray-faced sengi (Rhynchocyon udzungwensis) as revealed by camera traps

Three of the 4 species of giant sengis or elephant shrews (genus Rhynchocyon) have restricted geographic distributions in eastern Africa and are threatened by anthropogenic habitat loss. However, little is known about their ecology and habitat relationships. We used remotely triggered cameras to detect the gray-faced sengi (Rhynchocyon udzungwensis), which is endemic in the Udzungwa Mountains of Tanzania, with the aim of defining distributional limits, estimating occupancy patterns, and determining habitat requirements. We deployed 183 camera stations over 6 years and accumulated 4,600 camera trapping days. We refined the area of known occurrence to be 390 km 2 , thus confirming the species’ restricted range and vulnerability. We estimated the average occupancy at 56% of sites occupied on sites sampled, and found that occupancy was best predicted by the forest habitat type, with interior, closed-canopy forest supporting highest estimated sengi occupancy. Terrain slope and distance to the nearest park boundary were less important covariates, but nevertheless included among the best models. Camera-trapping rate (photographic events by day) was significantly correlated with subcanopy tree coverage. Combined, these habitat features may provide optimal conditions for antipredation vigilance (vegetation cover), and for nest-building and/or foraging on invertebrates in the thicker leaf litter on gentle slopes. Our results offer new insights into the ecology of giant sengis and confirm the potential utility of camera trapping for occupancy analysis of small, forest-dwelling mammals.

Petrosal and Bony Labyrinth Morphology Supports Paraphyly of Elephantulus Within Macroscelididae (Mammalia, Afrotheria)

Interest in the phylogeny of Macroscelididae (sengis or elephant shrews) has been prompted by molecular studies indicating that Elephantulus rozeti is best placed as the sister group of Petrodromus tetradactylus (this clade being in turn the sister taxon to Macroscelides proboscideus) than among other species of the genus Elephantulus. Until now, no discrete morphological characters have been proposed to support the grouping of E. rozeti, Petrodromus, and Macroscelides into this single so-called ‘Panelephantulus’ clade. Here, we employed μCT scanning in order to investigate the petrosal and bony labyrinth (bony capsule of the inner ear) morphology of most species of extant Macroscelididae. We performed a cladistic analysis on ear traits and found that despite some convergences (e.g., concerning the bony arterial canals in Macroscelides and Rhynchocyon) the middle and inner ear morphology furnishes significant support for the ‘Panelephantulus’ clade. In our analysis, this clade is unambigously supported by the presence of a fully ossified stapediofacial tube. Two additional characters (the presence of a bony septum at the mouth of the fenestra cochleae dividing the D3 sinus into two distinct cavities and the absence of an accessory lateral pneumatic fossa) could also support ‘Panelephantulus.’ These newly discovered morphological characters support the molecular phylogenies published and highlight the importance of coding hitherto difficult to sample morphologies within cladistic analyses using micro-CT techniques. Taxonomic implications are briefly discussed.

RETROSPECTIVE ANALYSIS OF MORTALITIES IN ELEPHANT SHREWS (MACROSCELIDIDAE) AND TREE SHREWS (TUPAIIDAE) AT THE SMITHSONIAN NATIONAL ZOOLOGICAL PARK, USA

Investigations into the cause of mortality and other important findings at necropsy were made into two families of small mammals at the Smithsonian National Zoological Park (SNZP; USA). Necropsy reports from 1976 through 2008 were reviewed for all elephant shrews in family Macroscelididae (n = 118) and all tree shrews in family Tupaiidae (n = 90) that lived for greater than 30 days at the SNZP. Causes of mortality were classified by body system and etiology to identify prevalent diseases and trends across demographics for each family. In elephant shrews, gastrointestinal disease (n = 18) and respiratory disease (n = 22) were important causes of mortality with an increased prevalence of pneumonia in adult males. Trauma was a common cause of mortality in tree shrews (n = 22). Cryptococcosis was an important cause of mortality in both families (n = 8 elephant shrews; n = 13 tree shrews). Bacterial infections, often systemic at time of mortality, were also common (n = 16 elephant shrews; n = 17 tree shrews). Arteriosclerosis was a common comorbid pathology noted at necropsy in certain populations, seen only in Elephantulus rufescens in the family Macroscelididae (n = 22) and in only males in the family Tupaiidae (n = 11). Gongylonemiasis was seen commonly in tree shrews (n = 15), as a comorbid finding, or in 5 cases directly leading to mortality. Awareness of the prevalence of these diseases can help guide prevention and intervention strategies.

The hippocampus of the eastern rock sengi: cytoarchitecture, markers of neuronal function, principal cell numbers, and adult neurogenesis.

