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Life history consequences of mammal sibling rivalry.

Mammal life history traits relating to growth and reproduction are extremely diverse. Sibling rivalry may contribute to selection pressures influencing this diversity, because individuals that are relatively large at birth typically have an advantage in competition for milk. However, selection for increased growth rate is likely to be constrained by kin selection and physiological costs. Here, we present and test a model examining the ESS (evolutionarily stable strategy) balance between these constraints and advantages associated with increased prenatal growth in mammal sibling rivalry. Predictions of the model are supported by results of comparative analyses for the Carnivora and Insectivora, which demonstrate an increase in prenatal growth rate with increasing intensity of postnatal scramble competition, and a decrease in postnatal growth rate relative to size at birth. Because increased prenatal growth rates are predicted to select for reduced gestation length under certain conditions, our study also indicates that sibling rivalry may contribute to selection pressures influencing variation in altriciality and precociality among mammals.

Animal Nutritional Physiological Phenomena↗

Protein sequence signatures support the African clade of mammals.

DNA sequence evidence supports a superordinal clade of mammals that comprises elephants, sea cows, hyraxes, aardvarks, elephant shrews, golden moles, and tenrecs, which all have their origins in Africa, and therefore are dubbed Afrotheria. Morphologically, this appears an unlikely assemblage, which challenges-by including golden moles and tenrecs-the monophyly of the order Lipotyphla (Insectivora). We here identify in three proteins unique combinations of apomorphous amino acid replacements that support this clade. The statistical support for such "sequence signatures" as unambiguous synapomorphic evidence for the naturalness of the Afrotherian clade is reported. Using likelihood, combinatorial, and Bayesian methods we show that the posterior probability of the mammalian tree containing the Afrotherian clade is effectively 1.0, based on conservative assumptions. Presenting sequence data for another African insectivore, the otter shrew Micropotamogale lamottei, we demonstrate that such signatures are diagnostic for including newly investigated species in the Afrotheria. Sequence signatures provide "protein-morphological" synapomorphies that may aid in visualizing monophyletic groupings.

Africa↗

Phenotypic and genotypic identification of staphylococci isolated from wild small mammals.

229 strains of Staphylococcus isolated from the intestines of wild small mammals (Insectivora and rodents) were analysed by phenotypic tests and 16S-23S rDNA intergenic spacer PCR amplification (ITS-PCR). Based on the results of three methods (phenotypical tests, API numerical profiles and ITS-PCR) 65.5% of all staphylococcal strains could be identified. The remaining strains were compared by cluster analysis with reference and identified strains. It is quite possible that, a part of unidentified strains represent hitherto undescribed species, all the more as their ITS types were not found among other members of the genus Staphylococcus. Moreover, the results show that small mammals provides a suitable new habitat for certain staphylococci.

Animals↗

Additional support for Afrotheria and Paenungulata, the performance of mitochondrial versus nuclear genes, and the impact of data partitions with heterogeneous base composition.

We concatenated sequences for four mitochondrial genes (12S rRNA, tRNA valine, 16S rRNA, cytochrome b) and four nuclear genes [aquaporin, alpha 2B adrenergic receptor (A2AB), interphotoreceptor retinoid-binding protein (IRBP), von Willebrand factor (vWF)] into a multigene data set representing 11 eutherian orders (Artiodactyla, Hyracoidea, Insectivora, Lagomorpha, Macroscelidea, Perissodactyla, Primates, Proboscidea, Rodentia, Sirenia, Tubulidentata). Within this data set, we recognized nine mitochondrial partitions (both stems and loops, for each of 12S rRNA, tRNA valine, and 16S rRNA; and first, second, and third codon positions of cytochrome b) and 12 nuclear partitions (first, second, and third codon positions, respectively, of each of the four nuclear genes). Four of the 21 partitions (third positions of cytochrome b, A2AB, IRBP, and vWF) showed significant heterogeneity in base composition across taxa. Phylogenetic analyses (parsimony, minimum evolution, maximum likelihood) based on sequences for all 21 partitions provide 99-100% bootstrap support for Afrotheria and Paenungulata. With the elimination of the four partitions exhibiting heterogeneity in base composition, there is also high bootstrap support (89-100%) for cow + horse. Statistical tests reject Altungulata, Anagalida, and Ungulata. Data set heterogeneity between mitochondrial and nuclear genes is most evident when all partitions are included in the phylogenetic analyses. Mitochondrial-gene trees associate cow with horse, whereas nuclear-gene trees associate cow with hedgehog and these two with horse. However, after eliminating third positions of A2AB, IRBP, and vWF, nuclear data agree with mitochondrial data in supporting cow + horse. Nuclear genes provide stronger support for both Afrotheria and Paenungulata. Removal of third positions of cytochrome b results in improved performance for the mitochondrial genes in recovering these clades.

