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Heterogeneous but conserved natural killer receptor gene complexes in four major orders of mammals.

The natural killer (NK) receptor gene complex (NKC) encodes a large number of C-type lectin-like receptors, which are expressed on NK and other immune-related cells. These receptors play an important role in regulating NK-cell cytolytic activity, protecting cells against virus infection and tumorigenesis. To understand the evolutionary history of the NKC, we characterized the C-type lectin-like NKC genes and their organization from four major orders of placental mammals, primates (human), rodents (mouse and rat), carnivores (dog), and artiodactyls (cattle) and then conducted phylogenetic analysis of these genes. The results indicate that the NKC of placental mammals is highly heterogeneous in terms of the gene content and rates of birth and death of different gene lineages, but the NKC is also remarkably conserved in its gene organization and persistence of orthologous gene lineages. Among the 28 identified NKC gene lineages, 4, KLRA1, KLRB1, CLEC2D, and CLEC4A/B/C, have expanded rapidly in rodents only. The high birth and death rate of these 4 gene families might be due to functional differentiation driven by positive selection. Identification of putative NKC sequences in opossum and chicken genomes implies that the expansion of the NKC gene families might have occurred before the radiation of placental mammals but after the divergence of birds from mammals.

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Revised age estimates for the later Paleogene mammal faunas of Egypt and Oman.

The Jebel Qatrani Formation of northern Egypt has produced Afro-Arabia's primary record of Paleogene mammalian evolution, including the world's most complete remains of early anthropoid primates. Recent studies of Fayum mammals have assumed that the Jebel Qatrani Formation contains a significant Eocene component ( approximately 150 of 340 m), and that most taxa from that succession are between 35.4 and 33.3 million years old (Ma), i.e., latest Eocene to earliest Oligocene in age. Reanalysis of the chronological evidence shared by later Paleogene strata exposed in Egypt and Oman (Taqah and Thaytiniti areas, Dhofar Province) reveals that this hypothesis is no longer tenable. Revised correlation of the Fayum and Dhofar magnetostratigraphies indicates that (i) only the lowest 48 m of the Jebel Qatrani Formation are likely to be Eocene in age; (ii) the youngest Fayum anthropoids, including well known species such as Aegyptopithecus zeuxis and Apidium phiomense, are probably between 30.2 and 29.5 Ma, approximately 3-4 Ma younger than previously thought; (iii) oligopithecid anthropoids did not go extinct at the Eocene-Oligocene boundary but rather persisted for at least another 2.5 Ma; (iv) propliopithecid anthropoids first appear in the Fayum area at approximately 31.5 Ma, long after the Eocene-Oligocene boundary; and (v) the youngest Fayum mammals may be only approximately 1 Ma older than the 28- to 27-Ma mammals from Chilga, Ethiopia, and not 4-5 Ma older, as previously thought. Whatever gap exists in the Oligocene record of Afro-Arabian mammal evolution is now limited primarily to a poorly sampled 27- to 23-Ma window in the latest Oligocene.

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Global mammal distributions, biodiversity hotspots, and conservation.

Hotspots, which have played a central role in the selection of sites for reserves, require careful rethinking. We carried out a global examination of distributions of all nonmarine mammals to determine patterns of species richness, endemism, and endangerment, and to evaluate the degree of congruence among hotspots of these three measures of diversity in mammals. We then compare congruence of hotspots in two animal groups (mammals and birds) to assess the generality of these patterns. We defined hotspots as the richest 2.5% of cells in a global equal-area grid comparable to 1 degrees latitude x 1 degrees longitude. Hotspots of species richness, "endemism," and extinction threat were noncongruent. Only 1% of cells and 16% of species were common to the three types of mammalian hotspots. Congruence increased with increases in both the geographic scope of the analysis and the percentage of cells defined as being hotspots. The within-mammal hotspot noncongruence was similar to the pattern recently found for birds. Thus, assigning global conservation priorities based on hotspots is at best a limited strategy.

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Ticks associated with wild mammals in Ghana.

