Elephant
Bibliographic
Database

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Elephant Bibliographic Database
www.elephantcare.org

References updated October 2009 by date of publication, most recent first.

Cerling, T.E., Wittemyer, G., Ehleringer, J.R., Remien, C.H., Douglas-Hamilton, I., 2009. History of Animals using Isotope Records (HAIR): a 6-year dietary history of one family of African elephants
76. Proc. Natl. Acad. Sci. U. S. A 106, 8093-8100.
Abstract: The dietary and movement history of individual animals can be studied using stable isotope records in animal tissues, providing insight into long-term ecological dynamics and a species niche. We provide a 6-year history of elephant diet by examining tail hair collected from 4 elephants in the same social family unit in northern Kenya. Sequential measurements of carbon, nitrogen, and hydrogen isotope rations in hair provide a weekly record of diet and water resources. Carbon isotope ratios were well correlated with satellite-based measurements of the normalized difference vegetation index (NDVI) of the region occupied by the elephants as recorded by the global positioning system (GPS) movement record; the absolute amount of C(4) grass consumption is well correlated with the maximum value of NDVI during individual wet seasons. Changes in hydrogen isotope ratios coincided very closely in time with seasonal fluctuations in rainfall and NDVI whereas diet shifts to relatively high proportions of grass lagged seasonal increases in NDVI by approximately 2 weeks. The peak probability of conception in the population occurred approximately 3 weeks after peak grazing. Spatial and temporal patterns of resource use show that the only period of pure browsing by the focal elephants was located in an over-grazed, communally managed region outside the protected area. The ability to extract time-specific longitudinal records on animal diets, and therefore the ecological history of an organism and its environment, provides an avenue for understanding the impact of climate dynamics and land-use change on animal foraging behavior and habitat relations

Clark, C.J., Poulsen, J.R., Malonga, R., Elkan, P.W., Jr., 2009. Logging concessions can extend the conservation estate for Central African tropical forests
56. Conserv. Biol. 23, 1281-1293.
Abstract: The management of tropical forest in timber concessions has been proposed as a solution to prevent further biodiversity loss. The effectiveness of this strategy will likely depend on species-specific, population-level responses to logging. We conducted a survey (749 line transects over 3450 km) in logging concessions (1.2 million ha) in the northern Republic of Congo to examine the impact of logging on large mammal populations, including endangered species such as the elephant (Loxodonta africana), gorilla (Gorilla gorilla), chimpanzee (Pan troglodytes), and bongo (Tragelaphus eurycerus). When we estimated species abundance without consideration of transect characteristics, species abundances in logged and unlogged forests were not different for most species. When we modeled the data with a hurdle model approach, however, analyzing species presence and conditional abundance separately with generalized additive models and then combining them to calculate the mean species abundance, species abundance varied strongly depending on transect characteristics. The mean species abundance was often related to the distance to unlogged forest, which suggests that intact forest serves as source habitat for several species. The mean species abundance responded nonlinearly to logging history, changing over 30 years as the forest recovered from logging. Finally the distance away from roads, natural forest clearings, and villages also determined the abundance of mammals. Our results suggest that logged forest can extend the conservation estate for many of Central Africa's most threatened species if managed appropriately. In addition to limiting hunting, logging concessions must be large, contain patches of unlogged forest, and include forest with different logging histories

Freeman, E.W., Guagnano, G., Olson, D., Keele, M., Brown, J.L., 2009. Social factors influence ovarian acyclicity in captive African elephants (Loxodonta africana). Zoo. Biol. 28, 1-15.
Abstract: Nearly one-third of reproductive age African elephants in North America that are hormonally monitored fail to exhibit estrous cycle activity, which exacerbates the nonsustainability of the captive population. Three surveys were distributed to facilities housing female African elephants to determine how social and environmental variables contribute to cyclicity problems. Forty-six facilities returned all three surveys providing information on 90% of the SSP population and 106 elephants (64 cycling, 27 noncycling and 15 undetermined). Logistic analyses found that some physiological and social history variables were related to ovarian acyclicity. Females more likely to be acyclic had a larger body mass index and had resided longer at a facility with the same herdmates. Results suggest that controlling the weight of an elephant might be a first step to helping mitigate estrous cycle problems. Data further show that transferring females among facilities has no major impact on ovarian activity. Last, social status appears to impact cyclicity status; at 19 of 21 facilities that housed both cycling and noncycling elephants, the dominant female was acyclic. Further studies on how social and environmental dynamics affect hormone levels in free-living, cycling elephants are needed to determine whether acyclicity is strictly a captivity-related phenomenon

Fulka, J., Jr., Loi, P., Ptak, G., Fulka, H., John, J.S., 2009. Hope for the mammoth? Cloning Stem Cells 11, 1-4.

Gheerbrant, E., 2009. Paleocene emergence of elephant relatives and the rapid radiation of African ungulates
43. Proc. Natl. Acad. Sci. U. S. A 106, 10717-10721.
Abstract: Elephants are the only living representatives of the Proboscidea, a formerly diverse mammalian order whose history began with the 55-million years (mys) old Phosphatherium. Reported here is the discovery from the early late Paleocene of Morocco, ca. 60 mys, of the oldest and most primitive elephant relative, Eritherium azzouzorum n.g., n.sp., which is one of the earliest known representatives of modern placental orders. This well supported stem proboscidean is extraordinarily primitive and condylarth-like. It provides the first dental evidence of a resemblance between the proboscideans and African ungulates (paenungulates) on the one hand and the louisinines and early macroscelideans on the other. Eritherium illustrates the origin of the elephant order at a previously unknown primitive stage among paenungulates and "ungulates." The primitive morphology of Eritherium suggests a recent and rapid paenungulate radiation after the Cretaceous-Tertiary boundary, probably favoured by early endemic African paleoecosystems. At a broader scale, Eritherium provides a new old calibration point of the placental tree and supports an explosive placental radiation. The Ouled Abdoun basin, which yields the oldest known African placentals, is a key locality for elucidating phylogeny and early evolution of paenungulates and other related endemic African lineages

Greenwald, R., Lyashchenko, O., Esfandiari, J., Miller, M., Mikota, S., Olsen, J.H., Ball, R., Dumonceaux, G., Schmitt, D., Moller, T., Payeur, J.B., Harris, B., Sofranko, D., Waters, W.R., Lyaschenko, K.P., 2009. Highly accurate antibody assays for early and rapid detection of tuberculosis in African and Asian elephants. Clinical and Vaccine Immunology 16, 605-612.
Abstract: Tuberculosis (TB) in elephants is a reemerging zoonotic disease caused primarily by Mycobacterium tuberculosis. Current methods for screening and diagnosis rely on trunk wash culture, which has serious limitations due to low test sensitivity, slow turnaround time, and variable sample quality. Innovative and more efficient diagnostic tools are urgently needed. We describe three novel serologic techniques, the ElephantTB Stat-Pak kit, multiantigen print immunoassay, and dual-path platform VetTB test, for rapid antibody detection in elephants. The study was performed with serum samples from 236 captive African and Asian elephants from 53 different locations in the United States and Europe. The elephants were divided into three groups based on disease status and history of exposure: (i) 26 animals with culture-confirmed TB due to M. tuberculosis or Mycobacterium bovis, (ii) 63 exposed elephants from known-infected herds that had never produced a culture-positive result from trunk wash samples, and (iii) 147 elephants without clinical symptoms suggestive of TB, with consistently negative trunk wash culture results, and with no history of potential exposure to TB in the past 5 years. Elephants with culture-confirmed TB and a proportion of exposed but trunk wash culture-negative elephants produced robust antibody responses to multiple antigens of M. tuberculosis, with seroconversions detectable years before TB-positive cultures were obtained from trunk wash specimens. ESAT-6 and CFP10 proteins were immunodominant antigens recognized by elephant antibodies during disease. The serologic assays demonstrated 100% sensitivity and 95 to 100% specificity. Rapid and accurate antibody tests to identify infected elephants will likely allow earlier and more efficient treatment, thus limiting transmission of infection to other susceptible animals and to humans.

Greenwald, R., Lyashchenko, O., Esfandiari, J., Miller, M., Mikota, S., Olsen, J.H., Ball, R., Dumonceaux, G., Schmitt, D., Moller, T., Payeur, J.B., Harris, B., Sofranko, D., Waters, W.R., Lyashchenko, K.P., 2009. Highly accurate antibody assays for early and rapid detection of tuberculosis in African and Asian elephants. Clin. Vaccine Immunol. 16, 605-612.
Abstract: Tuberculosis (TB) in elephants is a reemerging zoonotic disease caused primarily by Mycobacterium tuberculosis. Current methods for screening and diagnosis rely on trunk wash culture, which has serious limitations due to low test sensitivity, slow turnaround time, and variable sample quality. Innovative and more efficient diagnostic tools are urgently needed. We describe three novel serologic techniques, the ElephantTB Stat-Pak kit, multiantigen print immunoassay, and dual-path platform VetTB test, for rapid antibody detection in elephants. The study was performed with serum samples from 236 captive African and Asian elephants from 53 different locations in the United States and Europe. The elephants were divided into three groups based on disease status and history of exposure: (i) 26 animals with culture-confirmed TB due to M. tuberculosis or Mycobacterium bovis, (ii) 63 exposed elephants from known-infected herds that had never produced a culture-positive result from trunk wash samples, and (iii) 147 elephants without clinical symptoms suggestive of TB, with consistently negative trunk wash culture results, and with no history of potential exposure to TB in the past 5 years. Elephants with culture-confirmed TB and a proportion of exposed but trunk wash culture-negative elephants produced robust antibody responses to multiple antigens of M. tuberculosis, with seroconversions detectable years before TB-positive cultures were obtained from trunk wash specimens. ESAT-6 and CFP10 proteins were immunodominant antigens recognized by elephant antibodies during disease. The serologic assays demonstrated 100% sensitivity and 95 to 100% specificity. Rapid and accurate antibody tests to identify infected elephants will likely allow earlier and more efficient treatment, thus limiting transmission of infection to other susceptible animals and to humans

Grus, W.E., Zhang, J., 2009. Origin of the genetic components of the vomeronasal system in the common ancestor of all extant vertebrates. Molecular Biology and Evolution 26, 407-419.
Abstract: Comparative genomics provides a valuable tool for inferring the evolutionary history of physiological systems, particularly when this information is difficult to ascertain by morphological traits. One such example is the vomeronasal system (VNS), a vertebrate nasal chemosensory system that is responsible for detecting intraspecific pheromonal cues as well as environmental odorants. The morphological components of the VNS are found only in tetrapods, but the genetic components of the system have been found in teleost fish, in addition to tetrapods. To determine when the genetic components of the VNS originated, we searched for the VNS-specific genes in the genomes of two early diverging vertebrate lineages: the sea lamprey from jawless fishes and the elephant shark from cartilaginous fishes. Genes encoding vomeronasal type 1 receptors (V1Rs) and Trpc2, two components of the vomeronasal signaling pathway, are present in the sea lamprey genome, and both are expressed in the olfactory organ, revealing that the genetic components of the present-day VNS existed in the common ancestor of all extant vertebrates. Additionally, all three VNS genes, Trpc2, V1Rs, and vomeronasal type 2 receptors (V2Rs), are found in the elephant shark genome. Because V1Rs and V2Rs are related to two families of taste receptors, we also searched the early diverging vertebrate genomes for taste system genes and found them in the shark genome but not in the lamprey. Coupled with known distributions of the genetic components of the vertebrate main olfactory system, our results suggest staggered origins of vertebrate sensory systems. These findings are important for understanding the evolution of vertebrate sensory systems and illustrate the utility of the genome sequences of early diverging vertebrates for uncovering the evolution of vertebrate-specific traits

Hakeem, A.Y., Sherwood, C.C., Bonar, C.J., Butti, C., Hof, P.R., Allman, J.M., 2009. Von Economo neurons in the elephant brain. Anat. Rec. (Hoboken. ) 292, 242-248.
Abstract: Von Economo neurons (VENs), previously found in humans, all of the great ape species, and four cetacean species, are also present in African and Indian elephants. The VENs in the elephant are primarily found in similar locations to those in the other species. They are most abundant in the frontoinsular cortex (area FI) and are also present at lower density in the anterior cingulate cortex. Additionally, they are found in a dorsolateral prefrontal area and less abundantly in the region of the frontal pole. The VEN morphology appears to have arisen independently in hominids, cetaceans, and elephants, and may reflect a specialization for the rapid transmission of crucial social information in very large brains

Kaim, U., Paltian, V., Krudewig, C., Nieder, A., Wohlsein, P., 2009. Pulmonary aspergillosis in an African elephant (Loxodonta africana)
64. Dtsch. Tierarztl. Wochenschr. 116, 148-151.
Abstract: A 26-year-old female African elephant (Loxodonta africana) with a history of purulent pododermatitis, recurrent abdominal pain, and severe weight loss died spontaneously after a period of deteriorating disease. The main pathological finding was a severe bilateral pyogranulomatous, partially necrotizing pneumonia with numerous intralesional fungal hyphae. At microbiological examination Aspergillus spp. were isolated. The present case indicates that mycotic pneumonia should to be considered as a differential diagnosis of pulmonary disorders in elephants

Kun, A., Scheuring, I., 2009. Evolution of cooperation on dynamical graphs. Biosystems 96, 65-68.
Abstract:
There are two key characteristics of animal and human societies: (1) degree heterogeneity, meaning that not all individual have the same number of associates; and (2) the interaction topology is not static, i.e. either individuals interact with different set of individuals at different times of their life, or at least they have different associations than their parents. Earlier works have shown that population structure is one of the mechanisms promoting cooperation. However, most studies had assumed that the interaction network can be described by a regular graph (homogeneous degree distribution). Recently there are an increasing number of studies employing degree heterogeneous graphs to model interaction topology. But mostly the interaction topology was assumed to be static. Here we investigate the fixation probability of the cooperator strategy in the prisoner's dilemma, when interaction network is a random regular graph, a random graph or a scale-free graph and the interaction network is allowed to change.We show that the fixation probability of the cooperator strategy is lower when the interaction topology is described by a dynamical graph compared to a static graph. Even a limited network dynamics significantly decreases the fixation probability of cooperation, an effect that is mitigated stronger by degree heterogeneous networks topology than by a degree homogeneous one. We have also found that from the considered graph topologies the decrease of fixation probabilities due to graph dynamics is the lowest on scale-free graphs.

Leshchinskiy, S.V., 2009. Mineral deficiency, enzootic diseases and extinction of mammoth of northern Eurasia
82. Dokl. Biol. Sci. 424, 72-74.

Mueller, T. A near-perfect frozen mammoth resurfaces after 40,000 years, bearing clues to a great vanished species. National Geographic [May]. 2009.
Ref Type: Magazine Article

Murata, Y., Yonezawa, T., Kihara, I., Kashiwamura, T., Sugihara, Y., Nikaido, M., Okada, N., Endo, H., Hasegawa, M., 2009. Chronology of the extant African elephant species and case study of the species identification of the small African elephant with the molecular phylogenetic method
70. Gene 441, 176-186.
Abstract: Despite vigorous genetic studies of African elephants (Loxodonta africana and L. cyclotis) during the last decade, their evolutionary history is still obscure. Phylogenetic studies and coalescence time estimation using longer nucleotide sequence data from denser samplings are necessary to better understand the natural history of African elephants. Further, species identification among African elephants is sometimes very difficult using only the external morphological characteristics. This is a serious problem for making an adequate breeding plan in zoological gardens. In this paper, we investigated the continent-wide phylogeographical pattern of the African elephants and estimated the coalescence times among them. From these molecular data and geological evidence, we proposed an evolutionary scenario for the African elephants. We further demonstrated the effectiveness of molecular phylogenetic methods in species identification.

Opazo, J.C., Sloan, A.M., Campbell, K.L., Storz, J.F., 2009. Origin and ascendancy of a chimeric fusion gene: the beta/delta-globin gene of paenungulate mammals
84. Molecular Biology and Evolution 26, 1469-1478.
Abstract: The delta-globin gene (HBD) of eutherian mammals exhibits a propensity for recombinational exchange with the closely linked beta-globin gene (HBB) and has been independently converted by the HBB gene in multiple lineages. Here we report the presence of a chimeric beta/delta fusion gene in the African elephant (Loxodonta africana) that was created by unequal crossing-over between misaligned HBD and HBB paralogs. The recombinant chromosome that harbors the beta/delta fusion gene in elephants is structurally similar to the "anti-Lepore" duplication mutant of humans (the reciprocal exchange product of the hemoglobin Lepore deletion mutant). However, the situation in the African elephant is unique in that the chimeric beta/delta fusion gene supplanted the parental HBB gene and is therefore solely responsible for synthesizing the beta-chain subunits of adult hemoglobin. A phylogenetic survey of beta-like globin genes in afrotherian and xenarthran mammals revealed that the origin of the chimeric beta/delta fusion gene and the concomitant inactivation of the HBB gene predated the radiation of "Paenungulata," a clade of afrotherian mammals that includes three orders: Proboscidea (elephants), Sirenia (dugongs and manatees), and Hyracoidea (hyraxes). The reduced fitness of the human Hb Lepore deletion mutant helps to explain why independently derived beta/delta fusion genes (which occur on an anti-Lepore chromosome) have been fixed in a number of mammalian lineages, whereas the reciprocal delta/beta fusion gene (which occurs on a Lepore chromosome) has yet to be documented in any nonhuman mammal. This illustrates how the evolutionary fates of chimeric fusion genes can be strongly influenced by their recombinational mode of origin

Roca, A.L., Ishida, Y., Nikolaidis, N., Kolokotronis, S.O., Fratpietro, S., Stewardson, K., Hensley, S., Tisdale, M., Boeskorov, G., Greenwood, A.D., 2009. Genetic variation at hair length candidate genes in elephants and the extinct woolly mammoth. BMC. Evol. Biol. 9, 232.
Abstract: BACKGROUND: Like humans, the living elephants are unusual among mammals in being sparsely covered with hair. Relative to extant elephants, the extinct woolly mammoth, Mammuthus primigenius, had a dense hair cover and extremely long hair, which likely were adaptations to its subarctic habitat. The fibroblast growth factor 5 (FGF5) gene affects hair length in a diverse set of mammalian species. Mutations in FGF5 lead to recessive long hair phenotypes in mice, dogs, and cats; and the gene has been implicated in hair length variation in rabbits. Thus, FGF5 represents a leading candidate gene for the phenotypic differences in hair length notable between extant elephants and the woolly mammoth. We therefore sequenced the three exons (except for the 3' UTR) and a portion of the promoter of FGF5 from the living elephantid species (Asian, African savanna and African forest elephants) and, using protocols for ancient DNA, from a woolly mammoth. RESULTS: Between the extant elephants and the mammoth, two single base substitutions were observed in FGF5, neither of which alters the amino acid sequence. Modeling of the protein structure suggests that the elephantid proteins fold similarly to the human FGF5 protein. Bioinformatics analyses and DNA sequencing of another locus that has been implicated in hair cover in humans, type I hair keratin pseudogene (KRTHAP1), also yielded negative results. Interestingly, KRTHAP1 is a pseudogene in elephantids as in humans (although fully functional in non-human primates). CONCLUSION: The data suggest that the coding sequence of the FGF5 gene is not the critical determinant of hair length differences among elephantids. The results are discussed in the context of hairlessness among mammals and in terms of the potential impact of large body size, subarctic conditions, and an aquatic ancestor on hair cover in the Proboscidea

Rodriguez Delgado, C.L., Waters, P.D., Gilbert, C., Robinson, T.J., Graves, J.A., 2009. Physical mapping of the elephant X chromosome: conservation of gene order over 105 million years. Chromosome. Res.
Abstract: All therian mammals (eutherians and marsupials) have an XX female/XY male sex chromosome system or some variant of it. The X and Y evolved from a homologous pair of autosomes over the 166 million years since therian mammals diverged from monotremes. Comparing the sex chromosomes of eutherians and marsupials defined an ancient X conserved region that is shared between species of these mammalian clades. However, the eutherian X (and the Y) was augmented by a recent addition (XAR) that is autosomal in marsupials. XAR is part of the X in primates, rodents, and artiodactyls (which belong to the eutherian clade Boreoeutheria), but it is uncertain whether XAR is part of the X chromosome in more distantly related eutherian mammals. Here we report on the gene content and order on the X of the elephant (Loxodonta africana)-a representative of Afrotheria, a basal endemic clade of African mammals-and compare these findings to those of other documented eutherian species. A total of 17 genes were mapped to the elephant X chromosome. Our results support the hypothesis that the eutherian X and Y chromosomes were augmented by the addition of autosomal material prior to eutherian radiation. Not only does the elephant X bear the same suite of genes as other eutherian X chromosomes, but gene order appears to have been maintained across 105 million years of evolution, perhaps reflecting strong constraints posed by the eutherian X inactivation system

Schwarz, C., Debruyne, R., Kuch, M., McNally, E., Schwarcz, H., Aubrey, A.D., Bada, J., Poinar, H., 2009. New insights from old bones: DNA preservation and degradation in permafrost preserved mammoth remains. Nucleic Acids Res March 24.