Animals↗

Regional and laminar distribution of cortical neurons projecting to either superior or inferior colliculus in the hedgehog tenrec.

Retrograde tracer substances were injected into either the inferior or the superior colliculus in the Madagascan hedgehog tenrec, Echinops telfairi (Insectivora), to reveal the laminar and regional distribution of corticotectal cells and to correlate the labeled areas with architectural data. The tenrecs, taken from our breeding colony, have one of the least differentiated cerebral cortices among mammals, and experimental investigations of such brains are important for the understanding of the evolution and intrinsic organization of the more highly differentiated cerebral cortex in other placental mammals. Following injections into the inferior colliculus, cortical neurons were labeled bilaterally, with an ipsilateral predominance. Most labeled cells were found in the caudolateral hemisphere, area 4 as defined by Rehkämper (1981); some were in the somatosensorimotor cortex, as defined in a previous study. The labeled neurons in area 4 were located in layers V and VI, forming two bands of cells separated from each other by a poorly labeled interspace. A further subdivision of this presumed auditory region could not be achieved. This entire area was also weakly labeled following tracer injections into the superior colliculus. The labeled cells, however, were restricted to layer V of the ipsilateral side. The most consistent sites of labeled cells following injections into the superior colliculus were found in layer V of the ipsilateral caudomedial hemisphere, Rehkämper's caudal area 3, and the transitional zone adjacent to the retrosplenial cortex. This area is small in comparison to the entire region that was found in this study to project to the superior colliculus. The superior colliculus also receives projections from the ipsilateral sensorimotor and cingulate cortices. The latter projections are particularly striking in comparison to other mammals because they originate from along the entire rostrocaudal extent of the cingulate/retrosplenial region.

Afferent Pathways↗

Cream, a new coat-color mutant in the musk shrew.

A coat-color mutant was found in the wild musk shrew (Suncus murinus, Insectivora). Five musk shrews with gray pelage, the common coat color of this species, were captured in the village of Tambum near Jakarta, Indonesia. Two males and two females were transported to Japan and mated. Matings between one male and two females segregated several cream-colored offspring, a color that had never been seen before in this species. From the pedigree record and data on mating experiments, it was confirmed that this mutant coat color was expressed in the homozygote by an autosomal recessive gene designated cr, and at least three of the four wild shrews examined were carriers of this gene. The cr gene was associated with failure of normal pigmentation in the pelage and skin. The mutant shrews also showed some behavioral abnormalities.

Animals↗

Origin and evolution of homologous repeated sequences in the mitochondrial DNA control region of shrews.

The complete mitochondrial DNA (mtDNA) control region was amplified and directly sequenced in two species of shrew, Crocidura russula and Sorex araneus (Insectivora, Mammalia). The general organization is similar to that found in other mammals: a central conserved region surrounded by two more variable domains. However, we have found in shrews the simultaneous presence of arrays of tandem repeats in potential locations where repeats tend to occur separately in other mammalian species. These locations correspond to regions which are associated with a possible interruption of the replication processes, either at the end of the three-stranded D-loop structure or toward the end of the heavy-strand replication. In the left domain the repeated sequences (R1 repeats) are 78 bp long, whereas in the right domain the repeats are 12 bp long in C. russula and 14 bp long in S. araneus (R2 repeats). Variation in the copy number of these repeated sequences results in mtDNA control region length differences. Southern blot analysis indicates that level of heteroplasmy (more than one mtDNA form within an individual) differs between species. A comparative study of the R2 repeats in 12 additional species representing three shrew subfamilies provides useful indications for the understanding of the origin and the evolution of these homologous tandemly repeated sequences. An asymmetry in the distribution of variants within the arrays, as well as the constant occurrence of shorter repeated sequences flanking only one side of the R2 arrays, could be related to asymmetry in the replication of each strand of the mtDNA molecule. The pattern of sequence and length variation within and between species, together with the capability of the arrays to form stable secondary structures, suggests that the dominant mechanism involved in the evolution of these arrays in unidirectional replication slippage.

Animals↗

Distinctive features of the gametes and reproductive tracts of the Asian musk shrew, Suncus murinus.