The host ranges of a collection of 21 tick species found on wild mammals in the savanna, forests and coastal zone of Ghana suggested that most species were adapted to feeding mainly on host species within a single mammalian order, i.e. on artiodactyls (bovids/suids), carnivores, rodents or pholidotes (pangolins). Only a few species were dispersed evenly across a range of orders. Seven out of ten of the most common ticks on forest mammals were significantly associated with a particular host species or a group of closely related host species, which could be viewed as their major host or hosts, but they were also recorded much less frequently on a wide range of host species. Two other species were confined to their major hosts. Only one species appeared to be widely dispersed on forest mammals and to lack a particular major host. The majority of tick species therefore occurred on hosts with very distinctive biological, behavioural and ecological characteristics. The study provided no evidence to support the view that host specificity is an artefact of sampling. Finding that the tick species on Ghanaian wild mammals occurred on particular hosts, as well as in distinct habitats, indicated that tick-host associations are important for tick survival and confirmed the importance of climate and vegetation in tick distribution.

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Mammals in which females are larger than males.

Females are larger than males in more species of mammals than is generally supposed. A provisional list of the mammalian cases is provided. The phenomenon is not correlated with an unusually large degree of male parental investment, polyandry, greater aggressiveness in females than in males, greater development of weapons in females, female dominance, or matriarchy. The phenomenon may have evolved in a variety of ways, but it is rarely, if ever, the result of sexual selection acting upon the female sex. The most common selective pressures favoring large size in female mammals are probably those associated with the fact that a big mother is often a better mother and those resulting from more intense competintion among females for some resource than among males. It appears that, in general, more than one such pressure must affect the females of a species, and that their combined effects must not be countered by even stronger selective pressures favoring large size in males, before the result is that of larger size in the female sex. Sexual selection may often be operating upon the male sux in mammals even when it is smaller. Present knowledge about the species of mammals in which females are lager than males is quite rudimentary. Much more information is needed before we will be able to speak of the selective pressures accounting for the phenomenon with any reasomable degree of certainty. Perhaps the most fruitful approach would be a series of field studies of groups of related species in which females are larger in some species and males are larger in others.

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Host associations of ticks (Acari: Ixodidae) parasitizing medium-sized mammals in a Lyme disease endemic area of southern New York.

Ticks of eight medium-sized mammal species were studied in an area of endemic Lyme disease in Westchester County, N.Y., from 1 April 1984 to 31 March 1985. Most (81%) of the 266 total mammal captures were raccoon, Procyon lotor (L.) (47%), opossum, Didelphis virginiana (Kerr) (19%), and striped skunk, Mephitis mephitis Schreber (15%); these host species accounted for 91% of the 1,519 ticks collected. Although the total number of ticks was rather evenly distributed among these mammals, species composition of ticks on each host species differed markedly. Ixodes dammini Spielman, Clifford, Piesman, and Corwin was the most abundant tick species and accounted for 45% (690) of the total ticks collected. Immatures were most prevalent (56%) on opossum, and nearly all (86%) adults were found on this host species. I. cookei Marx was second in abundance (34%) and was most prevalent (60%) on skunk. I. texanus Banks and Dermacentor variabilis (Say) were less abundant (less than 20% collectively) and were most prevalent on raccoon. I. dentatus Marx on eastern cottontail, Sylvilagus floridanus (Allen), and I. marxi Banks on gray squirrel, Sciurus carolinensis Gmelin, were least abundant (less than 2% collectively). The prevalence of I. dammini on medium-sized mammals in southern New York may influence the epizoötiology of Lyme disease.

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Cytochrome b phylogeny and the taxonomy of great apes and mammals.