Schwarz, C., Debruyne, R., Kuch, M., McNally, E., Schwarcz, H., Aubrey, A.D., Bada, J., Poinar, H., 2009. New insights from old bones: DNA preservation and degradation in permafrost preserved mammoth remains
89. Nucleic Acids Res. 37, 3215-3229.
Abstract: Despite being plagued by heavily degraded DNA in palaeontological remains, most studies addressing the state of DNA degradation have been limited to types of damage which do not pose a hindrance to Taq polymerase during PCR. Application of serial qPCR to the two fractions obtained during extraction (demineralization and protein digest) from six permafrost mammoth bones and one partially degraded modern elephant bone has enabled further insight into the changes which endogenous DNA is subjected to during diagenesis. We show here that both fractions exhibit individual qualities in terms of the prevailing type of DNA (i.e. mitochondrial versus nuclear DNA) as well as the extent of damage, and in addition observed a highly variable ratio of mitochondrial to nuclear DNA among the six mammoth samples. While there is evidence suggesting that mitochondrial DNA is better preserved than nuclear DNA in ancient permafrost samples, we find the initial DNA concentration in the bone tissue to be as relevant for the total accessible mitochondrial DNA as the extent of DNA degradation post-mortem. We also evaluate the general applicability of indirect measures of preservation such as amino-acid racemization, bone crystallinity index and thermal age to these exceptionally well-preserved samples

Sherwood, C.C., Stimpson, C.D., Butti, C., Bonar, C.J., Newton, A.L., Allman, J.M., Hof, P.R., 2009. Neocortical neuron types in Xenarthra and Afrotheria: implications for brain evolution in mammals. Brain Struct. Funct. 213, 301-328.
Abstract: Interpreting the evolution of neuronal types in the cerebral cortex of mammals requires information from a diversity of species. However, there is currently a paucity of data from the Xenarthra and Afrotheria, two major phylogenetic groups that diverged close to the base of the eutherian mammal adaptive radiation. In this study, we used immunohistochemistry to examine the distribution and morphology of neocortical neurons stained for nonphosphorylated neurofilament protein, calbindin, calretinin, parvalbumin, and neuropeptide Y in three xenarthran species-the giant anteater (Myrmecophaga tridactyla), the lesser anteater (Tamandua tetradactyla), and the two-toed sloth (Choloepus didactylus)-and two afrotherian species-the rock hyrax (Procavia capensis) and the black and rufous giant elephant shrew (Rhynchocyon petersi). We also studied the distribution and morphology of astrocytes using glial fibrillary acidic protein as a marker. In all of these species, nonphosphorylated neurofilament protein-immunoreactive neurons predominated in layer V. These neurons exhibited diverse morphologies with regional variation. Specifically, high proportions of atypical neurofilament-enriched neuron classes were observed, including extraverted neurons, inverted pyramidal neurons, fusiform neurons, and other multipolar types. In addition, many projection neurons in layers II-III were found to contain calbindin. Among interneurons, parvalbumin- and calbindin-expressing cells were generally denser compared to calretinin-immunoreactive cells. We traced the evolution of certain cortical architectural traits using phylogenetic analysis. Based on our reconstruction of character evolution, we found that the living xenarthrans and afrotherians show many similarities to the stem eutherian mammal, whereas other eutherian lineages display a greater number of derived traits

Vidya, T.N., Sukumar, R., Melnick, D.J., 2009. Range-wide mtDNA phylogeography yields insights into the origins of Asian elephants. Proc. Biol. Sci. 276, 893-902.
Abstract: Recent phylogeographic studies of the endangered Asian elephant (Elephas maximus) reveal two highly divergent mitochondrial DNA (mtDNA) lineages, an elucidation of which is central to understanding the species's evolution. Previous explanations for the divergent clades include introgression of mtDNA haplotypes between ancestral species, allopatric divergence of the clades between Sri Lanka or the Sunda region and the mainland, historical trade of elephants, and retention of divergent lineages due to large population sizes. However, these studies lacked data from India and Myanmar, which host approximately 70 per cent of all extant Asian elephants. In this paper, we analyse mtDNA sequence data from 534 Asian elephants across the species's range to explain the current distribution of the two divergent clades. Based on phylogenetic reconstructions, estimates of times of origin of clades, probable ancestral areas of origin inferred from dispersal-vicariance analyses and the available fossil record, we believe both clades originated from Elephas hysudricus. This probably occurred allopatrically in different glacial refugia, the alpha clade in the Myanmar region and the beta clade possibly in southern India-Sri Lanka, 1.6-2.1Myr ago. Results from nested clade and dispersal-vicariance analyses indicate a subsequent isolation and independent diversification of the beta clade in both Sri Lanka and the Sunda region, followed by northward expansion of the clade. We also find more recent population expansions in both clades based on mismatch distributions. We therefore suggest a contraction-expansion scenario during severe climatic oscillations of the Quaternary, with range expansions from different refugia during warmer interglacials leading to the varying geographical overlaps of the two mtDNA clades. We also demonstrate that trade in Asian elephants has not substantially altered the species's mtDNA population genetic structure

Wallis, M., 2009. Prolactin in the Afrotheria: characterization of genes encoding prolactin in elephant (Loxodonta africana), hyrax (Procavia capensis) and tenrec (Echinops telfairi). J. Endocrinol. 200, 233-240.
Abstract: Pituitary prolactin shows an episodic pattern of molecular evolution, with occasional short bursts of rapid change imposed on a generally rather slow evolutionary rate. In mammals, episodes of rapid change occurred in the evolution of primates, cetartiodactyls, rodents and the elephant. The bursts of rapid evolution in cetartiodactyls and rodents were followed by duplications of the prolactin gene that gave rise to large families of prolactin-related proteins including placental lactogens, while in primates the burst was followed by corresponding duplications of the related GH gene. The position in elephant is less clear. Extensive data relating to the genomic sequences of elephant and two additional members of the group Afrotheria are now available, and have been used here to characterize the prolactin genes in these species and explore whether additional prolactin-related genes are present. The results confirm the rapid evolution of elephant (Loxodonta africana) prolactin - the sequence of elephant prolactin is substantially different from that predicted for the ancestral placental mammal. Hyrax (Procavia capensis) prolactin is even more divergent but tenrec (Echinops telfairi) prolactin is strongly conserved. No evidence was obtained from searches of public databases for additional genes encoding prolactin-like proteins in any of these species. Detailed analysis of evolutionary rates, and other factors, indicates that the episode of rapid change in hyrax, and probably elephant, was adaptive, though the nature of the associated biological change(s) is not clear

Williams, A.C., Johnsingh, A.J.T., Krausman, P.R., 2009.  Population estimation and demography of the Rajaji National Park elephants, Northwest India. Journal of the Bombay Natural History Society 104, 142-152.
Abstract: The Asian Elephant (Elephas maximus) population in Rajaji National Park, north-west India is an important part of India's heritage, but has not been intensively studied until recently. Understanding the population dynamics is important for managers if the population is to remain viable. We used marked adult male Asian Elephants in a mark re-sight method to estimate the male segment of the population and the estimated number of female and associated young using their proportions relative to the adult male segment from classification data. We collected data on inter-calving period and calf survival from adult females present in groups with radio collared females. The number of adult males in the study area was estimated to be 31 (95% CI = 23-41). We computed the relative proportions of other age-sex classes to the adult males and estimated 188 elephants (95% CI = 139-248). Ninety per cent of the adult males had tusks (tuskers) and the adult male to adult female ratio was 1:1.87. This is one of the least skewed sex ratios reported for Asian Elephants and is comparable to areas in Sri Lanka where 95% of males are tuskless. Over 90% of the adult females were accompanied by juveniles or calves <5 years old. We estimated the inter-calving period to be around 4.23 years and the calf survival over the first year was almost 100%. One calf was killed when hit by a train. The high proportion of males, low inter-calving period, and high neonate survival of the Rajaji elephant population indicates that the population is demographically healthy. However, more adult elephants died in train accidents than due to natural causes and viability of this small population could be seriously threatened if losses to train accidents continue.

Wittemyer, G., Okello, J.B., Rasmussen, H.B., Arctander, P., Nyakaana, S., Douglas-Hamilton, I., Siegismund, H.R., 2009. Where sociality and relatedness diverge: the genetic basis for hierarchical social organization in African elephants. Proc Royal Soc Biol 276, 3513-3521.
Abstract: Hierarchical properties characterize elephant fission-fusion social organization whereby stable groups of individuals coalesce into higher order groups or split in a predictable manner. This hierarchical complexity is rare among animals and, as such, an examination of the factors driving its emergence offers unique insight into the evolution of social behaviour. Investigation of the genetic basis for such social affiliation demonstrates that while the majority of core social groups (second-tier affiliates) are significantly related, this is not exclusively the case. As such, direct benefits received through membership of these groups appear to be salient to their formation and maintenance. Further analysis revealed that the majority of groups in the two higher social echelons (third and fourth tiers) are typically not significantly related. The majority of third-tier members are matrilocal, carrying the same mtDNA control region haplotype, while matrilocality among fourth-tier groups was slightly less than expected at random. Comparison of results to those from a less disturbed population suggests that human depredation, leading to social disruption, altered the genetic underpinning of social relations in the study population. These results suggest that inclusive fitness benefits may crystallize elephant hierarchical social structuring along genetic lines when populations are undisturbed. However, indirect benefits are not critical to the formation and maintenance of second-, third- or fourth-tier level bonds, indicating the importance of direct benefits in the emergence of complex, hierarchical social relations among elephants. Future directions and conservation implications are discussed

Wong, K., 2009. Decoding the mammoth
108. Scientific American 300, 26-27.

Asher, R.J., Lehmann, T., 2008. Dental eruption in afrotherian mammals. BMC. Biol. 6, 14.
Abstract: BACKGROUND: Afrotheria comprises a newly recognized clade of mammals with strong molecular evidence for its monophyly. In contrast, morphological data uniting its diverse constituents, including elephants, sea cows, hyraxes, aardvarks, sengis, tenrecs and golden moles, have been difficult to identify. Here, we suggest relatively late eruption of the permanent dentition as a shared characteristic of afrotherian mammals. This characteristic and other features (such as vertebral anomalies and testicondy) recall the phenotype of a human genetic pathology (cleidocranial dysplasia), correlations with which have not been explored previously in the context of character evolution within the recently established phylogeny of living mammalian clades. RESULTS: Although data on the absolute timing of eruption in sengis, golden moles and tenrecs are still unknown, craniometric comparisons for ontogenetic series of these taxa show that considerable skull growth takes place prior to the complete eruption of the permanent cheek teeth. Specimens showing less than half (sengis, golden moles) or two-thirds (tenrecs, hyraxes) of their permanent cheek teeth reach or exceed the median jaw length of conspecifics with a complete dentition. With few exceptions, afrotherians are closer to median adult jaw length with fewer erupted, permanent cheek teeth than comparable stages of non-afrotherians. Manatees (but not dugongs), elephants and hyraxes with known age data show eruption of permanent teeth late in ontogeny relative to other mammals. While the occurrence of delayed eruption, vertebral anomalies and other potential afrotherian synapomorphies resemble some symptoms of a human genetic pathology, these characteristics do not appear to covary significantly among mammalian clades. CONCLUSION: Morphological characteristics shared by such physically disparate animals such as elephants and golden moles are not easy to recognize, but are now known to include late eruption of permanent teeth, in addition to vertebral anomalies, testicondy and other features. Awareness of their possible genetic correlates promises insight into the developmental basis of shared morphological features of afrotherians and other vertebrates

Aupperle, H., Reischauer, A., Bach, F., Hildebrandt, T., Goritz, F., Jager, K., Scheller, R., Klaue, H.J., Schoon, H.A., 2008. Chronic endometritis in an Asian elephant (Elephas maximus). J. Zoo. Wildl. Med. 39, 107-110.
Abstract: A 48-yr-old female Asian elephant with a history of pododermatitis developed recurrent hematuria beginning in 2002. Transrectal ultrasonography and endoscopic examination in 2004 identified the uterus as the source of hematuria and excluded hemorrhagic cystitis. Treatment with Desloreline implants, antibiotics, and homeopathic drugs led to an improved general condition of the elephant. In July 2005, the elephant was suddenly found dead. During necropsy, the severely enlarged uterus contained about 250 L of purulent fluid, and histopathology revealed ulcerative suppurative endometritis with high numbers of Streptococcus equi ssp. zooepidemicus and Escherichia coli identified on aerobic culture. Additional findings at necropsy included: multifocal severe pododermatitis, uterine leiomyoma, and numerous large calcified areas of abdominal fat necrosis. Microbiologic culture of the pododermatitis lesion revealed the presence of Streptococcus agalactiae, Streptococcus equi ssp. zooepidemicus, Staphylococcus sp., Corynebacterium sp., and Entercoccus sp

Gilbert, M.T., Drautz, D.I., Lesk, A.M., Ho, S.Y., Qi, J., Ratan, A., Hsu, C.H., Sher, A., Dalen, L., Gotherstrom, A., Tomsho, L.P., Rendulic, S., Packard, M., Campos, P.F., Kuznetsova, T.V., Shidlovskiy, F., Tikhonov, A., Willerslev, E., Iacumin, P., Buigues, B., Ericson, P.G., Germonpre, M., Kosintsev, P., Nikolaev, V., Nowak-Kemp, M., Knight, J.R., Irzyk, G.P., Perbost, C.S., Fredrikson, K.M., Harkins, T.T., Sheridan, S., Miller, W., Schuster, S.C., 2008. Intraspecific phylogenetic analysis of Siberian woolly mammoths using complete mitochondrial genomes. Proc. Natl. Acad. Sci. U. S. A 105, 8327-8332.
Abstract: We report five new complete mitochondrial DNA (mtDNA) genomes of Siberian woolly mammoth (Mammuthus primigenius), sequenced with up to 73-fold coverage from DNA extracted from hair shaft material. Three of the sequences present the first complete mtDNA genomes of mammoth clade II. Analysis of these and 13 recently published mtDNA genomes demonstrates the existence of two apparently sympatric mtDNA clades that exhibit high interclade divergence. The analytical power afforded by the analysis of the complete mtDNA genomes reveals a surprisingly ancient coalescence age of the two clades, approximately 1-2 million years, depending on the calibration technique. Furthermore, statistical analysis of the temporal distribution of the (14)C ages of these and previously identified members of the two mammoth clades suggests that clade II went extinct before clade I. Modeling of protein structures failed to indicate any important functional difference between genomes belonging to the two clades, suggesting that the loss of clade II more likely is due to genetic drift than a selective sweep

Hofreiter, M., 2008. DNA sequencing: Mammoth genomics. Nature 456, 330-331.

Huang, S., 2008. Ancient fossil specimens of extinct species are genetically more distant to an outgroup than extant sister species are. Riv. Biol. 101, 93-108.
Abstract: There exists a remarkable correlation between genetic distance as measured by protein or DNA dissimilarity and time of species divergence as inferred from fossil records. This observation has provoked the molecular clock hypothesis. However, data inconsistent with the hypothesis have steadily accumulated in recent years from studies of extant organisms. Here the published DNA and protein sequences from ancient fossil specimens were examined to see if they would support the molecular clock hypothesis. The hypothesis predicts that ancient specimens cannot be genetically more distant to an outgroup than extant sister species are. Also, two distinct ancient specimens cannot be genetically more distant than their extant sister species are. The findings here do not conform to these predictions. Neanderthals are more distant to chimpanzees and gorillas than modern humans are. Dinosaurs are more distant to frogs than extant birds are. Mastodons are more distant to opossums than other placental mammals are. The genetic distance between dinosaurs and mastodons is greater than that between extant birds and mammals. Therefore, while the molecular clock hypothesis is consistent with some data from extant organisms, it has yet to find support from ancient fossils. Far more damaging to the hypothesis than data from extant organisms, which merely question the constancy of mutation rate, the study of ancient fossil organisms here challenges for the first time the fundamental premise of modern evolution theory that genetic distances had always increased with time in the past history of life on Earth

Liu, A.G., Seiffert, E.R., Simons, E.L., 2008. Stable isotope evidence for an amphibious phase in early proboscidean evolution. Proc. Natl. Acad. Sci. U. S. A 105, 5786-5791.
Abstract: The order Proboscidea includes extant elephants and their extinct relatives and is closely related to the aquatic sirenians (manatees and dugongs) and terrestrial hyracoids (hyraxes). Some analyses of embryological, morphological, and paleontological data suggest that proboscideans and sirenians shared an aquatic or semiaquatic common ancestor, but independent tests of this hypothesis have proven elusive. Here we test the hypothesis of an aquatic ancestry for advanced proboscideans by measuring delta(18)O in tooth enamel of two late Eocene proboscidean genera, Barytherium and Moeritherium, which are sister taxa of Oligocene-to-Recent proboscideans. The combination of low delta(18)O values and low delta(18)O standard deviations in Barytherium and Moeritherium matches the isotopic pattern seen in aquatic and semiaquatic mammals, and differs from that of terrestrial mammals. delta(13)C values of these early proboscideans suggest that both genera are likely to have consumed freshwater plants, although a component of C(3) terrestrial vegetation cannot be ruled out. The simplest explanation for the combined evidence from isotopes, dental functional morphology, and depositional environments is that Barytherium and Moeritherium were at least semiaquatic and lived in freshwater swamp or riverine environments, where they grazed on freshwater vegetation. These results lend new support to the hypothesis that Oligocene-to-Recent proboscideans are derived from amphibious ancestors

Lotfy, W.M., Brant, S.V., DeJong, R.J., Le, T.H., Demiaszkiewicz, A., Rajapakse, R.P., Perera, V.B., Laursen, J.R., Loker, E.S., 2008. Evolutionary origins, diversification, and biogeography of liver flukes (Digenea, Fasciolidae). American Journal of Tropical Medicine and Hygiene 79, 248-255.
Abstract: Fasciolid flukes are among the largest and best known digenetic trematodes and have considerable historical and veterinary significance. Fasciola hepatica is commonly implicated in causing disease in humans. The origins, patterns of diversification, and biogeography of fasciolids are all poorly known. We have undertaken a molecular phylogenetic study using 28S, internal transcribed spacer 1 and 2 (ITS-1 and ITS-2) of nuclear ribosomal DNA, and mitochondrial nicotinamide dehydrogenase subunit 1 (nad1) that included seven of the nine recognized species in the family. The fasciolids examined comprise a monophyletic group with the most basal species recovered from African elephants. We hypothesize fasciolids migrated from Africa to Eurasia, with secondary colonization of Africa. Fasciolids have been conservative in maintaining relatively large adult body size, but anatomical features of their digestive and reproductive systems are available. These flukes have been opportunistic, with respect to switching to new snail (planorbid to lymnaeid) and mammalian hosts and from intestinal to hepatic habitats within mammals

Lynch, V.J., Tanzer, A., Wang, Y., Leung, F.C., Gellersen, B., Emera, D., Wagner, G.P., 2008. Adaptive changes in the transcription factor HoxA-11 are essential for the evolution of pregnancy in mammals. Proc. Natl. Acad. Sci. U. S. A 105, 14928-14933.
Abstract: Evolutionary change in gene regulation can result from changes in cis-regulatory elements, leading to differences in the temporal and spatial expression of genes or in the coding region of transcription factors leading to novel functions or both. Although there is a growing body of evidence supporting the importance of cis-regulatory evolution, examples of protein-mediated evolution of novel developmental pathways have not been demonstrated. Here, we investigate the evolution of prolactin (PRL) expression in endometrial cells, which is essential for placentation/pregnancy in eutherian mammals and is a direct regulatory target of the transcription factor HoxA-11. Here, we show that (i) endometrial PRL expression is a derived feature of placental mammals, (ii) the PRL regulatory gene HoxA-11 experienced a period of strong positive selection in the stem-lineage of eutherian mammals, and (iii) only HoxA-11 proteins from placental mammals, including the reconstructed ancestral eutherian gene, are able to up-regulate PRL from the promoter used in endometrial cells. In contrast, HoxA-11 from the reconstructed therian ancestor, opossum, platypus, and chicken are unable to up-regulate PRL expression. These results demonstrate that the evolution of novel gene expression domains is not only mediated by the evolution of cis-regulatory elements but can also require evolutionary changes of transcription factor proteins themselves

Makarieva, A.M., Gorshkov, V.G., Li, B.L., Chown, S.L., Reich, P.B., Gavrilov, V.M., 2008. Mean mass-specific metabolic rates are strikingly similar across life's major domains: Evidence for life's metabolic optimum. Proc. Natl. Acad. Sci. U. S. A 105, 16994-16999.
Abstract: A fundamental but unanswered biological question asks how much energy, on average, Earth's different life forms spend per unit mass per unit time to remain alive. Here, using the largest database to date, for 3,006 species that includes most of the range of biological diversity on the planet-from bacteria to elephants, and algae to sapling trees-we show that metabolism displays a striking degree of homeostasis across all of life. We demonstrate that, despite the enormous biochemical, physiological, and ecological differences between the surveyed species that vary over 10(20)-fold in body mass, mean metabolic rates of major taxonomic groups displayed at physiological rest converge on a narrow range from 0.3 to 9 W kg(-1). This 30-fold variation among life's disparate forms represents a remarkably small range compared with the 4,000- to 65,000-fold difference between the mean metabolic rates of the smallest and largest organisms that would be observed if life as a whole conformed to universal quarter-power or third-power allometric scaling laws. The observed broad convergence on a narrow range of basal metabolic rates suggests that organismal designs that fit in this physiological window have been favored by natural selection across all of life's major kingdoms, and that this range might therefore be considered as optimal for living matter as a whole

Meyers, D.A., Isaza, R., MacNeill, A. Evaluation of acute phase proteins for diagnosis of inflammation in Asian elephants ( Elephas maximus). Proc American Associaton of Zoo Veterinarians and Assoc of Reptile and Amphibian Veterinarians.  128. 2008. 11-10-2008.
Ref Type: Conference Proceeding
Abstract: In many domestic species, routine hematology assays are useful diagnostic tools to diagnose inflammatory conditions. Unlike other species, these hematologic tests apparently are insensitive indicators of inflammation in elephants.1 We studied a novel group of blood proteins, called acute phase proteins, which increase during inflammatory conditions, for their usefulness in diagnosing elephants with inflammatory diseases. Although these proteins currently are useful in humans and domestic animals, each species has a different set of important proteins that must be individually investigated.2 We tested several acute phase proteins (C-reactive protein, alpha-1 glycoprotein, alpha-1 antitrypsin, serum amyloid A, haptoglobin, fibrinogen, ceruloplasmin, and albumin) as well as complete blood counts, chemistry panels, serum protein electrophoresis, and 3-D gel electrophoresis to determine their usefulness for diagnosing different types of inflammatory conditions in Asian elephants (Elephas maximus). Animals with inflammatory conditions were classified as those individuals with known illnesses such as mycobacteriosis, arthritis, nail bed abscesses, and malignant tumors. Control animals were thoseanimals that were suspected to not have any inflammation and be healthy at the time of testing as determined by physical examination and obtaining a thorough medical history.
LITERATURE CITED
1. Lyashchenko, K., R. Greenwald, J. Esfandiari, J. Olsen, R. Ball, G. Dumonceaux, F. Dunker, C. Buckley, M.
Richard, S. Murray, J.B. Payeur, P. Anderson, J.M. Pollock, S. Mikota, M. Miller, D. Sofranko, and W.R.
Waters. 2006. Tuberculosis in Elephants: Antibody responses to defined antigens of Mycobacterium
tuberculosis, potential for early diagnosis, and monitoring of treatment. Clin. Vacc. Immunol. 13: 722-732.
2. Murata H., N. Shimada, M. Yoshioka. 2004. Current research on acute phase proteins in veterinary diagnosis:
an overview. Vet J. 168: 28-40.