Insectivora are of special interest as the most primitive of the eutherian mammals, but essentially nothing is known of their gamete function. In this respect, the Asian musk shrew (Suncus murinus), investigated in the present study, displays many idiosyncrasies. In the epididymis, the giant acrosome undergoes further stabilization, its unusual resilience being especially evident in a "rim" created by a persistent close alignment of the outer acrosomal and overlying plasma membranes. However, until spermatozoa reached a gland on the vas deferens, no post-testicular change was demonstrable in the sperm head surface, the unusual nature of which was indicated by a dorso-ventral differentiation, by an inability to auto-agglutinate or to bind to the homologous zona pellucida, and by an insensitivity to anti-sperm immunoglobulin IgG in fresh serum. At mating, only about 1 x 10(6) spermatozoa are inseminated as far as the anterior vagina with plug formation. Within the small (6 mm) fallopian tube, the isthmus and ampulla are sharply delineated by their contractile activity and epithelial character; there is evidence of some sperm entry into isthmic crypts and a tendency for ipsilateral ovarian control of sperm transport to the tubal ampulla. The cumulus oophorus does not undergo preovulatory mucification and expansion, is characterized by persistent intercellular gap junctions, and is insensitive to hyaluronidase and trypsin. It is unclear how the compact cumulus is penetrated at fertilization. The giant acrosome contains acrosin and an unusually temperature-dependent cumulytic activity; it is intact in motile ampullary spermatozoa but appears to be discharged before reaching the zona pellucida. Since eggs were not penetrated in the presence of ampullary spermatozoa until 4-10 h after ovulation, Suncus spermatozoa spend a long period in the female before they can fertilize. The determinants of sperm function, including capacitation and the acrosome reaction (AR), may depend on a different set of controls in Suncus, perhaps as a legacy of the resilient giant acrosome. This possibility could be examined in other Crociduran and Soricine shrews selected according to acrosome size.

Acrosome↗

Development of microarray-based diagnostics of voles and shrews for use in biodiversity monitoring studies, and evaluation of mitochondrial cytochrome oxidase I vs. cytochrome b as genetic markers.

Molecular methods are widely used for species identification of mammals. In particular, the mitochondrial cytochrome b gene sequence has proven helpful for this purpose. Microarray technology can now open up new perspectives for biodiversity monitoring. With microarrays, many thousands of genetically based characteristics can be tested on one microscopic glass slide called a 'chip'. A 'Mammalia-Chip', for example, could include redundant diagnostic markers to unambiguously identify all European mammal species. Of broader use, and therefore economically more relevant, could be a 'Biodiversity-Chip', containing diagnostic features to distinguish key species in the taxa of bacteria, lichen, molluscs, insects, fungi, mammals, etc. An important prerequisite for any mixed-phyla chip is a standardization of methods. One of the most promising genes as a universal marker for all eukaryotes is cytochrome oxidase I. We show that cytochrome oxidase I is adequate for the discrimination of different species of voles and shrews with cluster analysis. Based on these results we present a diagnostic microarray-chip using cytochrome oxidase I sequences for the identification of three species of Sorex (Soricidae, Insectivora) and four species of Microtus (Arvicolinae, Rodentia). We conclude that cytochrome oxidase I can be used as an alternative marker to cytochrome b in a mixed-phyla chip, or both genes can be used in combination to enhance redundance and thus robustness of a specific chip including small mammals.

Animals↗

Some corrections in Haemogregarine (Apicomplexa: Protozoa) nomenclature.

The nomenclature of three genera in the family Haemogregarinidae (Haemogregarina, Karyolysus, and Hepatozoon) has been reviewed and the following new names are introduced to replace homonyms or for previously unnamed species: haemogregarina carlosi n. nom., in the erythrocytes of the lizard Lacerta ocellata; Haemogregarina tincae n. nom., in the stomach and intestine of the tench Tinca tinca; Hepatozoon insectivorae n. sp., in the leucocytes of the shrews Sorex araneus and Crocidura leucodon; Hepatozoon krampitzi n. sp., in the leucocytes of the vole Microtus oeconomus; Hepatozoon peromysci n. sp., in the leucocytes of the deermice Peromyscus boylii and P. truei gilberti; and Hepatozoon pallida (Pessoa et al., 1971) n. comb., in the erythrocytes of the snake Thamnodynastes pallidus nattereri.

Animals↗

Purgatorius, an early paromomyid primate (Mammalia).