In the Linnaean system of classification, the generic status of a species is part of its binomial name, and it is therefore important that the classification at the level of genus is consistent at least in related groups of organisms. Using maximum-likelihood phylogenetic trees constructed from a large number of complete or nearly complete mammalian cytochrome b sequences, I show that the distributions of intrageneric and intergeneric distances derived from these trees are clearly separated, which allows the limits for a more rational generic classification of mammals to be established. The analysis of genetic distances among hominids in this context provides strong support for the inclusion of humans and chimpanzees in the same genus. It is also of interest to decipher the main reasons for the possible biases in the mammalian classification. I found by correlation analysis that the classification of mammals of large body size tends to be oversplit, whereas that of small mammals has an excess of lumping, which may be a manifestation of the larger difficulty in finding diagnostic characters in the classification of small animals. In addition, and contrary to some previous observations, there is no correlation between body size and rate of cytochrome b evolution in mammals, which excludes the difference in evolutionary rates as the cause of the observed body size taxonomic bias.

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Four new mitochondrial genomes and the increased stability of evolutionary trees of mammals from improved taxon sampling.

We have sequenced four new mitochondrial genomes to improve the stability of the tree for placental mammals; they are two insectivores (a gymnure, Echinosorex gymnurus and Formosan shrew Soriculus fumidus); a Formosan lesser horseshoe bat (Rhinolophus monoceros); and the New Zealand fur seal (Arctocephalus forsteri). A revision to the hedgehog sequence (Erinaceus europaeus) is also reported. All five are from the Laurasiatheria grouping of eutherian mammals. On this new data set there is a strong tendency for the hedgehog and its relative, the gymnure, to join with the other Laurasiatherian insectivores (mole and shrews). To quantify the stability of trees from this data we define, based on nuclear sequences, a major four-way split in Laurasiatherians. This ([Xenarthra, Afrotheria], [Laurasiatheria, Supraprimates]) split is also found from mitochondrial genomes using either protein-coding or RNA (rRNA and tRNA) data sets. The high similarity of the mitochondrial and nuclear-derived trees allows a quantitative estimate of the stability of trees from independent data sets, as detected from a triplet Markov analysis. There are significant changes in the mutational processes within placental mammals that are ignored by current tree programs. On the basis of our quantitative results, we expect the evolutionary tree for mammals to be resolved quickly, and this will allow other problems to be solved.

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The maximum oxygen consumption and aerobic scope of birds and mammals: getting to the heart of the matter.

Resting or basal metabolic rates, compared across a wide range of organisms, scale with respect to body mass as approximately the 0.75 power. This relationship has recently been linked to the fractal geometry of the appropriate transport system or, in the case of birds and mammals, the blood vascular system. However, the structural features of the blood vascular system should more closely reflect maximal aerobic metabolic rates rather than submaximal function. Thus, the maximal aerobic metabolic rates of birds and mammals should also scale as approximately the 0.75 power. A review of the literature on maximal oxygen consumption and factorial aerobic scope (maximum oxygen consumption divided by basal metabolic rate) suggests that body mass influences the capacity of the cardiovascular system to raise metabolic rates above those at rest. The results show that the maximum sustainable metabolic rates of both birds and mammals are similar and scale as approximately the 0.88 +/- 0.02 power of body mass (and aerobic scope as approximately the 0.15 +/- 0.05 power), when the measurements are standardized with respect to the differences in relative heart mass and haemoglobin concentration between species. The maximum heart beat frequency of birds and mammals is predicted to scale as the -0.12 +/- 0.02 power of body mass, while that at rest should scale as -0.27 +/- 0.04.

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The postcranial skeletons of the Triassic mammals Eozostrodon, Megazostrodon and Erythrotherium.