Miller, W., Drautz, D.I., Ratan, A., Pusey, B., Qi, J., Lesk, A.M., Tomsho, L.P., Packard, M.D., Zhao, F., Sher, A., Tikhonov, A., Raney, B., Patterson, N., Lindblad-Toh, K., Lander, E.S., Knight, J.R., Irzyk, G.P., Fredrikson, K.M., Harkins, T.T., Sheridan, S., Pringle, T., Schuster, S.C., 2008. Sequencing the nuclear genome of the extinct woolly mammoth. Nature 456, 387-390.
Abstract: In 1994, two independent groups extracted DNA from several Pleistocene epoch mammoths and noted differences among individual specimens. Subsequently, DNA sequences have been published for a number of extinct species. However, such ancient DNA is often fragmented and damaged, and studies to date have typically focused on short mitochondrial sequences, never yielding more than a fraction of a per cent of any nuclear genome. Here we describe 4.17 billion bases (Gb) of sequence from several mammoth specimens, 3.3 billion (80%) of which are from the woolly mammoth (Mammuthus primigenius) genome and thus comprise an extensive set of genome-wide sequence from an extinct species. Our data support earlier reports that elephantid genomes exceed 4 Gb. The estimated divergence rate between mammoth and African elephant is half of that between human and chimpanzee. The observed number of nucleotide differences between two particular mammoths was approximately one-eighth of that between one of them and the African elephant, corresponding to a separation between the mammoths of 1.5-2.0 Myr. The estimated probability that orthologous elephant and mammoth amino acids differ is 0.002, corresponding to about one residue per protein. Differences were discovered between mammoth and African elephant in amino-acid positions that are otherwise invariant over several billion years of combined mammalian evolution. This study shows that nuclear genome sequencing of extinct species can reveal population differences not evident from the fossil record, and perhaps even discover genetic factors that affect extinction

Nathan, R., Getz, W.M., Revilla, E., Holyoak, M., Kadmon, R., Saltz, D., Smouse, P.E., 2008. A movement ecology paradigm for unifying organismal movement research. Proc. Natl. Acad. Sci. U. S. A 105, 19052-19059.
Abstract: Movement of individual organisms is fundamental to life, quilting our planet in a rich tapestry of phenomena with diverse implications for ecosystems and humans. Movement research is both plentiful and insightful, and recent methodological advances facilitate obtaining a detailed view of individual movement. Yet, we lack a general unifying paradigm, derived from first principles, which can place movement studies within a common context and advance the development of a mature scientific discipline. This introductory article to the Movement Ecology Special Feature proposes a paradigm that integrates conceptual, theoretical, methodological, and empirical frameworks for studying movement of all organisms, from microbes to trees to elephants. We introduce a conceptual framework depicting the interplay among four basic mechanistic components of organismal movement: the internal state (why move?), motion (how to move?), and navigation (when and where to move?) capacities of the individual and the external factors affecting movement. We demonstrate how the proposed framework aids the study of various taxa and movement types; promotes the formulation of hypotheses about movement; and complements existing biomechanical, cognitive, random, and optimality paradigms of movement. The proposed framework integrates eclectic research on movement into a structured paradigm and aims at providing a basis for hypothesis generation and a vehicle facilitating the understanding of the causes, mechanisms, and spatiotemporal patterns of movement and their role in various ecological and evolutionary processes. "Now we must consider in general the common reason for moving with any movement whatever." (Aristotle, De Motu Animalium, 4th century B.C.)

Nicholls, H., 2008. Darwin 200: Let's make a mammoth. Nature 456, 310-314.

Nishihara, H., Okada, N., 2008. Retroposons: genetic footprints on the evolutionary paths of life. Methods Mol. Biol. 422, 201-225.
Abstract: Retroposons such as short interspersed elements (SINEs) and long interspersed elements are abundant transposable elements in eukaryote genomes. Recent large-scale comparative genome analyses have revealed that retroposons are a major component of genomes, wherein they provide structural diversity between species and uniqueness to each species. SINEs have been used as powerful markers in phylogenetic analyses of various species. This approach, which has been termed the SINE insertion method, infers phylogenetic relationships based on the presence/absence of SINEs among lineages. However, the method is not yet used extensively among biologists, especially molecular phylogenetists, because it is based on an understanding of the molecular mechanisms of retroposition, which may be unfamiliar to many researchers. Moreover, the method may require a large amount of bench work to characterize a new SINE family and to screen genomic libraries of the species of interest. In this chapter, we present the basic theory and detailed technical steps involved in a SINE insertion analysis. Furthermore, we explain the isolation and characterization of a new SINE family from the genome of a species of interest using as an example a known SINE family in mammals

Nogues-Bravo, D., Rodriguez, J., Hortal, J., Batra, P., Araujo, M.B., 2008. Climate change, humans, and the extinction of the woolly mammoth. PLoS. Biol. 6, e79.
Abstract: Woolly mammoths inhabited Eurasia and North America from late Middle Pleistocene (300 ky BP [300,000 years before present]), surviving through different climatic cycles until they vanished in the Holocene (3.6 ky BP). The debate about why the Late Quaternary extinctions occurred has centred upon environmental and human-induced effects, or a combination of both. However, testing these two hypotheses-climatic and anthropogenic-has been hampered by the difficulty of generating quantitative estimates of the relationship between the contraction of the mammoth's geographical range and each of the two hypotheses. We combined climate envelope models and a population model with explicit treatment of woolly mammoth-human interactions to measure the extent to which a combination of climate changes and increased human pressures might have led to the extinction of the species in Eurasia. Climate conditions for woolly mammoths were measured across different time periods: 126 ky BP, 42 ky BP, 30 ky BP, 21 ky BP, and 6 ky BP. We show that suitable climate conditions for the mammoth reduced drastically between the Late Pleistocene and the Holocene, and 90% of its geographical range disappeared between 42 ky BP and 6 ky BP, with the remaining suitable areas in the mid-Holocene being mainly restricted to Arctic Siberia, which is where the latest records of woolly mammoths in continental Asia have been found. Results of the population models also show that the collapse of the climatic niche of the mammoth caused a significant drop in their population size, making woolly mammoths more vulnerable to the increasing hunting pressure from human populations. The coincidence of the disappearance of climatically suitable areas for woolly mammoths and the increase in anthropogenic impacts in the Holocene, the coup de grace, likely set the place and time for the extinction of the woolly mammoth

Okello, J.B., Wittemyer, G., Rasmussen, H.B., Arctander, P., Nyakaana, S., Douglas-Hamilton, I., Siegismund, H.R., 2008. Effective population size dynamics reveal impacts of historic climatic events and recent anthropogenic pressure in African elephants. Mol. Ecol. 17, 3788-3799.
Abstract: Two hundred years of elephant hunting for ivory, peaking in 1970-1980s, caused local extirpations and massive population declines across Africa. The resulting genetic impacts on surviving populations have not been studied, despite the importance of understanding the evolutionary repercussions of such human-mediated events on this keystone species. Using Bayesian coalescent-based genetic methods to evaluate time-specific changes in effective population size, we analysed genetic variation in 20 highly polymorphic microsatellite loci from 400 elephants inhabiting the greater Samburu-Laikipia region of northern Kenya. This area experienced a decline of between 80% and 90% in the last few decades when ivory harvesting was rampant. The most significant change in effective population size, however, occurred approximately 2500 years ago during a mid-Holocene period of climatic drying in tropical Africa. Contrary to expectations, detailed analyses of four contemporary age-based cohorts showed that the peak poaching epidemic in the 1970s caused detectable temporary genetic impacts, with genetic diversity rebounding as juveniles surviving the poaching era became reproductively mature. This study demonstrates the importance of climatic history in shaping the distribution and genetic history of a keystone species and highlights the utility of coalescent-based demographic approaches in unravelling ancestral demographic events despite a lack of ancient samples. Unique insights into the genetic signature of mid-Holocene climatic change in Africa and effects of recent poaching pressure on elephants are discussed

Organ, C.L., Schweitzer, M.H., Zheng, W., Freimark, L.M., Cantley, L.C., Asara, J.M., 2008. Molecular phylogenetics of mastodon and Tyrannosaurus rex. Science 320, 499.
Abstract: We report a molecular phylogeny for a nonavian dinosaur, extending our knowledge of trait evolution within nonavian dinosaurs into the macromolecular level of biological organization. Fragments of collagen alpha1(I) and alpha2(I) proteins extracted from fossil bones of Tyrannosaurus rex and Mammut americanum (mastodon) were analyzed with a variety of phylogenetic methods. Despite missing sequence data, the mastodon groups with elephant and the T. rex groups with birds, consistent with predictions based on genetic and morphological data for mastodon and on morphological data for T. rex. Our findings suggest that molecular data from long-extinct organisms may have the potential for resolving relationships at critical areas in the vertebrate evolutionary tree that have, so far, been phylogenetically intractable

Perelygin, A.A., Zharkikh, A.A., Astakhova, N.M., Lear, T.L., Brinton, M.A., 2008. Concerted evolution of vertebrate CCR2 and CCR5 genes and the origin of a recombinant equine CCR5/2 gene. J. Hered. 99, 500-511.
Abstract: Chemokine receptors (CCRs) play an essential role in the initiation of an innate immune host response. Several of these receptors have been shown to modulate the outcome of viral infections. The recent availability of complete genome sequences from a number of species provides a unique opportunity to analyze the evolution of the CCR genes. A phylogenetic analysis revealed that the CCR2 gene evolved in concert with the paralogous CCR5 gene, but not with another paralogous gene, CCR3, in the opossum, platypus, rabbit, guinea pig, cat, and rodent lineages. In addition, evidence of concerted evolution of the CCR2 and CCR5 genes was observed in chicken and lizard genomes. A unique CCR5/2 gene that originated by unequal crossing over between the CCR2 and CCR5 genes was detected in the domestic horse. The CCR2, CCR5, and CCR5/2 genes were mapped to ECA16q21 using fluorescent in situ hybridization (FISH). Single-nucleotide polymorphisms identified in the equine CCR5 gene and characterized within 5 horse breeds provide haplotype markers for future case/control studies investigating the genetic bases of horse susceptibility to infectious diseases

Tyack, P.L., 2008. Convergence of calls as animals form social bonds, active compensation for noisy communication channels, and the evolution of vocal learning in mammals. J. Comp Psychol. 122, 319-331.
Abstract: The classic evidence for vocal production learning involves imitation of novel, often anthropogenic sounds. Among mammals, this has been reported for dolphins, elephants, harbor seals, and humans. A broader taxonomic distribution has been reported for vocal convergence, where the acoustic properties of calls from different individuals converge when they are housed together in captivity or form social bonds in the wild. Vocal convergence has been demonstrated for animals as diverse as songbirds, parakeets, hummingbirds, bats, elephants, cetaceans, and primates. For most species, call convergence is thought to reflect a group-distinctive identifier, with shared calls reflecting and strengthening social bonds. A ubiquitous function for vocal production learning that is starting to receive attention involves modifying signals to improve communication in a noisy channel. Pooling data on vocal imitation, vocal convergence, and compensation for noise suggests a wider taxonomic distribution of vocal production learning among mammals than has been generally appreciated. The wide taxonomic distribution of this evidence for vocal production learning suggests that perhaps more of the neural underpinnings for vocal production learning are in place in mammals than is usually recognized

Wallis, M., 2008. Mammalian genome projects reveal new growth hormone (GH) sequences. Characterization of the GH-encoding genes of armadillo (Dasypus novemcinctus), hedgehog (Erinaceus europaeus), bat (Myotis lucifugus), hyrax (Procavia capensis), shrew (Sorex araneus), ground squirrel (Spermophilus tridecemlineatus), elephant (Loxodonta africana), cat (Felis catus) and opossum (Monodelphis domestica). Gen. Comp Endocrinol. 155, 271-279.
Abstract: Mammalian growth hormone (GH) sequences have been shown previously to display episodic evolution: the sequence is generally strongly conserved but on at least two occasions during mammalian evolution (on lineages leading to higher primates and ruminants) bursts of rapid evolution occurred. However, the number of mammalian orders studied previously has been relatively limited, and the availability of sequence data via mammalian genome projects provides the potential for extending the range of GH gene sequences examined. Complete or nearly complete GH gene sequences for six mammalian species for which no data were previously available have been extracted from the genome databases-Dasypus novemcinctus (nine-banded armadillo), Erinaceus europaeus (western European hedgehog), Myotis lucifugus (little brown bat), Procavia capensis (cape rock hyrax), Sorex araneus (European shrew), Spermophilus tridecemlineatus (13-lined ground squirrel). In addition incomplete data for several other species have been extended. Examination of the data in detail and comparison with previously available sequences has allowed assessment of the reliability of deduced sequences. Several of the new sequences differ substantially from the consensus sequence previously determined for eutherian GHs, indicating greater variability than previously recognised, and confirming the episodic pattern of evolution. The episodic pattern is not seen for signal sequences, 5' upstream sequence or synonymous substitutions-it is specific to the mature protein sequence, suggesting that it relates to the hormonal function. The substitutions accumulated during the course of GH evolution have occurred mainly on the side of the hormone facing away from the receptor, in a non-random fashion, and it is suggested that this may reflect interaction of the receptor-bound hormone with other proteins or small ligands

Archie, E.A., Hollister-Smith, J.A., Poole, J.H., Lee, P.C., Moss, C.J., Maldonado, J.E., Fleischer, R.C., Alberts, S.C., 2007. Behavioural inbreeding avoidance in wild African elephants. Molecular Ecology 16, 4138-4148.
Abstract: The costs of inbreeding depression, as well as the opportunity costs of inbreeding avoidance, determine whether and which mechanisms of inbreeding avoidance evolve. In African elephants, sex-biased dispersal does not lead to the complete separation of male and female relatives, and so individuals may experience selection to recognize kin and avoid inbreeding. However, because estrous females are rare and male-male competition for mates is intense, the opportunity costs of inbreeding avoidance may be high, particularly for males. Here we combine 28 years of behavioural and demographic data on wild elephants with genotypes from 545 adult females, adult males, and calves in Amboseli National Park, Kenya, to test the hypothesis that elephants engage in sexual behaviour and reproduction with relatives less often than expected by chance. We found support for this hypothesis: males engaged in proportionally fewer sexual behaviours and sired proportionally fewer offspring with females that were natal family members or close genetic relatives (both maternal and paternal) than they did with nonkin. We discuss the relevance of these results for understanding the evolution of inbreeding avoidance and for elephant conservation.

Aroch, I., King, R., Baneth, G., 2007. Hematology and serum biochemistry values of trapped, healthy, free-ranging rock hyraxes (Procavia capensis) and their association with age, sex, and gestational status. Vet. Clin. Pathol. 36, 40-48.
Abstract: BACKGROUND: The rock hyrax (Procavia capensis) is an herbivore prevalent from South Africa to Turkey, and a most common zoo animal. Although many studies of hyrax diseases and physiology are available, clinicopathologic data are limited. OBJECTIVES: The purpose of this study was to establish comprehensive hematologic and biochemical reference intervals for trapped, apparently healthy, free-ranging rock hyraxes using modern laboratory methods and to assess differences related to sex, gestation, and age. METHODS: Blood samples were obtained from 27 healthy, free-ranging hyraxes under anesthesia. Gender, body weight, and gestational status were recorded. Hematologic (n = 25) and serum biochemical (n = 22) analyses were performed using standard automated methodology. Data for male vs female, adult vs juvenile, and pregnant vs nonpregnant female hyraxes were compared using the Mann-Whitney U-test. Associations between variables were assessed using Pearson's or Spearman rank correlation tests. RESULTS: Significant age- and sex-related, but not gestation-related differences were observed in several variables. Serum alkaline phosphatase activity and phosphorus concentration were significantly higher in juveniles compared with adults. A unique type of monocyte comprised 1-3% of leukocytes in 4 hyraxes. Markedly high serum creatine kinase (CK) activity was observed in most hyraxes. CONCLUSIONS: The large number of animals and the availability of sex, age, and gestational data in this study will be useful to zoo and wildlife veterinarians working with rock hyraxes. High serum concentrations of betahydroxybutyric acid in the rock hyrax, compared with dogs, cats, and ruminants, may be related to its unique digestive system. High CK activity may have been the result of a capture myopathy-like syndrome. The unique monocytes in hyraxes resemble those of elephants and are a novel finding in this species

Asara, J.M., Schweitzer, M.H., Freimark, L.M., Phillips, M., Cantley, L.C., 2007. Protein sequences from mastodon and Tyrannosaurus rex revealed by mass spectrometry. Science 316, 280-285.
Abstract: Fossilized bones from extinct taxa harbor the potential for obtaining protein or DNA sequences that could reveal evolutionary links to extant species. We used mass spectrometry to obtain protein sequences from bones of a 160,000- to 600,000-year-old extinct mastodon (Mammut americanum) and a 68-million-year-old dinosaur (Tyrannosaurus rex). The presence of T. rex sequences indicates that their peptide bonds were remarkably stable. Mass spectrometry can thus be used to determine unique sequences from ancient organisms from peptide fragmentation patterns, a valuable tool to study the evolution and adaptation of ancient taxa from which genomic sequences are unlikely to be obtained. Division of Signal Transduction, Beth Israel Deaconess Medical Center, Boston, MA 02115, USA. jasara@bidmc.harvard.edu

Barnes, I., Shapiro, B., Lister, A., Kuznetsova, T., Sher, A., Guthrie, D., Thomas, M.G., 2007. Genetic structure and extinction of the woolly mammoth, Mammuthus primigenius. Curr. Biol. 17, 1072-1075.
Abstract: The interval since circa 50 Ka has been a period of significant species extinctions among the large mammal fauna. However, the relative roles of an increasing human presence and a synchronous series of complex environmental changes in these extinctions have yet to be fully resolved. Recent analyses of fossil material from Beringia have clarified our understanding of the spatiotemporal pattern of Late Pleistocene extinctions, identifying periods of population turnover well before the last glacial maximum (LGM: circa 21 Ka) or subsequent human expansion. To examine the role of pre-LGM population changes in shaping the genetic structure of an extinct species, we analyzed the mitochondrial DNA of woolly mammoths in western Beringia and across its range. We identify genetic signatures of a range expansion of mammoths, from eastern to western Beringia, after the last interglacial (circa 125 Ka), and then an extended period during which demographic inference indicates no population-size increase. The most marked change in diversity at this time is the loss of one of two major mitochondrial lineages

Bates, L.A., Byrne, R.W., 2007. Creative or created: using anecdotes to investigate animal cognition. Methods 42, 12-21.
Abstract: In non-human animals, creative behaviour occurs spontaneously only at low frequencies, so is typically missed by standardised observational methods. Experimental approaches have tended to rely overly on paradigms from child development or adult human cognition, which may be inappropriate for species that inhabit very different perceptual worlds and possess quite different motor capacities than humans. The analysis of anecdotes offers a solution to this impasse, provided certain conditions are met. To be reliable, anecdotes must be recorded immediately after observation, and only the records of scientists experienced with the species and the individuals concerned should be used. Even then, interpretation of a single record is always ambiguous, and analysis is feasible only when collation of multiple records shows that a behaviour pattern occurs repeatedly under similar circumstances. This approach has been used successfully to study a number of creative capacities of animals: the distribution, nature and neural correlates of deception across the primate order; the occurrence of teaching in animals; and the neural correlates of several aptitudes--in birds, foraging innovation, and in primates, innovation, social learning and tool-use. Drawing on these approaches, we describe the use of this method to investigate a new problem, the cognition of the African elephant, a species whose sheer size and evolutionary distance from humans renders the conventional methods of comparative psychology of little use. The aim is both to chart the creative cognitive capacities of this species, and to devise appropriate experimental methods to confirm and extend previous findings