Fragmentary mandibles of Purgatorius unio Van Valen and Sloan from the Puercan (approximately early Paleocene), Garbani Locality, Montana, preserve associated postcanines. Their morphology indicates that this mammal was an early paromomyid primate and suggests that primate ancestry does not include currently known members of the palaeoryctid and leptictid Insectivora or of the Condylarthra.

Animals↗

Parvalbumin immunoreactive neurons in the main olfactory bulb of the house musk shrew, Suncus murinus.

The distribution, morphological features, and postnatal development of parvalbumin (PV) immunoreactive neurons in the main olfactory bulb (MOB) of the house musk shrew, Suncus murinus, were studied to report for the first time on PV positive bulbar interneurons in the order Insectivora. In adult animals, PV neurons are distributed in the glomerular layer (GL), the external plexiform layer (EPL), the internal plexiform layer (IPL) and the granule cell layer (GCL) of the MOB. These neurons were identified as a subpopulation of periglomerular cells and perinidal cells [Alonso et al., 1995] in the GL and at the GL-EPL border, respectively, and as bipolar and multipolar neurons in the EPL and four types of the interneurons (horizontal cells, Cajal cells, Golgi cells, and bitufted cells) in the layers deeper than the mitral cell layer. During development of PV neurons, neurons exhibiting extremely faint PV immunoreactivity first appeared in the GCL at postnatal day 14 and increased markedly in number and intensity of their PV immunoreactivity from postnatal days 14 to 28. At postnatal day 21, PV neurons were identified as periglomerular cells in the GL, perinidal cells at the GL-EPL border, and morphologically unidentifiable neurons in the EPL, IPL and GCL. At postnatal day 28, PV neurons exhibited a nearly adult pattern with respect to distribution and structural features. The present results strongly suggest that a wide variety of PV positive neurons in the MOB of the house musk shrew may develop postnatally.

Animals↗

Cortical layer VII and persistent subplate cells in mammalian brains.

Layer VII is the deepest cortical layer in rats, and consists of a thin layer of persistent subplate cells overlain by a cell-sparse, myelin-rich stratum through which many corticocortical axons travel. Layer VII neurons participate in local and long-distance corticocortical connections. The present study was undertaken to determine whether layer VII is a typical feature in rodent brains, and to determine which other mammalian taxa exhibit a layer VII. The adult brains of 144 species from 22 orders were examined. Of these, 43 species in 6 orders exhibit a layer VII. These include the sciurognath Rodentia, Insectivora, Paucituberculata, Paramelemorphia, some Xenarthra, and some Chiroptera. In all taxa interstitial cells were observed scattered throughout the white matter. The observed distribution of layer VII in this sample of mammalian taxa suggests that layer VII is a typical feature in some orders, but is not present in most orders. The heterogeneous distribution of layer VII in the Rodentia and Chiroptera suggests that species-level developmental dynamics are involved. It is hypothesized that the timing of subplate apoptosis in relation to the establishment of corticocortical connections is the major factor that determines whether layer VII is present in the adult stage.

Animals↗

Chromosome homologies between man and mountain zebra (Equus zebra hartmannae) and description of a new ancestral synteny involving sequences homologous to human chromosomes 4 and 8.

Using human chromosome painting probes, we looked for homologies between human and mountain zebra (Equus zebra hartmannae, Equidae, Perissodactyla) karyotypes. Except for two very short segments, all euchromatic regions were found to have a human homologous chromosome segment. Conserved syntenies previously described in various mammalian orders were detected. Each synteny corresponded to a chromosomal region homologous to two parts of human chromosomes: HSA3 and HSA21, HSA7 and HSA16, HSA12 and HSA22, and HSA16 and HSA19. Chromosomal segments homologous to a part of HSA11 and HSA19p are found syntenic in zebra, horse and donkey, suggesting that this group of synteny has been inherited from an Equidae or Perissodactyla common ancestor. A synteny of segments homologous to parts of HSA4 and HSA8 was observed in zebra and horse. It also exists in the rabbit (Lagomorpha) and several Carnivora species. A second group of taxa which does not have this region of synteny is composed of primates, Chiroptera and Insectivora, and possibly also Cetacea and Scandantia. Thus, the presence or absence of this region of synteny may separate two groups of eutherian mammals.

Animals↗

Sex chromosome aneuploidy in wild small mammals.