The brains of sengis (elephant shrews, order Macroscelidae) have long been known to contain a hippocampus that in terms of allometric progression indices is larger than that of most primates and equal in size to that of humans. In this report, we provide descriptions of hippocampal cytoarchitecture in the eastern rock sengi (Elephantulus myurus), of the distributions of hippocampal calretinin, calbindin, parvalbumin, and somatostatin, of principal neuron numbers, and of cell numbers related to proliferation and neuronal differentiation in adult hippocampal neurogenesis. Sengi hippocampal cytoarchitecture is an amalgamation of characters that are found in CA1 of, e.g., guinea pig and rabbits and in CA3 and dentate gyrus of primates. Correspondence analysis of total cell numbers and quantitative relations between principal cell populations relate this sengi to macaque monkeys and domestic pigs, and distinguish the sengi from distinct patterns of relations found in humans, dogs, and murine rodents. Calretinin and calbindin are present in some cell populations that also express these proteins in other species, e.g., interneurons at the stratum oriens/alveus border or temporal hilar mossy cells, but neurons expressing these markers are often scarce or absent in other layers. The distributions of parvalbumin and somatostatin resemble those in other species. Normalized numbers of PCNA+ proliferating cells and doublecortin-positive (DCX+) differentiating cells of neuronal lineage fall within the overall ranges of murid rodents, but differed from three murid species captured in the same habitat in that fewer DCX+ cells relative to PCNA+ were observed. The large and well-differentiated sengi hippocampus is not accompanied by correspondingly sized cortical and subcortical limbic areas that are the main hippocampal sources of afferents and targets of efferents. This points to intrinsic hippocampal information processing as the selective advantage of the large sengi hippocampus.

Life on high: the diversity of small mammals at high altitude in South Africa

The Great Escarpment is the major mountain system in South Africa, yet very few biological surveys have been conducted outside of the well-known Drakensberg section. This is surprising given the important role that mountains play in local and global biodiversity patterns. In this study, small mammal diversity and community composition were estimated at three high altitude (>1,700 m) sites within the Sneeuberg Mountain Complex (SMC) of the Great Escarpment, South Africa from June 2009 to May 2010. The influences of selected environmental variables on diversity were also tested. Of 423 live-captures, a total of 292 unique individuals of 12 small mammal species (one shrew, one elephant shrew and 10 rodents) were identified during 5,280 trap nights. No single environmental variable could account for the variation observed in diversity measurements but vegetation height appeared to be the most important factor to influence the number of individuals captured. It is hypothesised that the high species richness and diversity of small mammals observed in the SMC compared to other parts of the Great Escarpment is due to the SMC being located in a transition zone of the Grassland and Nama-Karoo biomes. Our results suggest that the SMC could be important in conserving small mammal species from western and eastern assemblages across South Africa.

Variation in body temperature is related to ambient temperature but not experimental manipulation of insulation in two small endotherms with different thermoregulatory patterns

AbstractMaintaining a high and constant body temperature (T b) is thought to enhance performance in endotherms, but such a thermoregulatory pattern is energetically expensive. Thus, some variation in T b is probably universal among endotherms, and several recent attempts have been made to generalize the factors that should cause this variation. Two factors that may be closely tied to the thermoregulatory pattern expressed are the cost of thermoregulation and food availability. To test these predictions, we measured T b of eastern rock elephant shrews (Elephantulus myurus; a traditionally defined heterotherm) and Namaqua rock mice (Micaelamys namaquensis; a homeotherm) in the field and under semi‐natural conditions after experimentally reducing insulation by shaving a patch of dorsal fur. After accounting for ambient temperature (T a), there was no significant difference in the level of variation in T b between shaven and unshaven elephant shrews. In rock mice, the increase in heat loss paradoxically led to decreased variation in T b in the field, but no effect was evident in captivity. Furthermore, as predicted, both species displayed significantly less variation in T b under semi‐natural conditions when given food ad libitum and predation risk involved with obtaining that food was low. Our results show that small mammals, both homeothermic and heterothermic, are capable of altering their thermoregulatory patterns in response to ecological conditions (e.g. rapid changes in food availability). However, increasing the cost of thermoregulation, at least as it was done in this experiment, does not appear to affect the expression of T b as strongly as does T a.