Binladen, J., Gilbert, M.T., Willerslev, E., 2007. 800,000 year old mammoth DNA, modern elephant DNA or PCR artefact? Biol. Lett. 3, 55-56.
Abstract: Poulakakis and colleagues (Poulakakis et al. 2006: Biol. Lett. 2, 451-454), report the recovery of 'authentic' mammoth DNA from an 800,000-year-old fragment of bone excavated on the island of Crete. In light of results from other ancient DNA studies that indicate how DNA survival is unlikely in samples, which are recovered from warm environments and are relatively old (e.g. more than 100,000 years), these findings come as a great surprise. Here, we show that problems exist with the methodological approaches used in the study. First, the nested PCR technique as reported is nonsensical--one of the second round 'nested' primers falls outside the amplicon of the first round PCR. More worryingly, the binding region of one of the first round primers (Elcytb320R) falls within the short 43 base pair reported mammoth sequence, specifically covering two of the three reportedly diagnostic Elephas polymorphisms. Finally, we demonstrate using a simple BLAST search in GenBank that the claimed 'uniquely derived character state' for mammoths is in fact also found within modern elephants

Fritsch, A., Hellmich, C., 2007. 'Universal' microstructural patterns in cortical and trabecular, extracellular and extravascular bone materials: micromechanics-based prediction of anisotropic elasticity
390. Journal of Theoretical Biology 244, 597-620.
Abstract: Bone materials are characterized by an astonishing variability and diversity. Still, because of 'architectural constraints' due to once chosen material constituents and their physical interaction, the fundamental hierarchical organization or basic building plans of bone materials remain largely unchanged during biological evolution. Such universal patterns of microstructural organization govern the mechanical interaction of the elementary components of bone (hydroxyapatite, collagen, water; with directly measurable tissue-independent elastic properties), which are here quantified through a multiscale homogenization scheme delivering effective elastic properties of bone materials: at a scale of 10nm, long cylindrical collagen molecules, attached to each other at their ends by approximately 1.5nm long crosslinks and hosting intermolecular water inbetween, form a contiguous matrix called wet collagen. At a scale of several hundred nanometers, wet collagen and mineral crystal agglomerations interpenetrate each other, forming the mineralized fibril. At a scale of 5-10microm, the extracellular solid bone matrix is represented as collagen fibril inclusions embedded in a foam of largely disordered (extrafibrillar) mineral crystals. At a scale above the ultrastructure, where lacunae are embedded in extracellular bone matrix, the extravascular bone material is observed. Model estimates predicted from tissue-specific composition data gained from a multitude of chemical and physical tests agree remarkably well with corresponding acoustic stiffness experiments across a variety of cortical and trabecular, extracellular and extravascular materials. Besides from reconciling the well-documented, seemingly opposed concepts of 'mineral-reinforced collagen matrix' and 'collagen-reinforced mineral matrix' for bone ultrastructure, this approach opens new possibilities in the exploitation of computer tomographic data for nano-to-macro mechanics of bone organs

Gilbert, M.T., Binladen, J., Miller, W., Wiuf, C., Willerslev, E., Poinar, H., Carlson, J.E., Leebens-Mack, J.H., Schuster, S.C., 2007. Recharacterization of ancient DNA miscoding lesions: insights in the era of sequencing-by-synthesis
422. Nucleic Acids Res. 35, 1-10.
Abstract: Although ancient DNA (aDNA) miscoding lesions have been studied since the earliest days of the field, their nature remains a source of debate. A variety of conflicting hypotheses exist about which miscoding lesions constitute true aDNA damage as opposed to PCR polymerase amplification error. Furthermore, considerable disagreement and speculation exists on which specific damage events underlie observed miscoding lesions. The root of the problem is that it has previously been difficult to assemble sufficient data to test the hypotheses, and near-impossible to accurately determine the specific strand of origin of observed damage events. With the advent of emulsion-based clonal amplification (emPCR) and the sequencing-by-synthesis technology this has changed. In this paper we demonstrate how data produced on the Roche GS20 genome sequencer can determine miscoding lesion strands of origin, and subsequently be interpreted to enable characterization of the aDNA damage behind the observed phenotypes. Through comparative analyses on 390,965 bp of modern chloroplast and 131,474 bp of ancient woolly mammoth GS20 sequence data we conclusively demonstrate that in this sample at least, a permafrost preserved specimen, Type 2 (cytosine-->thymine/guanine-->adenine) miscoding lesions represent the overwhelming majority of damage-derived miscoding lesions. Additionally, we show that an as yet unidentified guanine-->adenine analogue modification, not the conventionally argued cytosine-->uracil deamination, underpins a significant proportion of Type 2 damage. How widespread these implications are for aDNA will become apparent as future studies analyse data recovered from a wider range of substrates

Kellogg, M.E., Burkett, S., Dennis, T.R., Stone, G., Gray, B.A., McGuire, P.M., Zori, R.T., Stanyon, R., 2007. Chromosome painting in the manatee supports Afrotheria and Paenungulata. BMC. Evol. Biol. 7, 6.
Abstract: BACKGROUND: Sirenia (manatees, dugongs and Stellar's sea cow) have no evolutionary relationship with other marine mammals, despite similarities in adaptations and body shape. Recent phylogenomic results place Sirenia in Afrotheria and with elephants and rock hyraxes in Paenungulata. Sirenia and Hyracoidea are the two afrotherian orders as yet unstudied by comparative molecular cytogenetics. Here we report on the chromosome painting of the Florida manatee. RESULTS: The human autosomal and X chromosome paints delimited a total of 44 homologous segments in the manatee genome. The synteny of nine of the 22 human autosomal chromosomes (4, 5, 6, 9, 11, 14, 17, 18 and 20) and the X chromosome were found intact in the manatee. The syntenies of other human chromosomes were disrupted in the manatee genome into two to five segments. The hybridization pattern revealed that 20 (15 unique) associations of human chromosome segments are found in the manatee genome: 1/15, 1/19, 2/3 (twice), 3/7 (twice), 3/13, 3/21, 5/21, 7/16, 8/22, 10/12 (twice), 11/20, 12/22 (three times), 14/15, 16/19 and 18/19. CONCLUSION: There are five derived chromosome traits that strongly link elephants with manatees in Tethytheria and give implicit support to Paenungulata: the associations 2/3, 3/13, 8/22, 18/19 and the loss of the ancestral eutherian 4/8 association. It would be useful to test these conclusions with chromosome painting in hyraxes. The manatee chromosome painting data confirm that the associations 1/19 and 5/21 phylogenetically link afrotherian species and show that Afrotheria is a natural clade. The association 10/12/22 is also ubiquitous in Afrotheria (clade I), present in Laurasiatheria (clade IV), only partially present in Xenarthra (10/12, clade II) and absent in Euarchontoglires (clade III). If Afrotheria is basal to eutherians, this association could be part of the ancestral eutherian karyotype. If afrotherians are not at the root of the eutherian tree, then the 10/12/22 association could be one of a suite of derived associations linking afrotherian taxa

Kerr, R.A., 2007. Paleontology. Mammoth-killer impact gets mixed reception from Earth scientists. Science 316, 1264-1265.

Kullberg, M., Hallström, B., Arnason, U., Janke, A., 2007. Expressed sequence tags as a tool for phylogenetic analysis of placental mammal evolution. PLoS ONE E publication Aug 22;2(1):e775.
Abstract: BACKGROUND: We investigate the usefulness of expressed sequence tags, ESTs, for establishing divergences within the tree of placental mammals. This is done on the example of the established relationships among primates (human), lagomorphs (rabbit), rodents (rat and mouse), artiodactyls (cow), carnivorans (dog) and proboscideans (elephant). METHODOLOGY/PRINCIPAL FINDINGS: We have produced 2000 ESTs (1.2 mega bases) from a marsupial mouse and characterized the data for their use in phylogenetic analysis. The sequences were used to identify putative orthologous sequences from whole genome projects. Although most ESTs stem from single sequence reads, the frequency of potential sequencing errors was found to be lower than allelic variation. Most of the sequences represented slowly evolving housekeeping-type genes, with an average amino acid distance of 6.6% between human and mouse. Positive Darwinian selection was identified at only a few single sites. Phylogenetic analyses of the EST data yielded trees that were consistent with those established from whole genome projects. CONCLUSIONS: The general quality of EST sequences and the general absence of positive selection in these sequences make ESTs an attractive tool for phylogenetic analysis. The EST approach allows, at reasonable costs, a fast extension of data sampling from species outside the genome projects.

Murphy, W.J., Pringle, T.H., Crider, T.A., Springer, M.S., Miller, W., 2007. Using genomic data to unravel the root of the placental mammal phylogeny. Genome Res. 17, 413-421.
Abstract: The phylogeny of placental mammals is a critical framework for choosing future genome sequencing targets and for resolving the ancestral mammalian genome at the nucleotide level. Despite considerable recent progress defining superordinal relationships, several branches remain poorly resolved, including the root of the placental tree. Here we analyzed the genome sequence assemblies of human, armadillo, elephant, and opossum to identify informative coding indels that would serve as rare genomic changes to infer early events in placental mammal phylogeny. We also expanded our species sampling by including sequence data from >30 ongoing genome projects, followed by PCR and sequencing validation of each indel in additional taxa. Our data provide support for a sister-group relationship between Afrotheria and Xenarthra (the Atlantogenata hypothesis), which is in turn the sister-taxon to Boreoeutheria. We failed to recover any indels in support of a basal position for Xenarthra (Epitheria), which is suggested by morphology and a recent retroposon analysis, or a hypothesis with Afrotheria basal (Exafricoplacentalia), which is favored by phylogenetic analysis of large nuclear gene data sets. In addition, we identified two retroposon insertions that also support Atlantogenata and none for the alternative hypotheses. A revised molecular timescale based on these phylogenetic inferences suggests Afrotheria and Xenarthra diverged from other placental mammals approximately 103 (95-114) million years ago. We discuss the impacts of this topology on earlier phylogenetic reconstructions and repeat-based inferences of phylogeny

Orlando, L., Pages, M., Calvignac, S., Hughes, S., Hanni, C., 2007. Does the 43 bp sequence from an 800,000 year old Cretan dwarf elephantid really rewrite the textbook on mammoths? Biology Letters 3, 57-59.
Abstract: Pigmy elephants inhabited the islands from the Mediterranean region during the Pleistocene period but became extinct in the course of the Holocene. Despite striking distinctive anatomical characteristics related to insularity, some similarities with the lineage of extant Asian elephants have suggested that pigmy elephants could be most probably seen as members of the genus Elephas. Poulakakis et al. (2006) have recently challenged this view by recovering a short mtDNA sequence from an 800 000 year old fossil of the Cretan pigmy elephant (Elephas creticus). According to the authors of this study, a deep taxonomic revision of Cretan dwarf elephants would be needed, as the sequence exhibits clear affinities with woolly mammoth haplotypes. However, we point here many aspects that seriously weaken the strength of the ancient DNA evidence reported.

Orlando, L., Pages, M., Calvignac, S., Hughes, S., Hanni, C., 2007. Does the 43 bp sequence from an 800,000 year old cretan dwarf elephantid really rewrite the textbook on mammoths? Biol. Lett. 3, 57-59.
Abstract: Pigmy elephants inhabited the islands from the Mediterranean region during the Pleistocene period but became extinct in the course of the Holocene. Despite striking distinctive anatomical characteristics related to insularity, some similarities with the lineage of extant Asian elephants have suggested that pigmy elephants could be most probably seen as members of the genus Elephas. Poulakakis et al (2006) have recently challenged this view by recovering a short mtDNA sequence from an 800 000 year old fossil of the Cretan pigmy elephant (Elephas creticus). According to the authors of this study, a deep taxonomic revision of Cretan dwarf elephants would be needed, as the sequence exhibits clear affinities with woolly mammoth haplotypes. However, we point here many aspects that seriously weaken the strength of the ancient DNA evidence reported

Orlando, L., Hanni, C., Douady, C.J., 2007. Mammoth and Elephant Phylogenetic Relationships: Mammut Americanum, the Missing Outgroup. Evol. Bioinform. Online. 3, 45-51.
Abstract: At the morphological level, the woolly mammoth has most often been considered as the sister-species of Asian elephants, but at the DNA level, different studies have found support for proximity with African elephants. Recent reports have increased the available sequence data and apparently solved the discrepancy, finding mammoths to be most closely related to Asian elephants. However, we demonstrate here that the three competing topologies have similar likelihood, bayesian and parsimony supports. The analysis further suggests the inadequacy of using Sirenia or Hyracoidea as outgroups. We therefore argue that orthologous sequences from the extinct American mastodon will be required to definitively solve this long-standing question

Pan, D., 2007. Hippo signaling in organ size control. Genes Dev. 21, 886-897.
Abstract: The control of organ (or organism) size is a fundamental aspect of life that has long captured human imagination. What makes an elephant grow a million times larger than a mouse? How do our two hands develop independently of each other yet reach very similar size? How does a liver precisely regenerate its original mass when two-thirds of it is removed? The recent discovery of a novel signaling network in Drosophila, known as the Hippo (Hpo) pathway, might provide an important entry point to these fascinating questions. The Hpo pathway consists of several negative growth regulators acting in a kinase cascade that ultimately phosphorylates and inactivates Yorkie (Yki), a transcriptional coactivator that positively regulates cell growth, survival, and proliferation. Components of the Hpo pathway are highly conserved throughout evolution, suggesting that this pathway may function as a global regulator of tissue homeostasis in all metazoan animals. Here, I provide a historical review of this potent growth-regulatory pathway and highlight outstanding questions that will likely be the focus of future investigation

Pardini, A.T., O'Brien, P.C., Fu, B., Bonde, R.K., Elder, F.F., Ferguson-Smith, M.A., Yang, F., Robinson, T.J., 2007. Chromosome painting among Proboscidea, Hyracoidea and Sirenia: support for Paenungulata (Afrotheria, Mammalia) but not Tethytheria. Proc. Biol. Sci. 274, 1333-1340.
Abstract: Despite marked improvements in the interpretation of systematic relationships within Eutheria, particular nodes, including Paenungulata (Hyracoidea, Sirenia and Proboscidea), remain ambiguous. The combination of a rapid radiation, a deep divergence and an extensive morphological diversification has resulted in a limited phylogenetic signal confounding resolution within this clade both at the morphological and nucleotide levels. Cross-species chromosome painting was used to delineate regions of homology between Loxodonta africana (2n=56), Procavia capensis (2n=54), Trichechus manatus latirostris (2n=48) and an outgroup taxon, the aardvark (Orycteropus afer, 2n=20). Changes specific to each lineage were identified and although the presence of a minimum of 11 synapomorphies confirmed the monophyly of Paenungulata, no change characterizing intrapaenungulate relationships was evident. The reconstruction of an ancestral paenungulate karyotype and the estimation of rates of chromosomal evolution indicate a reduced rate of genomic repatterning following the paenungulate radiation. In comparison to data available for other mammalian taxa, the paenungulate rate of chromosomal evolution is slow to moderate. As a consequence, the absence of a chromosomal character uniting two paenungulates (at the level of resolution characterized in this study) may be due to a reduced rate of chromosomal change relative to the length of time separating successive divergence events

Redi, C.A., Garagna, S., Zuccotti, M., Capanna, E., 2007. Genome size: a novel genomic signature in support of Afrotheria. Journal of Molecular Evolution 64, 484-487.
Abstract: Molecular phylogenetic analyses suggest an emerging phylogeny for the extant Placentalia (eutherian) that radically departs from morphologically based constructions of the past. Placental mammals are partitioned into four supraordinal clades: Afrotheria, Xenarthra, Laurasiatheria, and Euarchontoglires. Afrotheria form an endemic African clade that includes elephant shrews, golden moles, tenrecs, aardvarks, hyraxes, elephants, dugongs, and manatees. Datamining databases of genome size (GS) shows that till today just one afrotherian GS has been evaluated, that of the aardvark Orycteropus afer. We show that the GSs of six selected representatives across the Afrotheria supraordinal group are among the highest for the extant Placentalia, providing a novel genomic signature of this enigmatic group. The mean GS value of Afrotheria, 5.3 +/- 0.7 pg, is the highest reported for the extant Placentalia. This should assist in planning new genome sequencing initiatives

Roca, A.L., Georgiadis, N., O'Brien, S.J., 2007. Cyto-nuclear genomic dissociation and the African elephant species question. Quat. Int. 169-170, 4-16.
Abstract: Studies of skull morphology and of nuclear DNA have strongly concluded that African elephants comprise two species. Nonetheless, Debruyne (2005) has suggested a single-species model for Loxodonta based on the polyphyly of a single genetic locus, mitochondrial DNA (mtDNA). Discordant patterns between mitochondrial and nuclear DNA markers were subsequently reported in some African savanna elephant populations, further supporting a two-species model, and prompting us to re-examine here the geographic distribution of different elephant morphotypes and their relationship to nuclear and mtDNA phylogeographic patterns. We used exact tests to compare the distribution of forest elephant-typical and savanna elephant-typical characteristics across eight published datasets containing morphological, mtDNA or nuclear DNA data for African elephants. Among the elephants examined by Debruyne (2005), we found that patterns of forest vs. savanna characteristics were significantly different (p < 10(-5)) between mtDNA and morphology, suggesting the presence of cyto-nuclear genomic dissociation. We show that the eight African elephant continent-wide datasets compared, including that of Debruyne (2005), together support a two-species model with cyto-nuclear genomic dissociation rather than a one-species model, and together indicate that Africa harbors two species of elephant

Rohland, N., Malaspinas, A.S., Pollack, J.L., Slatkin, M., Matheus, P., 2007. Proboscidean mitogenomics: Chronology and mode of elephant evolution using mastodon as outgroup. PLoS Biol 5,  e207. doi:10.1371/journal.pbio.0050207.
Abstract: We have sequenced the complete mitochondrial genome of the extinct American mastodon (Mammut americanum) from an Alaskan fossil that is between 50,000 and 130,000 y old, extending the age range of genomic analyses by almost a complete glacial cycle. The sequence we obtained is substantially different from previously reported partial mastodon mitochondrial DNA sequences. By comparing those partial sequences to other proboscidean sequences, we conclude that we have obtained the first sequence of mastodon DNA ever reported. Using the sequence of the mastodon, which diverged 24-28 million years ago (mya) from the Elephantidae lineage, as an outgroup, we infer that the ancestors of African elephants diverged from the lineage leading to mammoths and Asian elephants approximately 7.6 mya and that mammoths and Asian elephants diverged approximately 6.7 mya. We also conclude
that the nuclear genomes of the African savannah and forest elephants diverged approximately 4.0 mya, supporting the view that these two groups represent different species. Finally, we found the mitochondrial mutation rate of proboscideans to be roughly half of the rate in primates during at least the last 24 million years.