We describe four examples of the XO condition in wild mammals. One XO house mouse (Mus musculus domesticus) was caught in nature and subsequently gave birth to three litters in captivity, confirming for wild mice the fertility observed for XO laboratory mice. Two other XO house mice were produced from laboratory crosses of wild-caught mice. An immature XO common shrew (Sorex araneus) was caught in nature; this appears to be the first XO recorded in the order Insectivora. We collected data from researchers studying chromosome variation in house mice and common shrews and found an overall incidence of 0.22% sex chromosome aneuploidy in 4608 mice and 0.05% in 6625 shrews. The discrepancy related to a much higher frequency of XO's in mice than shrews. Single XXY and XYY shrews and an XXX mouse have been recorded in nature.

Animals↗

Historical biogeography of the strepsirhine primates of Madagascar.

Lying some 400 km off the coast of southeastern Africa, Madagascar is the world's largest oceanic island. It has been in roughly the same position relative to its parent continent for 120 million years, and as a consequence its mammal fauna is unusual in composition, with a low number of major taxa but a high diversity at lower taxonomic levels. Among Madagascar's native terrestrial mammals, only the orders Primates, Rodentia, Carnivora and Insectivora are represented (plus, until recently, the enigmatic and endemic Bibymalagasia, and Artiodactyla in the form of semiaquatic pygmy hippopotamuses). This reflects the fact that terrestrial mammals are notoriously poor over-water dispersers; yet at the same time the ancestors of all of Madagascar's mammals had to have crossed a wide oceanic barrier to get to the island at various points during the Tertiary. Here I examine the palaeogeographic evidence for potential land bridge or 'stepping-stone' connections with adjacent continents from the Mesozoic through the Cenozoic, and review the fossil records and phylogenies of each of Madagascar's mammalian groups in an attempt to estimate the minimum number of crossings necessary to produce the island's current faunal composition. Probable monophyletic origins for each major group, and thus a smaller rather than a larger number of crossings of the Mozambique Channel, imply that this water barrier has acted as a powerful filter; so powerful that it is unclear whether any crossings would have been possible without some form of subaerial connection, however ephemeral, at least from time to time during the Tertiary. Clarification of how Madagascar's terrestrial mammal fauna may have originated is thus as likely to emerge from the geology of the seafloor surrounding the island as it is to come from the fossil record or from the internal and external relationships of its various components.

Animal Migration↗

Phylogeny through brain traits: more characters for the analysis of mammalian evolution.

We have assembled data on nine brain traits, in addition to the fifteen we have previously described, which provide new evidence for assessing mammalian relationships. States of these characters are tabulated as they occur in each of 152 mammalian species, providing data in numerically ordered form, useful for multiple analyses of phylogenetic relationships in programs which take into account variations in several different characters simultaneously. Derived states of each of the nine traits are characteristic of certain restricted groups of mammals; (1) mirroring of the complete SI body representation in isocortex (anthropoid primates); (2) loss of the accessory olfactory bulbs (sirenians, cetaceans, most bats, catarrhine primates); (3) Rindenkerne, clumps of cell bodies in layer 6 of cerebral cortex (sirenians); (4) posteriorly-pointing digits in the SI body representation (bats, both mega- and micro-); (5) equivalent tectopetal connections to the anterior colliculus of one side from both retinas, rather than predominantly from the contralateral retina (primates and megabats); (6) loss of lamination in dorsal cochlear nuclei (anthropoid primates, bats, seals, sirenians, cetaceans); (7) separation of claustrum from cerebral cortex (diprotodont marsupials, carnivores, artiodactyls, perissodactyls, hyracoids, cetaceans and primates), (8) presence of a complete secondary (SII) somatic sensory region of cerebral cortex (therians-all extant mammals other than monotremes), and (9) presence of a distinct external cuneate nucleus among the nuclei of the dorsal columns (all mammalian groups except monotremes and sirenians). Two examples of phylogenetic trees derived from these data are presented. These sample trees maintain the segregation of the monotremes and the marsupials, and the basic dichotomy of placentals seen in our earlier trees based entirely on brain data. They also show: an orderly sequence of bifurcations (rather than the commonly seen multifurcation near the base of the radiation) in the reconstruction of placental relationships; extremes of derivation for the Cetacea, the Chiroptera, and the Sirenia (in concordance with trees based on other data); a ferungulate association of Carnivora, Perissodactyla, Artiodactyla, Hyracoidea and Sirenia; and an assemblage of related Dermoptera, Primates, Scandentia, and Chiroptera which in this model also includes Insectivora and Macroscelidea. Analyses based on brain characters can reinforce conclusions based on other data, while at the same time introducing new ideas about relationships. Neural traits provide a source of data independent of those commonly used in phylogenetic analysis, and are extremely valuable for testing old hypotheses and for introducing new ones.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