The shoulder musculature of Rhynchocyon cirnei

Traditionally sengis (elephant shrews, order Macroscelidea) have been studied in light of other insect‐eating mammals without a firm understanding of their phylogeny, and all prior work on their myology has been made to animals of similar size rather than similar ancestry. Sengis are now recognized as part of the Afrotheria. Afrotheres have a long evolutionary history and currently few morphological characters are known to unite them, in part due to a limited knowledge of their soft tissue anatomy within a phylogenetically‐meaningful context. One poorly studied afrothere is the giant sengi, Rhynchocyon. We conducted a dissection of the shoulder and arm musculature of R. cirnei. A single specimen preserved in ethanol was dissected using standard dissection techniques under a Zeiss microscope. Muscles were cleaned to determine origin(s), insertion(s) and extent. Each muscle was only reflected after it was documented and photographed. We also identified innervations of each muscle whenever possible. We confirmed prior observations on sengi myology and provide new data on innervations. Comparisons highlight the distinctiveness of sengis relative to other afrotheres where known but also highlight the need for considerable additional anatomical work to firmly establish which features are apomorphies and which may represent shared derived characters among afrotheres.This work is supported by a Midwestern University intramural grant

Differential feeding success of two paralysis-inducing ticks, Rhipicephalus warburtoni and Ixodes rubicundus on sympatric small mammal species, Elephantulus myurus and Micaelamys namaquensis

Rodents are recognised as important hosts of ixodid ticks and as reservoirs of tick-borne pathogens across the world. Sympatric insectivores are usually inconspicuous and often overlooked as hosts of ticks and reservoirs of disease. Elephant shrews or sengis of the order Macroscelidea are small insectivores that often occur in sympatry with rodents in southern Africa. Sengis are invariably parasitised by large numbers of immature ticks while sympatric rodents are infested with very few. The reason for the difference in tick parasitism rates between these hosts is unknown. While a number of mechanisms are possible, we hypothesised that certain tick species exhibit “true host specificity” and as such would only attach and feed successfully on their preferred host or a very closely related host species. To investigate this, we conducted feeding experiments using two economically important tick species, the brown paralysis tick, Rhipicephalus warburtoni and the Karoo paralysis tick, Ixodes rubicundus and two sympatric small mammal species as potential hosts, the eastern rock sengi, Elephantulus myurus and the Namaqua rock mouse, Micaelamys namaquensis. Ticks attached and fed readily on E. myurus, but did not attach or feed successfully on M. namaquensis suggesting that these ticks exhibit true host specificity. We suggest that a kairomonal cue originating from the odour of E. myurus may stimulate the attachment and feeding of these ticks and that they further possess immunosuppressive mechanisms specific to E. myurus, allowing them to feed on this host species but not on M. namaquensis. This study highlights the importance of small mammalian insectivores as potential hosts of ixodid tick species and hence their potential as reservoirs of tick-borne pathogens.

Seasonal changes in thermogenesis of a free-ranging afrotherian small mammal, the Western rock elephant shrew (Elephantulus rupestris)

We report on the seasonal metabolic adjustments of a small-sized member of the phylogenetically ancient Afrotheria, the Western rock elephant shrew (Elephantulus rupestris). We recorded body temperature (T (b)) patterns and compared the capacity for adrenergically induced nonshivering thermogenesis (NST) in E. rupestris captured in the wild in summer and winter. Noradrenaline (NA) treatment (0.4-0.5mg/kg, s.c.) induced a pronounced elevation in oxygen consumption compared to controls (saline), and the increase in oxygen consumption following injection of NA was 1.8-fold higher in winter compared to summer. This suggests that the smaller members of Afrotheria possess functional brown adipose tissue, which changes in thermogenic capacity depending on the season. Torpor was recorded in both seasons, but in winter the incidence of torpor was higher (n=205 out of 448 observations) and minimal T (b) during torpor was lower (T (b)min: 11.9°C) than in summer (n=24 out of 674 observations; T (b)min: 26°C). In addition to cold, high air humidity emerged as a likely predictor for torpor entry. Overall, E. rupestris showed a high degree of thermoregulatory plasticity, which was mainly reflected in a variable timing of torpor entry and arousal. We conclude that E. rupestris exhibits seasonal metabolic adjustments comparable to what has been long known for many Holarctic rodents.

Abiotic and biotic determinants of tick burdens in the eastern rock sengi (Elephantulus myurus)

Ticks are important vectors of pathogens of medical and veterinary importance worldwide. In spite of their economic importance, our current knowledge about the factors affecting tick prevalence and abundance in tropical and subtropical regions is rather limited. Both abiotic (e.g. temperature) as well as biotic variables (e.g. host sex) have been identified as key determinants of distributions. Eastern rock sengis or elephant shrews (Elephantulus myurus, Macroscelidea: Cacroscelididae, Thomas & Schwann) are widely distributed throughout Africa and can harbour a large number of tick species and substantial tick burdens. In the present study, we evaluated the contribution of climate and host factors on tick burdens of sengis. Throughout the year sengis carried high abundances of immature stages of a single tick species, Rhipicephalus sp. near warburtoni. There was no evidence that host parameters affected tick burdens. However, larval abundance decreased with increasing ambient temperatures and both larvae and nymphs were negatively affected by rainfall 2 months before the sampling month. In addition, nymphal burdens decreased with increasing minimum temperatures. Our results suggest that climate factors are the largest constraint for the immature stages of R. sp. near warburtoni and that eastern rock sengis could play a crucial role in the dynamics of tick-borne diseases as a result of the large tick burdens they can sustain.