Sreekumar, E., Janki, M.B., Arathy, D.S., Hariharan, R., Premraj, C.A., Rasool, T.J., 2007. Molecular characterization and expression of interferon-gamma of Asian elephant (Elephas maximus). Vet. Immunol. Immunopathol. 118, 75-83.
Abstract: Tuberculosis (TB) caused by Mycobacterial organisms has emerged as one of the major diseases in captive elephants. In vitro Interferon-gamma (IFN-gamma) assay is being used as an ancillary test for early detection of TB in domestic and captive wild animals. In the present study, basic sequence information and immunological cross-reactivity of this major cytokine of Asian elephants were explored. At predicted amino acid level, IFN-gamma of Asian elephant showed maximum identity to that of horse (73%). Other IFN-gamma amino acid sequences that showed high level identity were that of giant panda (72%), dog (71%), nine-banded armadillo (69%), cattle (63%) and human (62%). IFN-gamma promoter sequences of Asian elephant, human, cattle and mouse showed high level conservation of the putative transcription factor binding sites, TATA box and transcriptional start site. The functionally important human IFN-gamma promoter elements, such as AP-2IRE-BE, YY1-gammaIFN-BED, ATFCS and AP-1gammaINF binding sites, were absolutely conserved in the corresponding elephant sequence. There was only a single nucleotide variation in the other two important elements, NFAT-gammaINF and IFN-gammaPE, indicating the highly conserved regulation of IFN-gamma expression across different species. Phylogenetic analysis based on IFN-gamma protein sequences revealed a closer relation of Asian elephants and nine-banded armadillo. This shows a closer evolution of these members of Afrotheria and Xenarthra, respectively; and supports the previous reports based on mitochondrial DNA studies. In Western blot analysis, IFN-gamma of Asian elephant expressed in Escherichia coli was detected using an anti-bovine IFN-gamma monoclonal antibody, indicating immunological cross-reactivity

Tabuce, R., Marivaux, L., Adaci, M., Bensalah, M., Hartenberger, J.L., Mahboubi, M., Mebrouk, F., Tafforeau, P., Jaeger, J.J., 2007. Early Tertiary mammals from North Africa reinforce the molecular Afrotheria clade. Proc. Biol. Sci. 274, 1159-1166.
Abstract: The phylogenetic pattern and timing of the radiation of mammals, especially the geographical origins of major crown clades, are areas of controversy among molecular biologists, morphologists and palaeontologists. Molecular phylogeneticists have identified an Afrotheria clade, which includes several taxa as different as tenrecs (Tenrecidae), golden moles (Chrysochloridae), elephant-shrews (Macroscelididae), aardvarks (Tubulidentata) and paenungulates (elephants, sea cows and hyracoids). Molecular data also suggest a Cretaceous African origin for Afrotheria within Placentalia followed by a long period of endemic evolution on the Afro-Arabian continent after the mid-Cretaceous Gondwanan breakup (approx. 105-25 Myr ago). However, there was no morphological support for such a natural grouping so far. Here, we report new dental and postcranial evidence of Eocene stem hyrax and macroscelidid from North Africa that, for the first time, provides a congruent phylogenetic view with the molecular Afrotheria clade. These new fossils imply, however, substantial changes regarding the historical biogeography of afrotheres. Their long period of isolation in Africa, as assumed by molecular inferences, is now to be reconsidered inasmuch as Eocene paenungulates and elephant-shrews are here found to be related to some Early Tertiary Euramerican 'hyopsodontid condylarths' (archaic hoofed mammals). As a result, stem members of afrotherian clades are not strictly African but also include some Early Paleogene Holarctic mammals

Weiss, B., Faus, H., Haendler, B., 2007. Phylogenetic conservation of the androgen receptor AR45 variant form in placental mammals. Gene 399, 105-111.
Abstract: A cDNA coding for a tissue-specific AR45 variant form of the androgen receptor (AR) has recently been identified in humans, with highest expression levels found in heart. The deduced protein comprises the DNA-binding domain, hinge region and ligand-binding domain of the AR, but not the N-terminal domain which is replaced by a unique, short, seven amino-acid-long stretch. This sequence is encoded by the mutually exclusive exon 1B, located between exons 1 and 2 of the human AR gene. As transcript variants of the steroid receptor family have been shown to have important implications for hormone function, we set out to analyse the genomes of different organisms for potential AR45 expression. We found exon 1B to be conserved in the syntenic chromosomal region of non-human primates such as the chimpanzee Pan troglodytes, the orang-utan Pongo pygmaeus, the macaque Macaca mulatta and the marmoset Callithrix jacchus, and of the elephant Loxondonta africana, the pig Sus scrofa and the dog Canis familiaris. Quantification of AR45 transcript levels in heart, skeletal muscle and lung of Macaca fascicularis showed the heart to be the main organ of expression. A complete AR45 cDNA was furthermore isolated from the heart of this species. Comparative analysis of the identified AR45 exon 1B regions and of the deduced amino acids revealed a high conservation among species. The four N-terminal residues were identical in all eight species, whereas a few changes were seen in the other three residues in the marmoset, elephant and pig. In contrast, we observed more divergence in the mouse Mus musculus and rat Rattus norvegicus syntenic regions. Here a stop codon was found downstream of the potential start codon in the putatively deduced protein sequence and it can be inferred that no protein corresponding to AR45 exists in these two species. The existence of AR45 in different placental mammals with the exception of mouse and rat suggests a disappearance in rodents late in evolution, before the separation of the mouse and rat lineages, about 16 million years ago. In view of the potential function of AR45 as a regulator of AR function, and considering the multiple roles of androgens in normal physiology and in several diseases, these findings have important implications with regard to subtle differences in the action of the male sexual hormone in various organisms

Wellehan, J.F., Johnson, A.J., Childress, A.L., Harr, K.E., Isaza, R., 2007. Six novel gammaherpesviruses of Afrotheria provide insight into the early divergence of the Gammaherpesvirinae. Vet Microbiol 2007 Aug 19; [Epub ahead of print].
Abstract: The Afrotheria represent an early branching of placental mammals. Only two herpesviruses from Afrotheria have been previously identified, and the genus Proboscivirus in the subfamily Betaherpesvirinae has been proposed for them. Six novel gammaherpesviruses were identified in four species in the superorder Afrotheria by detection and analysis of their DNA polymerase genes. Elephantid herpesvirus 3 (ElHV3) and Elephantid herpesvirus 4 (ElHV4) were identified from conjunctival swabs from Asian elephants (Elephas maximus). ElHV3 was also found
in a vaginal swab from one elephant with vaginitis. Elephantid herpesvirus 5 (ElHV5) was identified from vaginal swabs of two Asian elephants with vaginal plaques. Elephantid herpesvirus 6 was discovered in a conjunctival swab from an African elephant (Loxodonta africana). Procavid herpesvirus 1 (PrHV1) was found in spleen and conjunctival swabs of rock hyrax (Procavia capensis). Trichechid herpesvirus 1 (TrHV1) was identified from skin and buffy coats of Florida manatees (Trichechus manatus latirostris). ElHV3 and ElHV4 form a distinct cluster, and ElHV5, ElHV6, TrHV1, and PrHV1 form a second cluster. These viruses may have codiverged with their host species. Phylogenetic analysis of these novel herpesviruses suggests that two separate groups of gammaherpesviruses may have codiverged with the Afrotheria.

Archie, E.A., Moss, C.J., Alberts, S.C., 2006. The ties that bind: genetic relatedness predicts the fission and fusion of social groups in wild African elephants
490. Proc. Biol. Sci. 273, 513-522.
Abstract: Many social animals live in stable groups. In contrast, African savannah elephants (Loxodonta africana) live in unusually fluid, fission-fusion societies. That is, 'core' social groups are composed of predictable sets of individuals; however, over the course of hours or days, these groups may temporarily divide and reunite, or they may fuse with other social groups to form much larger social units. Here, we test the hypothesis that genetic relatedness predicts patterns of group fission and fusion among wild, female African elephants. Our study of a single Kenyan population spans 236 individuals in 45 core social groups, genotyped at 11 microsatellite and one mitochondrial DNA (mtDNA) locus. We found that genetic relatedness predicted group fission; adult females remained with their first order maternal relatives when core groups fissioned temporarily. Relatedness also predicted temporary fusion between social groups; core groups were more likely to fuse with each other when the oldest females in each group were genetic relatives. Groups that shared mtDNA haplotypes were also significantly more likely to fuse than groups that did not share mtDNA. Our results suggest that associations between core social groups persist for decades after the original maternal kin have died. We discuss these results in the context of kin selection and its possible role in the evolution of elephant sociality

Bojesen, A.M., Olsen, K.E., Bertelsen, M.F., 2006. Fatal enterocolitis in Asian elephants (Elephas maximus) caused by Clostridium difficile
456. Vet. Microbiol. 116, 329-335.
Abstract: Two cases of fatal enteritis caused by Clostridium difficile in captive Asian elephants are reported from an outbreak affecting five females in the same zoo. Post mortem examination including histopathology demonstrated fibrinonecrotic enterocolitis. C. difficile was isolated by selective cultivation from two dead and a third severely affected elephant. Four isolates were obtained and found positive for toxin A and B by PCR. All isolates were positive in a toxigenic culture assay and toxin was demonstrated in the intestinal content from one of the fatal cases and in a surviving but severely affected elephant. PCR ribotyping demonstrated that the C. difficile isolates shared an identical profile, which was different from an epidemiologically unrelated strain, indicating that the outbreak was caused by the same C. difficile clone. It is speculated that the feeding of large quantities of broccoli, a rich source of sulforaphane, which has been shown to inhibit the growth of many intestinal microorganisms may have triggered a subsequent overgrowth by C. difficile. This is the first report of C. difficile as the main cause of fatal enterocolitis in elephants. The findings emphasize the need to regard this organism as potentially dangerous for elephants and caution is recommended concerning antibiotic treatment and feeding with diets containing antimicrobials, which may trigger an expansion of a C. difficile population in the gut

Capelli, C., MacPhee, R.D., Roca, A.L., Brisighelli, F., Georgiadis, N., O'Brien, S.J., Greenwood, A.D., 2006. A nuclear DNA phylogeny of the woolly mammoth (Mammuthus primigenius)
468. Mol. Phylogenet. Evol. 40, 620-627.

Cooper, A., 2006. The year of the mammoth. PLoS Biology 4, 1-3.
Abstract: Mammoth mitochondrial (mt) genomes are apparently on a similar schedule to London buses-you wait for ages and then suddenly three come along at once. Within the past six weeks, three studies [1-3] have independently determined all, or most, of the mammoth mt genome sequence, some 16,800 base pairs (bp). Encouragingly, the partial sequence was a byproduct of a study that generated some 13 million bp of mammoth genomic DNA using a new, massively parallel sequencing approach. The very divergent methods used in these three studies also neatly represent the past, present, and future of ancient DNA (aDNA) research. aDNA methods provide an opportunity to characterise the genetic composition of species and populations in the past, and to actually observe evolutionary change through real time. Such a record has great potential to reveal the processes that have generated the diversity and distribution of taxa in our modern environment, and to examine phenomena such as speciation, domestication, morphological evolution, and the impacts of major environmental changes. aDNA data also provide an important opportunity to test our ability to accurately reconstruct evolutionary history via the fossil record or via extrapolation from the genetic data of modern species. Unfortunately, the potential of aDNA remains largely untapped because research has been severely limited by the technical diffi culties of retrieving and studying the trace amounts of  highly fragmented DNA that survive in ancient specimens.

Cooper, D.W., Larsen, E., 2006. Immunocontraception of mammalian wildlife: Ecological and immunogenetic issues. Reproduction 132, 821-828.
Abstract: Immunocontraception involves stimulating immune responses against gametes or reproductive hormones thus preventing conception. The method is being developed for the humane control of pest and overabundant populations of mammalian wildlife. This paper examines three fundamental issues associated with its use: (1) the difficulties of obtaining responses to self-antigens, (2) the likely evolution of genetically based non-response to immunocontraceptive agents, and (3) the possible changes in the array of pathogens possessed by the target species after generations of immunocontraception. Our review of the literature demonstrates that the barriers to an effective immunocontraceptive are at present very basic. Should they be overcome, the effects of immunocontraception on the immunogenetic constitution of wildlife populations through the selection for nonresponders must be examined. We suggest that the attempt to use the animal's own immune system to modulate reproduction may be incompatible with the basic biological function of protection against infectious disease. Research programs on mammalian immunocontraception should involve measurement of the heritability of non-response and an assessment of the likely change in the response of the contracepted population to possible pathogens.

Ducummon, C.C., Berger, T., 2006. Localization of the Rho GTPases and some Rho effector proteins in the sperm of several mammalian species
440. Zygote. 14, 249-257.
Abstract: The acrosome reaction is a fundamental event in the biology of the sperm and is a prerequisite to fertilization of the egg. Members of the Rho family of GTPases and their effectors are present in the cytoplasm and/or plasma membrane overlying the acrosome of porcine sperm. We have implicated the Rho family of GTPases and the Rho-activated kinase, ROCK-1, in mediating the zona-pellucida-induced acrosome reaction. Others have implicated the Rho GTPase in regulating the ionophore-induced acrosome reaction in the sperm of several mammalian species as well as in motility of bovine sperm. In this study, the localization of the Rho GTPases (RhoA, RhoB, Rac1 and Cdc42) as well as the effectors RhoGDI, PI(4)P5K and ROCK-1, was determined in boar, human, rat, ram, bull and elephant sperm. The four GTPases were each present in the sperm head of all species examined. RhoGDI was expressed in the head and tail of sperm from all species except pig, where it was present only in the head. PI(4)P5K was expressed in both head and tail of sperm from all species, but expression was typically weaker in the tail. Finally, ROCK-1 was expressed in the heads and tails of all sperm except that of the boar, where it was present only in the acrosomal region. These observations taken together suggest that the expression of Rho GTPases in sperm has been conserved throughout mammalian evolution, most likely due to the role of these GTPases in regulating acrosomal exocytosis

Eckhart, L., Uthman, A., Sipos, W., Tschachler, E., 2006. Genome sequence comparison reveals independent inactivation of the caspase-15 gene in different evolutionary lineages of mammals
428. Molecular Biology and Evolution 23, 2081-2089.
Abstract: We have recently demonstrated that placental mammalian species such as pig and dog express a novel proapoptotic protease, caspase-15, whereas mouse and humans lack this enzyme. Here we investigated the evolutionary fate of the caspase-15 gene in different mammalian lineages by analyzing whole-genome shotgun sequences of 30 mammalian species for the presence of caspase-15 orthologs. Caspase-15 gene sequences were found in representatives of all major mammalian clades except for the superorders Afrotheria (tenrec, rock hyrax, and elephant) and Euarchontoglires (rodents, rabbit, tree shrew, and primates), which either lacked any caspase-15-like sequences or contained mutated remnants of the caspase-15 gene. Polymerase chain reaction screenings confirmed the results of the database searches and showed that the caspase-15 gene is expressed not only in various placental mammals but also in the marsupial, Monodelphis domestica. The observed species distribution implies that caspase-15 has originated in an early ancestor of modern mammals and has been conserved, over more than 180 Myr, in marsupials and many placental mammals, whereas it was independently lost in 2 phylogenetically distant clades of placental mammals, that is, Afrotheria and Euarchontoglires. Our data suggest that the inactivation of the caspase-15 gene was not counteracted by, and may even have been driven by, evolutionary constraints in these clades, and therefore, caution against the uncritical use of gene absence for the inference of phylogenetic relationships

Fischer, M.S., Blickhan, R., 2006. The tri-segmented limbs of therian mammals: kinematics, dynamics, and self-stabilization--a review
397. J. Exp. Zool. A Comp Exp. Biol. 305, 935-952.
Abstract: The evolution of therian mammals is to a large degree marked by changes in their motion systems. One of the decisive transitions has been from the sprawled, bi-segmented to the parasagittal, tri-segmented limb. Here, we review aspects of the tri-segmented limb in locomotion which have been elucidated in our research groups in the last 10 years. First, we report the kinematics of the tri-segmented therian limb from mouse to elephant in order to explore general principles of the therian limb configuration and locomotion. Torques will be reported from a previous paper (Witte et al., 2002. J Exp Biol 205:1339-1353) for a better understanding of the anti-gravity work of all limb joints. The stability of a limb in z-configuration will be explained and its advantage with respect to other potential solutions from modeling will be discussed. Finally, we describe how the emerging concept of self-stability can be explained for a tri-segmented leg template and how it affects the design of the musculoskeletal system and the operation of legs during locomotion. While locomotion has been considered as mainly a control problem in various disciplines, we stress the necessity to reduce control as much as possible. Central control can be cheap if the limbs are "intelligent" by means of their design. Gravity-induced movements and self-stability seem to be energy-saving mechanisms

Gee, H., 2006. Evolution: memories of mammoths
505. Nature 439, 673.

Gelvin-Reymiller, C., Reuther, J.D., Potter, B.A., Bowers, P.M., 2006. Technical aspects of a worked proboscidean tusk from Inmachuk River, Seward Peninsula, Alaska. Journal of Archaeological Science 33, 1088-1094.
Abstract: Prehistoric reduction sequences of proboscidean ivory have been described and discussed within the Russian and European Upper Paleolithic archaeological literature. A culturally modified proboscidean tusk (Mammuthus sp.) in Seward Peninsula, northwestern Alaska, displays longitudinal grooving, providing an insight into a reduction technique rarely described within North American archaeological literature. Similar reduction sequences have been described for the production of bone, antler and walrus ivory artifacts in the North American prehistoric record; however, examples on proboscidean ivory are extremely rare.

Henderson, D.M., 2006. Burly gaits: Centers of mass, stability, and the trackways of sauropod dinosaurs. Journal of Vertebrate Paleontology 26, 907-921.
Abstract: The narrow- and wide-gauge trackways attributed to sauropod dinosaurs are hypothesized to be a consequence of the relative positions of their centers of mass. This hypothesis was tested using three-dimensional, trackwayproducing computer models of two sauropods and studies of Asian elephants. Centers of mass of sauropod models were computed using density distributions that reflect the high degree of pneumatization of the skeletons and air sacs within the body. A close correspondence was found between the relative areas of hand and foot prints in different trackways and the relative fractions of the body weight borne by the forefeet and hindfeet in the different types of sauropods inferred to have made the trackways. Experimental studies of Asian elephants corroborated the close correspondence between relative areas of the hindfeet and forefeet and body weight distribution. Replicating actual sauropod trackways with the walking models enabled testing of proposed gaits for a sauropod model. Brachiosaurus brancai, with its more centrally positioned center of mass, was stable and possessed a wide safety margin only when replicating a wide trackway. Conversely, Diplodocics carnegii, with a more posteriorly placed center of mass, was most stable when replicating a narrow trackway. A trend for large sauropods (> 12 tons), independent of clade, to have more anteriorly positioned centers of mass was identified, and it is proposed that all large sauropods were restricted to producing wide-gauge trackways for stability reasons. The primitive gait state for Sauropodomorpha was determined to be one that produced narrow-gauge trackways.

Hilborn, R., Arcese, P., Borner, M., Hando, J., Hopcraft, G., Loibooki, M., Mduma, S., Sinclair, A.R., 2006. Effective enforcement in a conservation area
378. Science 314, 1266.
Abstract: Wildlife within protected areas is under increasing threat from bushmeat and illegal trophy trades, and many argue that enforcement within protected areas is not sufficient to protect wildlife. We examined 50 years of records from Serengeti National Park in Tanzania and calculated the history of illegal harvest and enforcement by park authorities. We show that a precipitous decline in enforcement in 1977 resulted in a large increase in poaching and decline of many species. Conversely, expanded budgets and antipoaching patrols since the mid-1980s have greatly reduced poaching and allowed populations of buffalo, elephants, and rhinoceros to rebuild

Hofreiter, M., Lister, A., 2006. Mammoths
459. Curr. Biol. 16, R347-R348.

Joubert, D., 2006. Hunting behaviour of lions (Panthera leo) on elephants (Loxodonta africana) in the Chobe National Park, Botswana. African Journal of Ecology 44, 279-281.
Abstract: Megaherbivores like elephants and rhinos have been regarded as invulnerable to predation as adults (Owen-Smith, 1988; G. B. Schaller pers. comm.), although Guthrie (1990) suggests that lions hunted such large prey during the Pleistocene. Recently, there have been a number of observations of elephants killed by lions in northern Botswana, going as far back as 1985 (M. Slogrove pers. comm.). The hunting behaviour of lions on elephants, and the age and sex structure of the elephants killed, were observed at a waterhole in the Savute region of Chobe National Park. The first observed elephant kill was recorded in August 1991. Systematic records of elephants killed were made between 1993 and 1996.

Kalmykov, N.P., Mashchenko, E.N., 2006. New data on the migration of the family Elephantidae (Mammalia, Proboscidea) in Eurasia
480. Dokl. Biol. Sci. 406, 103-105.

Krause, J., Dear, P.H., Pollack, J.L., Slatkin, M., Spriggs, H., Barnes, I., Lister, A.M., Ebersberger, I., Paabo, S., Hofreiter, M., 2006. Multiplex amplification of the mammoth mitochondrial genome and the evolution of Elephantidae
533. Nature 439, 724-727.
Abstract: In studying the genomes of extinct species, two principal limitations are typically the small quantities of endogenous ancient DNA and its degraded condition, even though products of up to 1,600 base pairs (bp) have been amplified in rare cases. Using small overlapping polymerase chain reaction products, longer stretches of sequences or even whole mitochondrial genomes can be reconstructed, but this approach is limited by the number of amplifications that can be performed from rare samples. Thus, even from well-studied Pleistocene species such as mammoths, ground sloths and cave bears, no DNA sequences of more than about 1,000 bp have been reconstructed. Here we report the complete mitochondrial genome sequence of the Pleistocene woolly mammoth Mammuthus primigenius. We used about 200 mg of bone and a new approach that allows the simultaneous retrieval of multiple sequences from small amounts of degraded DNA. Our phylogenetic analyses show that the mammoth was more closely related to the Asian than to the African elephant. However, the divergence of mammoth, African and Asian elephants occurred over a short time, corresponding to only about 7% of the total length of the phylogenetic tree for the three evolutionary lineages

Larke, A., Crews, D.E., 2006. Parental investment, late reproduction, and increased reserve capacity are associated with longevity in humans. J Physiol Anthropol 25, 119-131.
Abstract: Throughout the living world trade-offs between reproductive success and longevity have been observed. In general, two extremes of life history patterning are reported, r- and K-selected species. The latter tend toward larger body sizes, few offspring from any one pregnancy, few offspring over the female reproductive span, longer life spans, and greater parental investment (PI: all efforts and expenses associated with the production, gestation, post-natal care, feeding, and protection of young) (e.g., whales, elephants, hominids). r-selected species tend toward smaller body size, multiple births/litters per pregnancy, female production of many gametes and offspring over the life span, and low levels of PI (e.g., most plants, insects, mice). These differences have significant influences on physiological variation among human populations.Across human samples, reproductive success (RS: the number of offspring successfully birthed and reared to reproductive age) has been reported to vary positively, negatively, and not at all with longevity of women. This complexity may be in part due to the fact that both early-life and late-life fecundity are associated with longevity in women, while total parity seems a poor gauge of female longevity in humankind. Large variations in associations of RS with longevity in women suggest that multiple factors may confound this association. One confounding factor is that among women, RS is largely determined not by fecundity, but by the quality of PI available to offspring. Among modern humans, PI is more complex, longer lasting (both relatively and absolutely), and extensive than for any other ammal. This suggests that modern human life history is a reflection of the co-evolution of longevity and extensive PI as part of our species' biocultural evolution. The need for long-term PI has greatly shaped human physiological variation and patterns of longevity.