The purposes of this monograph are to describe the postcranial skeletons of the earliest known mammals, and to probe, in so far as possible by osteological study, biological questions concerning the habits and adaptations of these late Triassic forms. In this context, information on the background of this investigation is useful. Studies of Mesozoic mammals, begun some 150 years ago, are based on rare and fragmentary fossils, principally jaws and teeth. These investigations have yielded a bare outline of some 120 million years of mammalian evolution-about two-thirds of mammalian history. No assessment of the important biological changes occurring during this time can ever be complete, but major advances are possible as new discoveries provide material that is more complete or that represents a previously unknown evolutionary stage. So tenuous is the evidence that at least some concepts are re-evaluated with each discovery. Postcranial anatomy offers especially intriguing prospects for investigation because associated material (that can be positively assigned to a taxon below subclass) has been for the most part unknown, and indeed even dissociated bones are a rarity. Since G.G. Simpson's monographs of 1928 and 1929, progress in the study of Mesozoic mammals has been largely dependent on new finds. A major impetus to renewed investigation came from the discoveries of Mesozoic mammals by Walter Kühne in 1939 and during the immediate post-war years. Kühne first worked on fissures in the Carboniferous limestone quarries at Frome, Somerset, in southwest England where he collected a series of teeth of the problematical form Haramiya and two triconodont teeth which were placed in the genus Eozostrodon (Parrington 1941, 1946). The fissure faunas are generally thought to be of Upper Triassic (Rhaetic) age (Kühne 1946), although Kermack, Musset & Rigney (1973) believe that the evidence is insufficient to determine whether the deposits are Rhaetic or Lower Liassic. After the war Kühne carried his explorations farther west, eventually reaching the quarries at Bridgend in Glamorgan, Wales, where he not only found more triconodont teeth in some quantity (Kühne 1958) but also a symmetrodont tooth (Kühne 1950). Shortly after making these discoveries, Kühne returned to Germany and the work was continued by a team from University College, London, under the leadership of Dr K.A. Kermack.

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The origin of human chromosome 1 and its homologs in placental mammals.

Developing ordered gene maps from multiple mammalian species coupled with chromosome-painting data provide a powerful resource for resolving the evolutionary history of chromosomes and whole genomes. In this work, we recapitulate the evolutionary history of human chromosome 1 and its homologs in placental mammals, putatively the largest physical unit in the ancestral placental genome. Precise definition of translocation exchange breakpoints in human, carnivore, cetartiodactyl, and rodent-ordered gene maps demonstrate that chromosome breakpoints, previously considered as equivalent, actually represent distinct chromosome positions and exchange events. Multidirectional chromosome painting, using probes from homologs to chromosome 1 in seven mammal species from six orders of placental mammals, confirm the gene-mapping results and indicate that the multiple human chromosome 1 homologs in these species are derived from independent fissions of a single ancestral chromosome. Chromosome painting using human chromosome 1 probes identifies a single human chromosome 1 homolog in phylogenetically distant taxa, the two-toed sloth, cetaceans, and higher primates. The diverse phylogenetic occurrence of a single Hsa1 synteny among the major clades of placental mammals suggests that human chromosome 1 represents an intact ancestral chromosome, which was variously fissioned in the majority of placental species. We find that the number of human chromosome 1 fissions in a specific lineage reflects its general rate of genomic evolution. Further, historic chromosome exchange appears to have been disproportionately clustered in two breakpoint hotspots on the long arm.

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The evolution of body armor in mammals: plantigrade constraints of large body size.

Predictions associated with opposing selection generating minimum variance in basal metabolic rate (BMR) in mammals at a constrained body mass (CBM; 358 g) were tested. The CBM is presumed to be associated with energetic constraints linked to predation and variable resources at intermediate sizes on a logarithmic mass scale. Opposing selection is thought to occur in response to energetic constraints associated with predation and unpredictable resources. As body size approaches and exceeds the CBM, mammals face increasing risks of predation and daily energy requirements. Fast running speeds may require high BMRs, but unpredictable and low resources may select for low BMRs, which also reduce foraging time and distances and thus predation risks. If these two selection forces oppose each other persistently, minimum BMR variance may result. However, extreme BMR outliers at and close to the CBM should be indicative of unbalanced selection and predator avoidance alternatives (escapers vs. defenders), and may therefore provide indirect support for opposing selection. It was confirmed that body armor in defenders evolves at and above the CBM, and armored mammals had significantly lower BMRs than their nonarmored counterparts. However, analyses comparing the BMR of escapers--the fastest nonarmored runners (Lagomorpha)--with similar-sized counterparts were inconclusive and were confounded by limb morphology associated with speed optimization. These analyses suggest that the risks and costs of predation and the speed limitations of the plantigrade foot may constrain the evolution of large body sizes in plantigrade mammals.