3D techniques and fossil identification: An elephant shrew hemi-mandible from the Malapa site

Conventional methods for extracting fossilised bones from calcified clastic sediments, using air drills or chemical preparations, can damage specimens to the point of rendering them unidentifiable. As an alternative, we tested an in silico approach that extended preparation and identification possibilities beyond those realisable using physical methods, ultimately proving to be crucial in identifying a fragile fossil. Image data from a matrix-encased hemi-mandible of a micromammal that was collected from the Plio-Pleistocene site of Malapa, Cradle of Humankind, South Africa, were acquired using microtomography. From the resultant images, a 3D rendering of the fossil was digitally segmented. Diagnostic morphologies were evaluated on the rendering for comparison with extant comparative specimens, positively identifying the specimen as an elephant shrew (Elephantulus sp.). This specimen is the first positively identified micromammal in the Malapa faunal assemblage. Cutting-edge in silico preparation technology provides a novel tool for identifying fossils without endangering bone integrity, as is commonly risked with physical preparation.

Body temperature patterns in two syntopic elephant shrew species during winter

We measured body temperature (T b) in free-ranging individuals of two species of elephant shrews, namely western rock elephant shrews ( Elephantulus rupestris) and Cape rock elephant shrews ( E. edwardii), during winter in a winter-rainfall region of western South Africa. These syntopic species have similar ecologies and morphologies and thus potential for large overlaps in diet and habitat use. Unexpectedly, they displayed different T b patterns. Western rock elephant shrews were heterothermic, with all individuals decreasing T b below 30 °C on at least 34% of nights. The level of heterothermy expressed was similar to other species traditionally defined as daily heterotherms and was inversely related to T a, as is commonly seen in small heterothermic endotherms. In contrast, Cape rock elephant shrews rarely allowed their T b to decrease below 30 °C. The level of heterothermy was similar to species traditionally defined as homeotherms and there was no relationship between the level of heterothermy expressed and T a. In both species, the minimum daily T b was recorded almost exclusively at night, often shortly before sunrise, although in some individuals minimum T b occasionally occurred during the day. The interspecific variation in T b patterns among Elephantulus species recorded to date reiterates the importance of ecological determinants of heterothermy that interact with factors such as body mass and phylogeny.

Small Mammal Inventories in the East and West Usambara Mountains, Tanzania. 2. Families Soricidae (Shrews) and Macroscelididae (Elephant Shrews)

Abstract Field surveys conducted in the montane forests of the East and West Usambara Mountains between 1991 and 1993 provide data on the species of shrews (Family Soricidae, Subfamily Crocidurinae) and elephant shrews (Family Macroscelididae). In total, 11 species of crocidurine shrew and one elephant shrew were documented on the Usambara Mountains, which include 10 shrew and one elephant shrew species for the East Usambara, and six species of shrew and one elephant shrew in the West Usambara. Information is presented on the captured animals regarding their natural history, distribution, ecology, and reproduction, as well as external and cranio-dental measurements. Species richness and different levels of endemism between the insectivorous small mammal faunas of tropical Africa and Madagascar are compared.

Small Mammal Inventories in the East and West Usambara Mountains, Tanzania. 1. Study Areas, Methodologies, and General Results

The Eastern Arc Mountains of southeastern Kenya and Tanzania hold a rich and unique biodiversity. The most celebrated of these mountains are the East and West Usambaras. Although these massifs have been the subjects of biological study for over a century, little is known about the shrews, bats, and rodents occurring in the montane forests. Using pitfall buckets, small mammal traps, and mist-nets, between 1991 and 1993, we surveyed the small mammals in the Amani area in the East Usambara and Ambangulu area in the West Usambara to document the natural history of the local fauna. These surveys were conducted during the dry season annually between early July and early September.In this chapter, we outline the study sites, methodologies, capture rates, and other trapping details. A total of 18,563 sample-nights was accrued over the three-year study, and on these two massifs, we documented 12 crocidurine shrew, one macroscelidid elephant shrew, 19 bat, and 14 rodent species. Species accumulation curves suggest that additional trapping effort may be required in the East Usambara Mountains to have a more complete view of the local small mammal fauna. There was no significant difference in the number of individuals captured or measures of species diversity in the pitfall and trap lines that were installed in the same location during the course of this study, suggesting that repeated sampling did not negatively affect the small mammal fauna of the study areas.