Lyashchenko, K.P., Greenwald, R., Esfandiari, J., Olsen, J.H., Ball, R., Dumonceaux, G., Dunker, F., Buckley, C., Richard, M., Murray, S., Payeur, J.B., Andersen, P., Pollock, J.M., Mikota, S., Miller, M., Sofranko, D., Waters, W.R., 2006. Tuberculosis in elephants: antibody responses to defined antigens of Mycobacterium tuberculosis, potential for early diagnosis, and monitoring of treatment
438. Clin. Vaccine Immunol. 13, 722-732.
Abstract: Tuberculosis (TB) in elephants is a re-emerging zoonotic disease caused primarily by Mycobacterium tuberculosis. Current diagnosis relies on trunk wash culture, the only officially recognized test, which has serious limitations. Innovative and efficient diagnostic methods are urgently needed. Rapid identification of infected animals is a crucial prerequisite for more effective control of TB, as early diagnosis allows timely initiation of chemotherapy. Serology has diagnostic potential, although key antigens have not been identified and optimal immunoassay formats are not established. To characterize the humoral responses in elephant TB, we tested 143 serum samples collected from 15 elephants over time. These included 48 samples from five culture-confirmed TB cases, of which four were in Asian elephants infected with M. tuberculosis and one was in an African elephant with Mycobacterium bovis. Multiantigen print immunoassay (MAPIA) employing a panel of 12 defined antigens was used to identify serologic correlates of active disease. ESAT-6 was the immunodominant antigen recognized in elephant TB. Serum immunoglobulin G antibodies to ESAT-6 and other proteins were detected up to 3.5 years prior to culture of M. tuberculosis from trunk washes. Antibody levels to certain antigens gradually decreased in response to antitubercular therapy, suggesting the possibility of treatment monitoring. In addition to MAPIA, serum samples were evaluated with a recently developed rapid test (RT) based on lateral flow technology (ElephantTB STAT-PAK). Similarly to MAPIA, infected elephants were identified using the RT up to 4 years prior to positive culture. These findings demonstrate the potential for TB surveillance and treatment monitoring using the RT and MAPIA, respectively

Perelygin, A.A., Lear, T.L., Zharkikh, A.A., Brinton, M.A., 2006. Comparative analysis of vertebrate EIF2AK2 (PKR) genes and assignment of the equine gene to ECA15q24-q25 and the bovine gene to BTA11q12-q15
416. Genet. Sel Evol. 38, 551-563.
Abstract: The structures of the canine, rabbit, bovine and equine EIF2AK2 genes were determined. Each of these genes has a 5' non-coding exon as well as 15 coding exons. All of the canine, bovine and equine EIF2AK2 introns have consensus donor and acceptor splice sites. In the equine EIF2AK2 gene, a unique single nucleotide polymorphism that encoded a Tyr329Cys substitution was detected. Regulatory elements predicted in the promoter region were conserved in ungulates, primates, rodents, Afrotheria (elephant) and Insectifora (shrew). Western clawed frog and fugu EIF2AK2 gene sequences were detected in the USCS Genome Browser and compared to those of other vertebrate EIF2AK2 genes. A comparison of EIF2AK2 protein domains in vertebrates indicates that the kinase catalytic domains were evolutionarily more conserved than the nucleic acid-binding motifs. Nucleotide substitution rates were uniform among the vertebrate sequences with the exception of the zebrafish and goldfish EIF2AK2 genes, which showed substitution rates about 20% higher than those of other vertebrates. FISH was used to physically assign the horse and cattle genes to chromosome locations, ECA15q24-q25 and BTA11q12-15, respectively. Comparative mapping data confirmed conservation of synteny between ungulates, humans and rodents

Plotnik, J.M., de Waal, F.B., Reiss, D., 2006. Self-recognition in an Asian elephant
389. Proc. Natl. Acad. Sci. U. S. A 103, 17053-17057.
Abstract: Considered an indicator of self-awareness, mirror self-recognition (MSR) has long seemed limited to humans and apes. In both phylogeny and human ontogeny, MSR is thought to correlate with higher forms of empathy and altruistic behavior. Apart from humans and apes, dolphins and elephants are also known for such capacities. After the recent discovery of MSR in dolphins (Tursiops truncatus), elephants thus were the next logical candidate species. We exposed three Asian elephants (Elephas maximus) to a large mirror to investigate their responses. Animals that possess MSR typically progress through four stages of behavior when facing a mirror: (i) social responses, (ii) physical inspection (e.g., looking behind the mirror), (iii) repetitive mirror-testing behavior, and (iv) realization of seeing themselves. Visible marks and invisible sham-marks were applied to the elephants' heads to test whether they would pass the litmus "mark test" for MSR in which an individual spontaneously uses a mirror to touch an otherwise imperceptible mark on its own body. Here, we report a successful MSR elephant study and report striking parallels in the progression of responses to mirrors among apes, dolphins, and elephants. These parallels suggest convergent cognitive evolution most likely related to complex sociality and cooperation

Poinar, H.N., Schwarz, C., Qi, J., Shapiro, B., MacPhee, R.D., Buigues, B., Tikhonov, A., Huson, D.H., Tomsho, L.P., Auch, A., Rampp, M., Miller, W., Schuster, S.C., 2006. Metagenomics to paleogenomics: large-scale sequencing of mammoth DNA
529. Science 311, 392-394.
Abstract: We sequenced 28 million base pairs of DNA in a metagenomics approach, using a woolly mammoth (Mammuthus primigenius) sample from Siberia. As a result of exceptional sample preservation and the use of a recently developed emulsion polymerase chain reaction and pyrosequencing technique, 13 million base pairs (45.4%) of the sequencing reads were identified as mammoth DNA. Sequence identity between our data and African elephant (Loxodonta africana) was 98.55%, consistent with a paleontologically based divergence date of 5 to 6 million years. The sample includes a surprisingly small diversity of environmental DNAs. The high percentage of endogenous DNA recoverable from this single mammoth would allow for completion of its genome, unleashing the field of paleogenomics

Poulakakis, N., Parmakelis, A., Lymberakis, P., Mylonas, M., Zouros, E., Reese, D.S., Glaberman, S., Caccone, A., 2006. Ancient DNA forces reconsideration of evolutionary history of Mediterranean pygmy elephantids. Biol. Lett. 2, 451-454.
Abstract: During the Pleistocene pygmy elephantids, some only a quarter of their ancestors' size, were present on Mediterranean islands until about 10,000 years ago (y.a.). Using a new methodology for ancient DNA (aDNA) studies, the whole genomic multiple displacement amplification method, we were able to retrieve cytochrome b (cytb) DNA fragments from 4200 to 800,000 y.a. specimens from island and mainland samples, including pygmy and normal-sized forms. The short DNA sequence (43 bp) retrieved from the 800,000 y.a. sample is one of the oldest DNA fragment ever retrieved. Duplication of the experiments in two laboratories, the occurrence of three diagnostic sites and the results of the phylogenetic analyses strongly support its authenticity. Our results challenge the prevailing view that pygmy elephantids of the eastern Mediterranean originated exclusively from Elephas, suggesting independent histories of dwarfism and the presence of both pygmy mammoths and elephant-like taxa on these islands. Based on our molecular data, the origin of the Tilos and Cyprus elephantids from a lineage within the genus Elephas is confirmed, while the DNA sequence from the Cretan sample falls clearly within the mammoth clade. Thus, the name Mammuthus creticus rather than Elephas creticus, seems to be justified for this form. Our findings also suggest a need to re-evaluate the evolutionary history of the Sicilian/Maltese species, traditionally included in the genus Elephas

Rogaev, E.I., Moliaka, Y.K., Malyarchuk, B.A., Kondrashov, F.A., Derenko, M.V., Chumakov, I., Grigorenko, A.P., 2006. Complete mitochondrial genome and phylogeny of Pleistocene mammoth Mammuthus primigenius
507. PLoS. Biol. 4, e73.
Abstract: Phylogenetic relationships between the extinct woolly mammoth (Mammuthus primigenius), and the Asian (Elephas maximus) and African savanna (Loxodonta africana) elephants remain unresolved. Here, we report the sequence of the complete mitochondrial genome (16,842 base pairs) of a woolly mammoth extracted from permafrost-preserved remains from the Pleistocene epoch--the oldest mitochondrial genome sequence determined to date. We demonstrate that well-preserved mitochondrial genome fragments, as long as approximately 1,600-1700 base pairs, can be retrieved from pre-Holocene remains of an extinct species. Phylogenetic reconstruction of the Elephantinae clade suggests that M. primigenius and E. maximus are sister species that diverged soon after their common ancestor split from the L. africana lineage. Low nucleotide diversity found between independently determined mitochondrial genomic sequences of woolly mammoths separated geographically and in time suggests that north-eastern Siberia was occupied by a relatively homogeneous population of M. primigenius throughout the late Pleistocene

Rompler, H., Rohland, N., Lalueza-Fox, C., Willerslev, E., Kuznetsova, T., Rabeder, G., Bertranpetit, J., Schoneberg, T., Hofreiter, M., 2006. Nuclear gene indicates coat-color polymorphism in mammoths
439. Science 313, 62.
Abstract: By amplifying the melanocortin type 1 receptor from the woolly mammoth, we can report the complete nucleotide sequence of a nuclear-encoded gene from an extinct species. We found two alleles and show that one allele produces a functional protein whereas the other one encodes a protein with strongly reduced activity. This finding suggests that mammoths may have been polymorphic in coat color, with both dark- and light-haired individuals co-occurring

Rothschild, B.M., Martin, L.D., 2006. Did ice-age bovids spread tuberculosis? Naturwissenschaften 93, 565-569.
Abstract: Pathognomonic metacarpal undermining is a skeletal pathology that has been associated with Mycobacterium tuberculosis in bovids. Postcranial artiodactyl, perissodactyl, and carnivore skeletons were examined in major university and museum collections of North America and Europe for evidence of this and other pathology potentially attributable to tuberculosis. Among nonproboscidean mammals from pre-Holocene North America, bone lesions indicative of tuberculosis were restricted to immigrant bovids from Eurasia. No bone lesions compatible
with diagnosis of tuberculosis were found in large samples of other pre-Holocene (164 Oligocene, 397 Miocene, and 1,041 Plio-Pleistocene) North American mammals, including
114 antilocaprids. Given the unchanged frequency of bovid tubercular disease during the Pleistocene, it appears that most did not die from the disease but actually reached an
accommodation with it (as did the mastodon) (Rothschild and Laub 2006). Thus, they were sufficiently long-lived to assure greater spread of the disease. The relationships of the
proboscidean examples need further study, but present evidence suggests a Holarctic spread of tuberculosis during the Pleistocene, with bovids acting as vectors. While the role of other animals in the transmission of tuberculosis could be considered, the unique accommodation achieved by bovids and mastodons makes them the likely "culprits" in its spread.

Rothschild, B.M., Laub, R., 2006. Hyperdisease in the late Pleistocene:validation of an early 20th century hypothesis. Naturwissenschaften 93, 557-564.

Rubenstein, D.R., Sherman, P.W., Gavin, T.A., 2006. Pleistocene park: Does re-wilding North America represent sound conservation for the 21st century? Biological Conservation 132, 232-238.
Abstract: A group of conservation biologists recently proposed to populate western North America with African and Asian megafauna, including lions, elephants, cheetahs, and camels, to create a facsimile of a species assemblage that disappeared from the continent some 13,000 years ago. The goals of this program, known as "Pleistocene re-wilding", are to restore some of the evolutionary and ecological potential that was lost from North America during the Pleistocene extinctions, and help prevent the extinction of selected African and Asian mammals. Pleistocene re-wilders justify this conservation strategy on ethical and aesthetic grounds, arguing that humans have a moral responsibility to make amends for overexploitation by our ancestors. They believe that the flora of many North American terrestrial ecosystems has gone basically unchanged since the end of the Pleistocene, so re-wilding would help restore evolutionary and ecological potential and improve ecosystem functioning. This paper discusses some of the pros and cons of this proposal, including the ethical, aesthetic, ecological, and evolutionary issues, assesses its potential economic and political impacts on other conservation practices, both in North America and elsewhere, and reviews the realities of large mammal reintroductions. It is concluded that Pleistocene re-wilding with exotic species will not restore the evolutionary or ecological potential of native North American species nor extinct Pleistocene megafauna and their ancient ecosystems, but may instead jeopardize indigenous species and North American ecosystems. Resources would be better spent on preserving threatened organisms in their native habitats and reintroducing them to places in their historical ranges from which they were only recently extirpated.

Schmitt, D., Cartmill, M., Griffin, T.M., Hanna, J.B., Lemelin, P., 2006. Adaptive value of ambling gaits in primates and other mammals
460. J. Exp. Biol. 209, 2042-2049.
Abstract: At speeds between the walk and the gallop, most mammals trot. Primates almost never trot, and it has been claimed that they transition directly from a walk to a gallop without any distinctive mid-speed running gait. If true, this would be another characteristic difference between the locomotion of primates and that of most other quadrupedal mammals. Presently, however, few data exist concerning the actual presence or absence of intermediate-speed gaits (i.e. gaits that are used between a walk and a gallop) in primates. Video records of running in twelve primate species reveal that, unlike most other mammals, all the primates studied almost exclusively adopt an 'amble'--an intermediate-speed running gait with no whole-body aerial phase--rather than trot. Ambling is also common in elephants and some horses, raising the question of why ambling is preferred over trotting in these diverse groups of animals. Mathematical analyses presented here show that ambling ensures continuous contact of the body with the substrate while dramatically reducing vertical oscillations of the center of mass. This may explain why ambling appears to be preferable to trotting for extremely large terrestrial mammals such as elephants and for arboreal mammals like primates that move on unstable branches. These findings allow us to better understand the mechanics of these unusual running gaits and shed new light on primate locomotor evolution

Shoshani, J., Kupsky, W.J., Marchant, G.H., 2006. Elephant brain. Part I: gross morphology, functions, comparative anatomy, and evolution
446. Brain Res. Bull. 70, 124-157.
Abstract: We report morphological data on brains of four African, Loxodonta africana, and three Asian elephants, Elephas maximus, and compare findings to literature. Brains exhibit a gyral pattern more complex and with more numerous gyri than in primates, humans included, and in carnivores, but less complex than in cetaceans. Cerebral frontal, parietal, temporal, limbic, and insular lobes are well developed, whereas the occipital lobe is relatively small. The insula is not as opercularized as in man. The temporal lobe is disproportionately large and expands laterally. Humans and elephants have three parallel temporal gyri: superior, middle, and inferior. Hippocampal sizes in elephants and humans are comparable, but proportionally smaller in elephant. A possible carotid rete was observed at the base of the brain. Brain size appears to be related to body size, ecology, sociality, and longevity. Elephant adult brain averages 4783 g, the largest among living and extinct terrestrial mammals; elephant neonate brain averages 50% of its adult brain weight (25% in humans). Cerebellar weight averages 18.6% of brain (1.8 times larger than in humans). During evolution, encephalization quotient has increased by 10-fold (0.2 for extinct Moeritherium, approximately 2.0 for extant elephants). We present 20 figures of the elephant brain, 16 of which contain new material. Similarities between human and elephant brains could be due to convergent evolution; both display mosaic characters and are highly derived mammals. Humans and elephants use and make tools and show a range of complex learning skills and behaviors. In elephants, the large amount of cerebral cortex, especially in the temporal lobe, and the well-developed olfactory system, structures associated with complex learning and behavioral functions in humans, may provide the substrate for such complex skills and behavior

Shoshani, J., Kupsky, W.J., Marchant, G.H., 2006. Elephant brain. Part I: gross morphology, functions, comparative anatomy, and evolution. Brain Res Bull 70, 124-157.
Abstract: We report morphological data on brains of four African, Loxodonta africana, and three Asian elephants, Elephas maximus, and compare findings to literature. Brains exhibit a gyral pattern more complex and with more numerous gyri than in primates, humans included, and in carnivores, but less complex than in cetaceans. Cerebral frontal, parietal, temporal, limbic, and insular lobes are well developed, whereas the occipital lobe is relatively small. The insula is not as opercularized as in man. The temporal lobe is disproportionately large and expands laterally. Humans and elephants have three parallel temporal gyri: superior, middle, and inferior. Hippocampal sizes in elephants and humans are comparable, but proportionally smaller in elephant. A possible carotid rete was observed at the base of the brain. Brain size appears to be related to body size, ecology, sociality, and longevity. Elephant adult brain averages 4783 g, the largest among living and extinct terrestrial mammals; elephant neonate brain averages 50% of its adult brain weight (25% in humans). Cerebellar weight averages 18.6% of brain (1.8 times larger than in humans). During evolution, encephalization quotient has increased by 10-fold (0.2 for extinct Moeritherium, approximately 2.0 for extant elephants). We present 20 figures of the elephant brain, 16 of which contain new material. Similarities between human and elephant brains could be due to convergent evolution; both display mosaic characters and are highly derived mammals. Humans and elephants use and make tools and show a range of complex learning skills and behaviors. In elephants, the large amount of cerebral cortex, especially in the temporal lobe, and the well-developed olfactory system, structures associated with complex learning and behavioral functions in humans, may provide the substrate for such complex skills and behavior.

Shoshani, J., Walter, R.C., Abraha, M., Berhe, S., Tassy, P., Sanders, W.J., Marchant, G.H., Libsekal, Y., Ghirmai, T., Zinner, D., 2006. A proboscidean from the late Oligocene of Eritrea, a "missing link" between early Elephantiformes and Elephantimorpha, and biogeographic implications
384. Proc. Natl. Acad. Sci. U. S. A 103, 17296-17301.
Abstract: We report on a late Oligocene proboscidean species from Eritrea, dated to 26.8 +/- 1.5 Mya. This "missing link" between early elephantiformes and Elephantimorpha is the oldest known nongomphothere proboscidean to probably display horizontal tooth displacement, typical of elephants [Elephantimorpha consists of Mammutida (mastodons) and Elephantida, and Elephantida includes gomphotheres, stegodons, and elephants]. Together with the newly discovered late Oligocene gomphotheres from Chilga, Ethiopia, the Eritrean taxon points to the importance of East Africa as a major area for the knowledge of the early evolution of Elephantimorpha before the faunal exchange between Eurasia and Africa

Shoshani, J., Walter, R.C., Abraha, M., Berhe, S., Tassy, P., Sanders, W.J., Marchant, G.H., Libsekal, Y., Ghirmai, T., Zinner, D., 2006. A proboscidean from the late Oligocene of Eritrea, a "missing link" between early Elephantiformes and Elephantimorpha, and biogeographic implications. Proc Natl Acad Sci U S A. 103, 17296-17301pdf.
Abstract: We report on a late Oligocene proboscidean species from Eritrea, dated to 26.8 +/- 1.5 Mya. This "missing link" between early elephantiformes and Elephantimorpha is the oldest known nongomphothere proboscidean to probably display horizontal tooth displacement, typical of elephants [Elephantimorpha consists of Mammutida (mastodons) and Elephantida, and Elephantida includes gomphotheres, stegodons, and elephants]. Together with the newly discovered late Oligocene gomphotheres from Chilga, Ethiopia, the Eritrean taxon points to the importance of East Africa as a major area for the knowledge of the early evolution of Elephantimorpha before the faunal exchange between Eurasia and Africa.Department of Biology, University of Asmara, PO Box 1220, Asmara, Eritrea. jshosh@sun.science.wayne.edu

Uni, S., Bain, O., Agatsuma, T., Katsumi, A., Baba, M., Yanai, T., Takaoka, H., 2006. New filarial nematode from Japanese serows (Naemorhedus crispus: Bovidae) close to parasites from elephants
403. Parasite 13, 193-200.
Abstract: A new onchocercid species, Loxodontofilaria caprini n. sp. (Filarioidea: Nematoda), found in subcutaneous tissues of 37 (33%) of 112 serows (Noemorhedus crispus) examined in Japan, is described. The female worm had the characteristics of Loxodontofilaria, e.g., the large body size, well-developed esophagus with a shallow buccal cavity, and the long tail with three caudal lappets. The male worm of the new species, which was first described in the genus, had unequal length of spicules, 10 pairs of pre- and post-caudal papillae, and three terminal caudal lappets. Deirids were present in both sexes. Among four species of the genus loxodontofiloria: one from the hippopotamus and three from the Elepantidae, L. caprini n. sp. appears close to L. asiatica Bain, Baker & Chabaud, 1982, a subcutaneous parasite of Elephas indicus in Myanmar (Burma). However, L. caprini n. sp. is distinct from L. asiatica in that the Japanese female worm has an esophagus half as long and the microfilariae also half as long with a coiled posterior. The microfilariae were found in the skin of serows. The new parasite appears to clearly illustrate a major event in the evolution of onchocercids: the host-switching. This might have occurred on the Eurasian continent, where elephantids and the lineage of rupicaprines diversified during the Pliocene-Pleistocene, or in Japan, into which some of these hosts migrated

Vinogradov, I.V., Kochneva, G.V., Shchelkunov, S.N., Riabchikova, E.I., 2006. [Reproduction of cowpox virus strain EP-2 isolated from an elephant in primary fibroblast cultures and chorion-allantoic chick embryos]
451. Vopr. Virusol. 51, 44-48.
Abstract: Electron microscopy was used to study the reproduction of cowpox virus strain EP-2 in the cells of a primary fibroblast cultures (PFC) and chorion-allantoic membrane (CAM) of chick embryos (CE). The sequential stages of viral morphogenesis and the structure of A-type inclusions were described. The parameters of viral reproduction in PFC and CE CAM were compared. The formation of crystalloid tubular structures in PFC, unusual electron dense inclusions in the cells of CE CAN, and different variants of A-type inclusions in the cells of a pock was found. The histological and ultrastructural characteristics of pocks in CE CAM are described

Agnew, D.W., Hagey, L., Shoshani, J., 2005. The elephants of Zoba Gash Barka, Eritrea: part 4. Cholelithiasis in a wild African elephant (Loxodonta africana). J. Zoo. Wildl. Med. 36, 677-683.
Abstract: A 4.0-kg cholelith was found within the abdominal cavity of a dead wild African elephant (Loxodonta africana) in Eritrea. Analysis of this cholelith by histochemistry, electron microscopy, electrospray mass spectroscopy, and energy-dispersive x-ray spectroscopy revealed it was composed of bile alcohols but no calcium, bilirubin, or cholesterol. Bacteria were also found in the cholelith. Similar, but smaller, bile stones have been identified previously in other wild African elephants and an excavated mammoth (Mammuthus columbi). Choleliths have been reported only once in a captive Asian elephant (Elephas maximus). Elephants, along with hyraxes (Procavia capensis) and manatees (Trichechus manatus), are unique among mammals in producing only bile alcohols and no bile acids, which may predispose them to cholelithiasis, particularly in association with bacterial infection. Dietary factors may also play an important role in cholelith formation.