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Owl pellets as a source of DNA for genetic studies of small mammals.

Owl pellets contain a good skeletal record of the small mammals consumed, and correspond to the undigested portions of prey which are regurgitated. These pellets are easy to find at the roosting site of owls. As it has been demonstrated that amplifiable DNA can be isolated from ancient bone remains, the possibility of using owl pellets as a source of DNA for small mammal genetics studies via the polymerase chain reaction has been investigated. The main uncertainties when isolating DNA from such a material are firstly the possibility that the extracted DNA would be too degraded during the digestion in the stomach of the owl, and secondly that extensive cross-contaminations could occur among the different prey consumed. The results obtained clearly demonstrate that cross-contamination does not occur, and that mitochondrial and nuclear DNA can be amplified using skulls of small mammals found in owl pellets as a source of DNA. The relative efficiency of two methods of DNA extraction is estimated and discussed. Thus, owl pellets represent a non-invasive sampling technique which provides a valuable source of DNA for studying population genetics of small mammals.

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Single nucleotide polymorphism (SNP) discovery in mammals: a targeted-gene approach.

Single nucleotide polymorphisms (SNPs) have rarely been exploited in nonhuman and nonmodel organism genetic studies. This is due partly to difficulties in finding SNPs in species where little DNA sequence data exist, as well as to a lack of robust and inexpensive genotyping methods. We have explored one SNP discovery method for molecular ecology, evolution, and conservation studies to evaluate the method and its limitations for population genetics in mammals. We made use of 'CATS' (or 'EPIC') primers to screen for novel SNPs in mammals. Most of these primer sets were designed from primates and/or rodents, for amplifying intron regions from conserved genes. We have screened 202 loci in 16 representatives of the major mammalian clades. Polymerase chain reaction (PCR) success correlated with phylogenetic distance from the human and mouse sequences used to design most primers; for example, specific PCR products from primates and the mouse amplified the most consistently and the marsupial and armadillo amplifications were least successful. Approximately 24% (opossum) to 65% (chimpanzee) of primers produced usable PCR product(s) in the mammals tested. Products produced generally high but variable levels of readable sequence and similarity to the expected genes. In a preliminary screen of chimpanzee DNA, 12 SNPs were identified from six (of 11) sequenced regions, yielding a SNP on average every 400 base pairs (bp). Given the progress in genome sequencing, and the large numbers of CATS-like primers published to date, this approach may yield sufficient SNPs per species for population and conservation genetic studies in nonmodel mammals and other organisms.

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Population cycles in northern small mammals.

I. The regular multiannual oscillations of small mammals at northern latitudes have been a subject of intensive study from the beginning of this century. The existence of a subjective bias in the research due to different schools of study together with a long series of failures and seemingly contradictory results in experiments testing a multitude of hypotheses have brought confusion to the field of study. Much of this confusion has resulted from a failure to recognize sharply the problem studied, which in turn has masked the progress made during the years. Northern mammal cycles are not a single problem but a composition of many related problems. Every problem may have a single-factor explanation, but even with a single-factor explanation, one factor is not necessarily an answer to all of the related problems. 2. At present, we can state that the cyclicity is caused by a predator-prey interaction. Both the 8-11-year and the 3-5-year cycles may be special cases of a more general cycle, most likely caused by a herbivore-resident specialist predator interaction, where the period of the cycles is determined by size-related constraints affecting the increase rate of the populations. The factors determining the amplitude of the cycles probably vary regionally and/or temporally. The operation of generalist and nomadic predators is largely responsible for the regional and geographic synchrony in cycles, although climatic factors may also contribute to the geographic synchrony. The northern distribution of animal communities; both these factors affect the density of generalist predators, which act as a stabilizing factor in the system. The age-related survival pattern seems to be mainly caused by predation, and the cyclically fluctuating reproductive output and mean body mass may be caused by changes in prey behaviour in response to fluctuating predation risk. Thus, we can already give a plausible explanation for most problems related to northern mammal cycles. 3. In all problems discussed, predation seems to be involved, and in most problems, it seems to be the factor which explains the observed patterns. Thus, as a generalization, it can be said that predation seems to be the key factor in the explanation of the northern multiannual cycles of small mammals. 4. There seems to be a linkage between diversity and cyclicity, probably because the diversity of the community (the number of prey species available) may determine the diet choice of a predator, which in turn determines whether the predators have a stabilizing or a destabilizing impact on prey populations.(ABSTRACT TRUNCATED AT 400 WORDS)