Debruyne, R., 2005. A case study of apparent conflict between molecular phylogenies: the interrelationships of African elephants. Cladistics 21, 31-50.
Abstract: Recent molecular phylogenies of the African elephants suggest that there is an evolutionary structure within Loxodonta africana. Some nuclear results (Roca et al., 2001) support the separation of the forest African elephant subspecies L. a. cyclotis as a species distinct from the savannah elephant L. a. africana, on the basis of the recognition of both forming highly divergent (reciprocally monophyletic) clades. Conversely, a mitochondrial survey (Eggert et al., 2002), while admitting a geographic partitioning of the genetic structure within African elephants, suggests retaining the status quo. They recognize three diagnosible entities (western, central and south-eastern Africa) with non-overlapping ranges within L. africana sensu lato. In order to address these con.icting views (historical fragmentation and speciation or isolation by distance, respectively), we have sequenced two datasets of 1961 bp (for 50 elephants) and about 3700 bp, respectively (for 20 elephants) of the mitochondrial DNA for both forms of elephants (cyclotis and africana). They span the cytochrome b gene, the control region and several RNAs. When compared with former mtDNA data, they provide the most comprehensive view of the African elephant phylogeny (78 mtDNA haplotypes, of which 44 are new) and provide the .rst insight into populations from the Democratic Republic of Congo. The genetic diversity of mtDNA was appraised and the stability of alternative phylogenetic trees was investigated. Our results are inconsistent with both those prior studies. They revealed two highly divergent molecular clades referred to as F and S, that do not conform to the morphological delineations of cyclotis and africana. A non-negligible proportion of specimens of L. a. africana display haplotypes prevailing in forest elephant populations (clade F). The geographic distribution of clades and areas of their co-occurrence support the hypothesis of incomplete isolation between forest and savannah African elephant populations, followed by recurrent interbreeding between the two forms. We state that the conclusions of prior studies resulted from insufficient character and / or geographic sampling. We conclude that there is no satisfying argument which can recognize two or more species of African elephants. We briefly comment on the meaning of such an attitude in a conservation viewpoint.

Delves, P.J., Roitt, I.M., 2005. Vaccines for the control of reproduction--status in mammals, and aspects of comparative interest
592. Dev. Biol. (Basel) 121, 265-273.
Abstract: The objective of producing vaccines which target elements of the reproductive system to control fertility has been pursued for many years. Of the many targets for such vaccines, several sperm-associated antigens have been proposed for antibody-mediated intervention before fertilization but the very abundance of antigen to be neutralized has been a barrier. Zona pellucida antigens associated with the surface of the oocyte have also been targeted and used successfully for control of 'wild' elephant populations but worries concerning immunopathologically-mediated tissue damage have been mooted. Vaccines using human chorionic gonadotropin (hCG) which is required for the implantation and maintenance of the fertilized egg, although of interest for the development of fertility control in human populations, has no relevance in the context of the present conference because external fertilization of fish eggs is independent. The pathways by which gonadotropin-releasing hormone (GnRH) secreted by the hypothalamus promote release of luteinizing (LH) and follicle-stimulating hormone (FSH) which govern the physiological maturation and maintenance of the reproductive organs, provide many targets for immunological intervention. Most consistent success has been reported using GnRH-based vaccines which are immunosterilizing in a variety of mammalian species such as pigs, rodents and white-tailed deer. The fact that the structure of the decapeptide, GnRH, has been maintained over so many years of evolution and been conserved across so many animal species, encourages the view that a strategy for control of sexual maturation in fish based upon stimulation of GnRH antibodies may well prove to be a practical proposition, provided the formulation of an appropriate highly immunogenic vaccine can be achieved

Edwards, H.G., Jorge Villar, S.E., Nik Hassan, N.F., Arya, N., O'Connor, S., Charlton, D.M., 2005. Ancient biodeterioration: an FT-Raman spectroscopic study of mammoth and elephant ivory
576. Anal. Bioanal. Chem. 383, 713-720.
Abstract: Raman spectra of mammoth ivory specimens have been recorded using near-infrared excitation, and comparisons made with modern Asian and African elephant ivories. Whereas the most ancient mammoth ivory (60-65 ky) showed no evidence for an organic collagen component, more recent samples of mammoth ivory indicated that some preservation had occurred, although with biodeterioration of the protein structure exhibited by the amide I and III bands in the 1200-1700 cm(-1) region of the Raman spectrum. The consequent difficulties encountered when applying chemometrics methods to ancient ivory analysis (which are successful for modern specimens) are noted. In the most ancient mammoth ivory specimens, which are extensively fragmented, evidence of mineralization is seen, with the production of gypsum, calcite and limonite; Raman microscopic analysis of crystalline material inside the fissures of the mammoth ivory shows the presence of gypsum as well as cyanobacterial colonisation. The application of Raman spectroscopy to the nondestructive analysis of archaeological materials in order to gain information of relevance to their preservation or restoration is highlighted

Edwards, H.G.M., Hassan, N.F.N., Arya, N., 2005. Evaluation of Raman spectroscopy and application of chemometric methods for the differentiation of contemporary ivory specimens I: elephant and mammalian species. Journal of Raman Spectroscopy 37, 353-360.
Abstract: Specimens of mammoth, African and Asian ivory dentine, and other mammalian species were examined using Fourier-Transform (FT), conventional dispersive (confocal) and remote-sensing portable Raman spectroscopy, all with near-infrared laser excitation (1064 and 785 nm). FT-Raman spectroscopy produced the best quality spectra for differentiation purposes and the application of a fibre probe coupled to a portable Raman spectrometer has also been demonstrated and proposed for the in situ characterization of suspected contraband ivories at airports. In addition to the visual comparison of spectral features, chemometric methods are used to discriminate between African and Asian elephant dentine by analyzing normalized integrated band areas in ten selected wavenumber regions. Principal component analysis separates the spectra of both species into two well-defined groups based upon their organic and inorganic composition. By means of stepwise discriminant analysis almost 98% of the spectra are correctly classified to their species group memberships.

Gibbons, A., 2005. Ancient DNA. New methods yield mammoth samples
527. Science 310, 1889.

Glickman, S.E., Short, R.V., Renfree, M.B., 2005. Sexual differentiation in three unconventional mammals: spotted hyenas, elephants and tammar wallabies
566. Horm. Behav. 48, 403-417.
Abstract: The present review explores sexual differentiation in three non-conventional species: the spotted hyena, the elephant and the tammar wallaby, selected because of the natural challenges they present for contemporary understanding of sexual differentiation. According to the prevailing view of mammalian sexual differentiation, originally proposed by Alfred Jost, secretion of androgen and anti-Mullerian hormone (AMH) by the fetal testes during critical stages of development accounts for the full range of sexually dimorphic urogenital traits observed at birth. Jost's concept was subsequently expanded to encompass sexual differentiation of the brain and behavior. Although the central focus of this review involves urogenital development, we assume that the novel mechanisms described in this article have potentially significant implications for sexual differentiation of brain and behavior, a transposition with precedent in the history of this field. Contrary to the "specific" requirements of Jost's formulation, female spotted hyenas and elephants initially develop male-type external genitalia prior to gonadal differentiation. In addition, the administration of anti-androgens to pregnant female spotted hyenas does not prevent the formation of a scrotum, pseudoscrotum, penis or penile clitoris in the offspring of treated females, although it is not yet clear whether the creation of masculine genitalia involves other steroids or whether there is a genetic mechanism bypassing a hormonal mediator. Wallabies, where sexual differentiation occurs in the pouch after birth, provide the most conclusive evidence for direct genetic control of sexual dimorphism, with the scrotum developing only in males and the pouch and mammary glands only in females, before differentiation of the gonads. The development of the pouch and mammary gland in females and the scrotum in males is controlled by genes on the X chromosome. In keeping with the "expanded" version of Jost's formulation, secretion of androgens by the fetal testes provides the best current account of a broad array of sex differences in reproductive morphology and endocrinology of the spotted hyena, and androgens are essential for development of the prostate and penis of the wallaby. But the essential circulating androgen in the male wallaby is 5alpha androstanediol, locally converted in target tissues to DHT, while in the pregnant female hyena, androstenedione, secreted by the maternal ovary, is converted by the placenta to testosterone (and estradiol) and transferred to the developing fetus. Testicular testosterone certainly seems to be responsible for the behavioral phenomenon of musth in male elephants. Both spotted hyenas and elephants display matrilineal social organization, and, in both species, female genital morphology requires feminine cooperation for successful copulation. We conclude that not all aspects of sexual differentiation have been delegated to testicular hormones in these mammals. In addition, we suggest that research on urogenital development in these non-traditional species directs attention to processes that may well be operating during the sexual differentiation of morphology and behavior in more common laboratory mammals, albeit in less dramatic fashion

Glickman, S.E., Short, R.V., Renfree, M.B., 2005. Sexual differentiation in three unconventional mammals: Spotted hyenas, elephants and tammar wallabies. Hormones and Behaviour 48, 403-417.
Abstract: The present review explores sexual differentiation in three non-conventional species: the spotted hyena, the elephant and the tammar wallaby, selected because of the natural challenges they present for contemporary understanding of sexual differentiation. According to the prevailing view of mammalian sexual differentiation, originally proposed by Alfred Jost, secretion of androgen and anti-Mullerian hormone (AMH) by the fetal testes during critical stages of development accounts for the full range of sexually dimorphic urogenital traits observed at birth. Jost's concept was subsequently expanded to encompass sexual differentiation of the brain and behavior. Although the central focus of this review involves urogenital development, we assume that the novel mechanisms described in this article have potentially significant implications for sexual differentiation of brain and behavior, a transposition with precedent in the history of this field. Contrary to the ''specific'' requirements of Jost's formulation, female spotted hyenas and elephants initially develop male-type external genitalia prior to gonadal differentiation. In addition, the administration of anti-androgens to pregnant female spotted hyenas does not prevent the formation of a scrotum, pseudoscrotum, penis or penile clitoris in the offspring of treated females, although it is not yet clear whether the creation of masculine genitalia involves other steroids or whether there is a genetic mechanism bypassing a hormonal mediator. Wallabies, where sexual differentiation occurs in the pouch after birth, provide the most conclusive evidence for direct genetic control of sexual dimorphism, with the scrotum developing only in males and the pouch and mammary glands only in females, before differentiation of the gonads. The development of the pouch and mammary gland in females and the scrotum in males is controlled by genes on the X chromosome. In keeping with the ''expanded'' version of Jost's formulation, secretion of androgens by the fetal testes provides the best current account of a broad array of sex differences in reproductive morphology and endocrinology of the spotted hyena, and androgens are essential for development of the prostate and penis of the wallaby. But the essential circulating androgen in the male wallaby is 5α androstanediol, locally converted in target tissues to DHT, while in the pregnant female hyena, androstenedione, secreted by the maternal ovary, is converted by the placenta to testosterone (and estradiol) and transferred to the developing fetus. Testicular testosterone certainly seems to be responsible for the behavioral phenomenon of musth in male elephants. Both spotted hyenas and elephants display matrilineal social organization, and, in both species, female genital morphology requires feminine cooperation for successful copulation. We conclude that not all aspects of sexual differentiation have been delegated to testicular hormones in these mammals. In addition, we suggest that research on urogenital development in these non-traditional species directs attention to processes that may well be operating during the sexual differentiation of morphology and behavior in more common laboratory mammals, albeit in less dramatic fashion.

Hambler, C., Henderson, P.A., Speight, M.R., 2005. Elephants, Ecology and Nonequilibrium? Science 307, 673c-674c.
Abstract: Elephants and thrips may have something in common: It has been proposed that elephants in Africa do not reach carrying capacity because they inhabit "nonequilibrium" ecosystems with highly variable rainfall ("Space-the final frontier for economists and elephants," E. Bulte et al., Perspectives, 15 Oct., p. 420). Similarly, it has been proposed that thrips in Australia do not reach a carrying capacity because of climatic fluctuations. The nonequilibrium (density-independent) ideas of the 1950s are being reworked as "state-of-the-art" ecological theory by Bulte et al. We should remember, however, that a more sophisticated analysis of the same thrip populations revealed strongly density dependent population change and hence a carrying capacity. The suggestion that multispecies systems are unlikely to show density dependence is erroneous. In contrast, evidence is emerging of the very widespread occurrence of density dependence, even in complex marine systems. Detection of such effects typically takes over four generations; well over a hundred years might be needed to detect density dependence in an elephant population. The application of nonequilibrium hypotheses to savannah has been challenged on theoretical and empirical grounds. Models indicate that herbivores in semi-arid areas are in long-term equilibrium with a subset of their resources. Competitive regulation is now very clear in a number of species of large herbivores in Africa, including wildebeest. A review concluded there was no evidence of a paradigm shift to a nonequilibrium perspective among those researching grasslands. Large species are unlikely to exhibit metapopulation dynamics. The harvesting of elephants is, rightly, controversial. We suggest that the nonequilibrium perspective is unlikely to clarify how their populations might respond to management.

Kalmykov, N.P., Maschenko, E.N., 2005. The oldest representative of Elephantidae (Mammalia, Proboscidea) in Asia. Paleontological Journal 39, 652-659.
Abstract: The earliest representative of the family Elephantidae in Asia,  Archidiskodon sp. From the Udunga locality (western Transbaikalia, Early Pliocene, MN 15), is described. The age of the accompanying mammal fauna suggests simultaneous migrations of mammuthoid elephants from Africa to Europe and Asia. The material includes a right dp(3) and isolated tooth  plates from another dp(3). The structure of dp(3) indicates that the  Early  Pliocene members of Elephantidae from western Transbaikalia displayed the basic morphological features of this family, i.e., the successive  formation  of dp(3), plate number, hypsodonty, and enamel thickness.

Larsen, R.S., Kay, M., Triantis, J., Salman, M.D. Update on serological detection of Mycobacterium tuberculosis infection in Asian elephants. 2005 Proceedings AAZV, AAWV, AZA Nutrition Advisory Group.  62-63. 2005.
Ref Type: Conference Proceeding
Abstract: Tuberculosis has become an important disease in captive elephants, particularly Asian elephants (Elephas maximus). Diagnosing tuberculosis in elephants has been problematic as many tests have inadequate sensitivity or specificity.2-4 A multiple-antigen enzyme-linked immunosorbent assay (ELISA) was previously investigated for detecting infection in Asian elephants and African elephants (Loxodonta africana); this test had excellent sensitivity and specificity, but needed further evaluation.1 Modifications to the multiple-antigen ELISA panel have since been made. Valuable antigens were retained, other antigens were removed, and new ones were added.  This modified ELISA was re-evaluated, using serum from 68 Asian elephants. Sixteen had M. tuberculosis -positive trunk cultures, while 52 were either culture negative at necropsy or had a history of negative trunk cultures and no contact with infected elephants. Seven elephants were evaluated over time. The test was 100% (95% CI; 95-100%) specific and 94% (95% CI; 79-100%) sensitive using two of the six antigens (M. bovis strain AN5 culture filtrate and M. tuberculosis early secretory antigenic target 6). "Effectively-treated" elephants had decreasing seroreactivity, but those that were culture-positive post-treatment were more consistently seroreactive.  Although "effectivelytreated" elephants had declining seroreactivity, they still usually had higher values than animals that had never been infected. Serology continues to show great promise in detecting tuberculosis in elephants, often detecting infection months-to-years sooner than trunk wash culture.  Advances in techniques may soon make serology even more practical.  While serology should not replace trunk-wash culture, it is a useful adjunct for early detection of infection in elephants and for monitoring treatment.
ACKNOLWEDGMENTS We thank the many veterinarians, owners, caretakers, and managers of elephant-owning institutions that participated in this investigation, as well as Drs. Michele Miller and Susan Mikota for helping to coordinate sample collection. We also thank Kimberly Deines and other laboratory personnel who processed ELISA samples.  The study was partially funded by a grant from USDA, CSREES to Colorado State University Program of Economically Important Infectious Animal Diseases.
LITERATURE CITED
1.Larsen, R.S., M.D. Salman, S.K. Mikota, R. Isaza, R.J. Montali, and J. Triantis. 2000.  Evaluation of a multiple-antigen enzyme-linked immunosorbent assay for detection of Mycobacterium tuberculosis in captive elephants.  J. Zoo Wildl. Med. 31: 291-302.
2. Mikota, S.K., L. Peddie, J. Peddie, R. Isaza, F. Dunker, G. West, W. Lindsay, R.S. Larsen, M.D. Salman, D. Chatterjee, J. Payeur, D. Whipple, C. Thoen, D.S. Davis, R.J. Montali and J. Maslow.  2001. Epidemiology and diagnosis of Mycobacterium tuberculosis in six groups of elephants.  J. Zoo Wildl. Med. 32: 1-16.
3. Mikota, S.K., R.S. Larsen, and R.J. Montali.  2000.  Tuberculosis in elephants in North America.  Zoo Biol. 19: 393-403.
4. U.S. Department of Agriculture.  2003.  Guidelines for the control of tuberculosis in elephants.  Animal and Plant Health Inspection Service; Animal Care. Washington, D.C. http://www.aphis.usda.gov/ac/TBGuidelines2003.pdf.

Lewerin, S.S., Olsson, S.L., Eld, K., Roken, B., Ghebremichael, S., Koivula, T., Kallenius, G., Bolske, G., 2005. Outbreak of Mycobacterium tuberculosis infection among captive Asian elephants in a Swedish zoo
637. Veterinary Record 156, 171-175.
Abstract: Between 2001 and 2003, there was an outbreak of tuberculosis in a Swedish zoo which involved elephants, giraffes, rhinoceroses and buffaloes. Cultures of trunk lavages were used to detect infected elephants, tuberculin testing was used in the giraffes and buffaloes, and tracheal lavage and tuberculin testing were used in the rhinoceroses. The bacteria isolated were investigated by spoligotyping and restriction fragment length polymorphism. Five elephants and one giraffe were found to have been infected by four different strains of Mycobacterium tuberculosis

Loannidis, J.P.A., 2005. Why most published research findings are false. PLoS Med 2, e124.