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A Late Jurassic digging mammal and early mammalian diversification.

A fossil mammal from the Late Jurassic Morrison Formation, Colorado, has highly specialized teeth similar to those of xenarthran and tubulidentate placental mammals and different from the generalized insectivorous or omnivorous dentitions of other Jurassic mammals. It has many forelimb features specialized for digging, and its lumbar vertebrae show xenarthrous articulations. Parsimony analysis suggests that this fossil represents a separate basal mammalian lineage with some dental and vertebral convergences to those of modern xenarthran placentals, and reveals a previously unknown ecomorph of early mammals.

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An IS711 element downstream of the bp26 gene is a specific marker of Brucella spp. isolated from marine mammals.

DNA polymorphism of the bp26 gene, coding for a diagnostic protein antigen for brucellosis, was assessed by PCR and restriction fragment length polymorphism analysis using primers to amplify the bp26 gene with its flanking regions. Surprisingly, whereas PCR performed on DNA of the reference strains of the six recognized Brucella species produced a product of the expected size (1,029 bp), PCR performed on DNA of three representative strains from marine mammals (from a seal, a dolphin, and a porpoise) produced a larger product, of about 1,900 bp. Nucleotide sequencing of the 1,900-bp PCR products revealed the presence of an insertion sequence, IS711, downstream of the bp26 gene and adjacent to a Bru-RS1 element previously described as being a hot spot for IS711 insertion. PCR performed on a large number of field strains from different geographic origins and from marine mammal isolates indicated that the occurrence of an IS711 element downstream of the bp26 gene was a feature specific to the marine mammal Brucella strains. Thus, this PCR assay is able to differentiate Brucella terrestrial isolates from marine mammal isolates and could be applied for diagnostic purposes.

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Natural infection of small mammal species in Minnesota with the agent of human granulocytic ehrlichiosis.

The natural reservoirs for the agent of human granulocytic ehrlichiosis (HGE) are suspected to be the small mammals that host immature stages of Ixodes scapularis ticks. To determine if such small mammals are naturally infected, we collected blood and serum samples from small mammal species in rural and suburban areas of Minneapolis and St. Paul, Minn. Samples were collected from white-footed mice (Peromyscus leucopus), eastern chipmunks (Tamias striatus), southern red-backed voles (Clethrionomys gapperi), and insectivorous shrews (Blarina brevicauda and Sorex cinereus). Blood samples were tested by PCR for active infection with the HGE agent, and sera from P. leucopus mice were tested for serologic evidence of infection by indirect immunofluorescence. PCR analyses revealed the presence of HGE agent DNA in 20 of the 190 samples (10.5%) tested. Of the 119 P. leucopus mouse serum samples that were analyzed, 12 (10.1%) contained Ehrlichia equi antibodies. In 3 of 119 (2.5%) P. leucopus mice from which both blood and serum were collected. HGE agent DNA and antibodies against E. equi were present. Animals with evidence of infection with the HGE agent are widely distributed around the Minneapolis-St. Paul area in regions with known I. scapularis tick activity. Small mammals that are frequent hosts for larval I. scapularis ticks and that are found in areas where HGE occurs are likely to be a major reservoir from which infected ticks that bite humans are derived.

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