Nishihara, H., Satta, Y., Nikaido, M., Thewissen, J.G., Stanhope, M.J., Okada, N., 2005. A retroposon analysis of Afrotherian phylogeny
600. Molecular Biology and Evolution 22, 1823-1833.
Abstract: Recent comprehensive studies of DNA sequences support the monophyly of Afrotheria, comprising elephants, sirenians (dugongs and manatees), hyraxes, tenrecs, golden moles, aardvarks, and elephant shrews, as well as that of Paenungulata, comprising elephants, sirenians, and hyraxes. However, phylogenetic relationships among paenungulates, as well as among nonpaenungulates, have remained ambiguous. Here we applied an extensive retroposon analysis to these problems to support the monophyly of aardvarks, tenrecs, and golden moles, with elephant shrews as their sister group. Regarding phylogenetic relationships in Paenungulata, we could characterize only one informative locus, although we could isolate many insertions specific to each of three lineages, namely, Proboscidea, Sirenia, and Hyracoidea. These data prompted us to reexamine phylogenetic relationships among Paenungulata using 19 nuclear gene sequences resulting in three different analyses, namely, short interspersed element (SINE) insertions, nuclear sequence analyses, and morphological cladistics, supporting different respective phylogenies. We concluded that these three lineages diverged very rapidly in a very short evolutionary period, with the consequence that ancestral polymorphism present in the last common ancestor of Paenungulata results in such incongruence. Our results suggest the rapid fixation of many large-scale morphological synapomorphies for Tethytheria; implications of this in relation to the morphological evolution in Paenungulata are discussed

Palombo, M.R., Ferretti, M.P., 2005. Elephant fossil record from Italy: Knowledge, problems, and perspectives. Quaternary International 126-28, 107-136.
Abstract: The earliest occurrence of elephantines in Italy is in the middle Villafranchian (late Middle Pliocene; ca. 2.6 Ma), with a primitive representative of the mammoth lineage. In addition to this still poorly known taxon, four elephant species are clearly recognized in Plio-Pleistocene fossil mammal assemblages from the Italian peninsula: Mammuthus meridionalis, M. trogontherii, M. primigenius, and Elephas (Palaeoloxodon) antiquus. In Sicily, at least three different taxa are present during the Pleistocene: the dwarf E. falconeri, the medium sized E. (P.) "mnaidriensis", and a third poorly known large sized taxon, represented by isolated findings, in some cases apparently associated with E. (P.) "mnaidriensis". A number of specimens from Spinagallo and Luparello, of intermediate size between E. "mnaidriensis" and E falconeri, suggest the occurrence in Sicily of a further elephant species. Sardinia records the only case of an endemic small sized Mammuthus species from the western Mediterranean. The Italian elephant fossil recordclearly shows the influence of climatic, physiographic and paleogeographic conditions on the pattern of occurrence and dispersal of this mammal group in Southern Europe.

Perez-Barberia, F.J., Gordon, I.J., 2005. Gregariousness increases brain size in ungulates
586. Oecologia. 145, 41-52.
Abstract: The brain's main function is to organise the physiological and behavioural responses to environmental and social challenges in order to keep the organism alive. Here, we studied the effects that gregariousness (as a measurement of sociality), dietary habits, gestation length and sex have on brain size of extant ungulates. The analysis controlled for the effects of phylogeny and for random variability implicit in the data set. We tested the following groups of hypotheses: (1) Social brain hypothesis-gregarious species are more likely to have larger brains than non-gregarious species because the former are subjected to demanding and complex social interactions; (2) Ecological hypothesis-dietary habits impose challenging cognitive tasks associated with finding and manipulating food (foraging strategy); (3) Developmental hypotheses (a) energy strategy: selection for larger brains operates, primarily, on maternal metabolic turnover (i.e. gestation length) in relation to food quality because the majority of the brain's growth takes place in utero, and finally (b) sex hypothesis: females are expected to have larger brains than males, relative to body size, because of the differential growth rates of the soma and brain between the sexes. We found that, after adjusting for body mass, gregariousness and gestation length explained most of the variation in brain mass across the ungulate species studied. Larger species had larger brains; gregarious species and those with longer gestation lengths, relative to body mass, had larger brains than non-gregarious species and those with shorter gestation lengths. The effect of diet was negligible and subrogated by gestation length, and sex had no significant effect on brain size. The ultimate cause that could have triggered the co-evolution between gestation length and brain size remains unclear

Pichardo, M., 2005. Taxonomic revision of Central Mexican mammoths in Paleoindian sites
522. Anthropol. Anz. 63, 409-413.
Abstract: Central Mexican mammoth species taxonomy has been based on the quotient Molar length/Number of dental plates, which sorted three species, Mammuthus imperator, columbi and ?jeffersonii. New evidence from skull morphology sorts only two subspecies, M. columbi columbi and M. columbi felicis as being present during Paleoindian time

Roca, A.L., O'Brien, S.J., 2005. Genomic inferences from Afrotheria and the evolution of elephants
558. Curr. Opin. Genet. Dev. 15, 652-659.
Abstract: Recent genetic studies have established that African forest and savanna elephants are distinct species with dissociated cytonuclear genomic patterns, and have identified Asian elephants from Borneo and Sumatra as conservation priorities. Representative of Afrotheria, a superordinal clade encompassing six eutherian orders, the African savanna elephant was among the first mammals chosen for whole-genome sequencing to provide a comparative understanding of the human genome. Elephants have large and complex brains and display advanced levels of social structure, communication, learning and intelligence. The elephant genome sequence might prove useful for comparative genomic studies of these advanced traits, which have appeared independently in only three mammalian orders: primates, cetaceans and proboscideans

Roca, A.L., Georgiadis, N., O'Brien, S.J., 2005. Cytonuclear genomic dissociation in African elephant species
650. Nat. Genet. 37 , 96-100.
Abstract: African forest and savanna elephants are distinct species separated by a hybrid zone. Because hybridization can affect the systematic and conservation status of populations, we examined gene flow between forest and savanna elephants at 21 African locations. We detected cytonuclear dissociation, indicative of different evolutionary histories for nuclear and mitochondrial genomes. Both paternally (n = 205 males) and biparentally (n = 2,123 X-chromosome segments) inherited gene sequences indicated that there was deep genetic separation between forest and savanna elephants. Yet in some savanna locales distant from present-day forest habitats, many individuals with savanna-specific nuclear genotypes carried maternally transmitted forest elephant mitochondrial DNA. This extreme cytonuclear dissociation implies that there were ancient episodes of hybridization between forest females and savanna males, which are larger and reproductively dominant to forest or hybrid males. Recurrent backcrossing of female hybrids to savanna bulls replaced the forest nuclear genome. The persistence of residual forest elephant mitochondria in savanna elephant herds renders evolutionary interpretations based on mitochondrial DNA alone misleading and preserves a genomic record of ancient habitat changes.

Roth, G., Dicke, U., 2005. Evolution of the brain and intelligence
612. Trends Cogn Sci. 9, 250-257.
Abstract: Intelligence has evolved many times independently among vertebrates. Primates, elephants and cetaceans are assumed to be more intelligent than 'lower' mammals, the great apes and humans more than monkeys, and humans more than the great apes. Brain properties assumed to be relevant for intelligence are the (absolute or relative) size of the brain, cortex, prefrontal cortex and degree of encephalization. However, factors that correlate better with intelligence are the number of cortical neurons and conduction velocity, as the basis for information-processing capacity. Humans have more cortical neurons than other mammals, although only marginally more than whales and elephants. The outstanding intelligence of humans appears to result from a combination and enhancement of properties found in non-human primates, such as theory of mind, imitation and language, rather than from 'unique' properties

Shoshani, J., Tassy, P., 2005. Advances in proboscidean taxonomy & classification, anatomy & physiology, and ecology & behavior. Quaternary International 126-128, 5-20.
Abstract: With the addition of 13 new taxa, we recognized 175 species and subspecies of proboscideans, classified in 42 genera and 10 families. The three extant species are: forest African elephant (Loxodonta cyclotis), bush African elephant (L. africana), and Asian elephant (Elephas maximus, with three subspecies). Rigorous analysis of characters published or awaiting publication is imperative for better understanding of the cladistic relationships among currently recognized proboscideans. Here we focus on ''aquatic ancestry'' of Proboscidea, interordinal relationships within Placentalia, proboscidean taxonomy in general and South American in particular, anatomy and physiology and some ecological considerations. New taxa above the family level include sister taxa Mammutida and Elephantida, and Plesielephantiformes as a sister taxon to Elephantiformes. Neontological research is currently under way on the hyoid apparatus, lungs, brain, hearing, ecology and behavior. Topics for future research include: phylogenetic positions of anthracobunids, Moeritherium, tetralophodont gomphotheres, Stegolophodon and Stegodon, and intra-familial relationships among Loxodonta, Elephas and Mammuthus, and continuing studies on encephalization quotient. Certain anatomical features and functions (e.g., the hyoid apparatus that helps in food procurement, in production of infrasonic sounds, and in storing water to be used in time of stress) evolved about 25 million years ago, in time for diversification into new niches when grasses appeared in the landscape.

Sivasundaram, S., 2005. Trading knowledge: The East India Company's elephants in India and Britain. Historical Journal 48, 27-63.
Abstract: During the East India Company's rule of India, Britons observed the pervasiveness of elephants in local modes of warfare, hunting, trade, and religious symbolism. The colonizers appropriated this knowledge about elephants: for instance, in the taking-over of Mughal trade routes or Tipu Sultan's stables. What Indians knew about the elephant also fed into a metropolitan culture of anthropomorphism, exemplified in the celebrated shooting of the elephant Chuny in 1826. Anthropomorphic approaches to the elephant held by Britons worked alongside Sanskrit texts and Mughal paintings. These hybrid understandings gave way, by the mid-century to an allegedly objective and Christian science of animals, which could not be tainted by what was called pagan superstition. By using the elephant as a point of focus, this article urges the importance of popular traditions of colonial exchange in the emergence of science, and cautions against the reification of indigenous knowledge. The argument aims to show the strengths of a history of knowledge-making that is not focused on elites, the metropolis, or the periphery. A study of the uses of the elephant in colonialism also suggests the multiple and easily interchangeable meanings that animals could carry.

Stone, J., Telford, M., 2005. Fractal dimensions characterizing mammal teeth: A case study involving Elephantidae. Mammal Rev. 35, 123-128.
Abstract: 1. Dental features frequently have provided data for producing and deducing mammal taxonomies and phylogeny, yet quantitative or statistical analyses for describing intricacies that characterize tooth form are wanting. 2. A method for determining fractal dimensions D that characterize enamel ridges constituting occlusal surfaces for teeth in some mammal species is presented; D quantify complexity (i.e. convolution). The method is exemplified with an analysis that was conducted on teeth from the Family Elephantidae.

Stuart, A.J., 2005. The extinction of woolly mammoth ( Mammuthus primigenius) and straight-tusked elephant (Palaeoloxodon antiquus) in Europe. Quaternary International 126-28, 171-177.
Abstract: Together with several other megafaunal species in Northern Eurasia, Mammuthus primigenius and Palaeoloxodon antiquus became extinct in the Last Glacial-Interglacial cycle, but they had very different ecologies, times of extinction and `last stands' in different regions. The dramatic contraction in mammoth range ca. 12 kyr (uncalibrated C-14 chronology), after which known populations were confined to Northern Siberia (mainly Taymyr and Wrangel Island), correlates well with the extensive spread of trees in the Allerod phase of the Late Glacial Interstadial. The return of open steppe-tundra in the Younger Dryas cold phase, ca. 10.6-10 kyr, saw a limited re-expansion into NE Europe, followed by retraction and apparent extinction of mainland populations, which can be correlated with the marked loss of open habitats in the early Holocene. In contrast, at the end of the Last Interglacial, the retreat of P. antiquus to S. Europe, where it may have survived to ca. 50-34 kyr, can be linked to the loss of woodland habitats elsewhere. Although in both species, climate acting through vegetational changes evidently drove these range shifts, environmental change alone appears insufficient to account for extinctions. However, the possible role of human hunters is also still unclear.

Vidya, T.N., Fernando, P., Melnick, D.J., Sukumar, R., 2005. Population differentiation within and among Asian elephant (Elephas maximus) populations in southern India
675. Heredity 94, 71-80.
Abstract: Southern India, one of the last strongholds of the endangered Asian elephant (Elephas maximus), harbours about one-fifth of the global population. We present here the first population genetic study of free-ranging Asian elephants, examining within- and among-population differentiation by analysing mitochondrial DNA (mtDNA) and nuclear microsatellite DNA differentiation across the Nilgiris-Eastern Ghats, Anamalai, and Periyar elephant reserves of southern India. Low mtDNA diversity and 'normal' microsatellite diversity were observed. Surprisingly, the Nilgiri population, which is the world's single largest Asian elephant population, had only one mtDNA haplotype and lower microsatellite diversity than the two other smaller populations examined. There was almost no mtDNA or microsatellite differentiation among localities within the Nilgiris, an area of about 15,000 km2. This suggests extensive gene flow in the past, which is compatible with the home ranges of several hundred square kilometres of elephants in southern India. Conversely, the Nilgiri population is genetically distinct at both mitochondrial and microsatellite markers from the two more southerly populations, Anamalai and Periyar, which in turn are not genetically differentiated from each other. The more southerly populations are separated from the Nilgiris by only a 40-km-wide stretch across a gap in the Western Ghats mountain range. These results variably indicate the importance of population bottlenecks, social organization, and biogeographic barriers in shaping the distribution of genetic variation among Asian elephant populations in southern India

Zack, S.P., Penkrot, T.A., Bloch, J.I., Rose, K.D., 2005. Affinities of 'hyopsodontids' to elephant shrews and a Holarctic origin of Afrotheria
622. Nature 434, 497-501.
Abstract: Macroscelideans (elephant shrews or sengis) are small-bodied (25-540 g), cursorial (running) and saltatorial (jumping), insectivorous and omnivorous placental mammals represented by at least 15 extant African species classified in four genera. Macroscelidea is one of several morphologically diverse but predominantly African placental orders classified in the superorder Afrotheria by molecular phylogeneticists. The distribution of modern afrotheres, in combination with a basal position for Afrotheria within Placentalia and molecular divergence-time estimates, has been used to link placental diversification with the mid-Cretaceous separation of South America and Africa. Morphological phylogenetic analyses do not support Afrotheria and the fossil record favours a northern origin of Placentalia. Here we describe fossil postcrania that provide evidence for a close relationship between North American Palaeocene-Eocene apheliscine 'hyopsodontid' 'condylarths' (early ungulates or hoofed mammals) and extant Macroscelidea. Apheliscine postcranial morphology is consistent with a relationship to other ungulate-like afrotheres (Hyracoidea, Proboscidea) but does not provide support for a monophyletic Afrotheria. As the oldest record of an afrothere clade, identification of macroscelidean relatives in the North American Palaeocene argues against an African origin for Afrotheria, weakening support for linking placental diversification to the break-up of Gondwana

Zhu, M., Maslow, J.N., Mikota, S.K., Isaza, R., Dunker, F., Riddle, H., Peloquin, C.A., 2005. Population pharmacokinetics of pyrazinamide in elephants
564. J. Vet. Pharmacol. Ther. 28, 403-409.
Abstract: This study was undertaken to characterize the population pharmacokinetics (PK), therapeutic dose, and preferred route of administration for pyrazinamide (PZA) in elephants. Twenty-three African (Loxodonta africana) and Asian (Elephas maximus) elephants infected with or in contact with others culture positive for Mycobacterium tuberculosis were dosed under treatment conditions. PZA was dosed daily at 20-30 mg/kg via oral (fasting or nonfasting state) or rectal (enema or suppository) administration. Blood samples were collected 0-24 h postdose. Population PK was estimated using nonlinear mixed effect modeling. Drug absorption was rapid with T(max) at or before 2 h regardless of the method of drug administration. C(max) at a mean dose of 25.6 (+/-4.6) mg/kg was 19.6 (+/-9.5 microg/mL) for PZA given orally under fasting conditions. Under nonfasting conditions at a mean dose of 26.1 +/- 4.2 mg/kg, C(max) was 25% (4.87 +/- 4.89 microg/mL) and area under concentration curve (AUC) was 30% of the values observed under fasting conditions. Mean rectal dose of 32.6 +/- 15.2 mg/kg yielded C(max) of 12.3 +/- 6.3 microg/mL, but comparable AUC to PZA administered orally while fasting. Both oral and rectal administration of PZA appeared to be acceptable and oral dosing is preferred because of the higher C(max) and lower inter-subject variability. A starting dose of 30 mg/kg is recommended with drug monitoring between 1 and 2 h postdose. Higher doses may be required if the achieved C(max) values are below the recommended 20-50 microg/mL range

Agatsuma, T., Rajapakse, R.P., Kuruwita, V.Y., Iwagami, M., Rajapakse, R.C., 2004. Molecular taxonomic position of the elephant schistosome, Bivitellobilharzia nairi, newly discovered in Sri Lanka
745. Parasitol. Int. 53, 69-75.
Abstract: Bivitellobilharzia nairi (Mudaliar and Ramanujachar, 1945) Dutt and Srivastava, 1955 was first recorded in India. A number of adult worm specimens of this schistosome species were recovered from a domestic elephant, which died in 1999 in Sri Lanka. This is the first report of this schistosome from Sri Lanka. In the present study, in order to clarify the phylogenetic relationship with other species of schistosomes, sequences from the second internal transcribed spacer (ITS2) of the ribosomal gene repeat, part of the 28S ribosomal RNA gene (28S), and part of the mitochondrial cytochrome c oxidase subunit 1 (CO1) gene from B. nairi were analyzed. Two intraspecific variations were seen within 13 individuals in the ITS2 region. In the CO1 region of the mitochondrial DNA, there were four haplotypes in the nucleotide sequences and two haplotypes in the amino acid sequences. Phylogenetic analysis using the nuclear DNA showed that B. nairi was basal to all of species of the genus Schistosoma. The 28S tree also showed that the mammalian lineage was monophyletic. However, phylogenetic analysis using the mitochondrial DNA showed that B. nairi was nested within the genus Schistosoma. The taxonomical position for this species as well as the contradiction between the results from the nuclear and mitochondrial genes were discussed

Alter, S., 2004. Elephas maximus: A portrait of the Indian elephant. Harcourt Press.
Abstract: Review from Amazon: Mixing mythology and natural history, Stephen Alter lets readers share his lifelong love for the Indian elephant, Elephas maximus. While legends threaten to overwhelm facts in the tale, Alter has nonetheless presented an accurate portrait of his subject, true to centuries of Indian tradition. Beyond metaphors and fables, elephants occupy an important place in Sanskrit literature. Gajashastra, or "elephant science," was studied and recorded in several texts that are based on oral traditions.As much travelogue as science book, Elephas inextricably links the Indian elephant with the history of southern Asia itself. In pre-colonial India, elephants were wound up in religion and daily life; in modern times, the animals were first hunted then fetishized by Westerners. Alter reserves judgment on these issues, except to note that none of India's 20th-century history has been good for elephant populations, which are endangered or threatened nearly everywhere. He treks into parks and reserves, seeking out wild elephants and describing their awe-inspiring behaviors. The stories he uncovers along the way--of temple elephants, mysterious Elephanta  Island, seagoing elephants, and the god Ganesha--weave a spellbinding tale. --Therese Littleton

Buchanan, K.L., Goldsmith, A.R., 2004. Noninvasive endocrine data for behavioural studies: The importance of validation. Animal Behaviour 67, 183-185.
Abstract: There has been a substantial growth recently in the use of noninvasive methods to quantify hormone production, through the measurement of excreted hormones or hormone levels from saliva, sweat or hair (e.g.Wasser et al. 2000; Cook 2002; Pfeffer et al. 2002). These measures can quantify either current (e.g. Berg & Wynne-Edwards 2002; Maso et al. 2002) or past (e.g. Wasser et al. 2000; Ostner et al. 2002) levels of hormone production and the data can be used to determine the relations between a range of hormone levels and animal behaviour across taxa (Wasser et al. 2000). Such techniques have been used extensively to examine social stress (Goymann et al. 2001), the effects of environmental stress (Creel et al. 2002), reproductive cycles (Curtis et al. 2000) and social dominance (von Engelhardt et al. 2000; Langmore et al. 2002). They may have important applications in conservation science (Ishii 1999). There are several reasons why noninvasive methods of sampling are highly desirable. Importantly, animal suffering can potentially be reduced. In practical terms there are also several advantages: noninvasive methods allow samples to be obtained retrospectively, which represent average hormone production over a certain time frame, and the time spent handling the animal does not affect the levels obtained, which is advantageous for highly pulsatile hormones such as corticosteroids. In addition, the licensing constraints for noninvasive methods of sampling are less restrictive. However, such techniques also have disadvantages. In particular, faecal, hair or feather samples can indicate only average hormone levels over a considerable, and possibly unknown, period. Compared with plasma levels, noninvasive measures may result in a loss of sensitivity in any further analyses examining the relations between hormone levels and other variables (Shirtcliff et al. 2002). Furthermore, faecal samples in particular may not be available from known individuals a known amount of time after excretion, preventing reliable determination of individual hormone levels. It is also worth considering that while noninvasive sampling will not cause large increases in pulsatile 'stress' hormones as caused by capture and restraint, some increase may occur merely as a result of the presence of the sampler. In addition, there are a number of validation issues concerning the quantification of steroids from noninvasive samples which we outline below. Koren et al. (2002) documented a protocol for the extraction of testosterone and cortisol from hair obtained from the rock hyrax, Procavia capensis. They used this technique to quantify the levels of hormones contained in plucked hair samples, allowing hormone levels during the period of hair production to be determined, noninvasively. They found that the levels of testosterone extracted correlated positively with the dominance rank of male hyraxes. Although such methods are highly desirable, it is important to emphasize that all new methods of measuring levels of hormone production using hormone extracted from organic substrates should be appropriately validated, such that the limitations of the technique can be defined. This requires: (1) that the assay is validated for each new species and substrate and (2) that the extraction efficiency is determined for the target hormone in the species and substrate of interest. Although ready-made endocrine kits are provided with some data on the assay validation, the validation is relevant only for the species and substrate tested by the commercial supplier, generally in a limited range of biological media. It is essential to extend these validations for the species and substrate to which the kit is being applied. For example, a methanol extract of hair may contain substances that interfere with the assay procedure and thus would give misleading results.

Carter, A.M., Enders, A.C., 2004. Comparative aspects of trophoblast development and placentation. Reproductive Biology and Endocrinology 2.
Abstract: Based on the number of tissues separating maternal from fetal blood, placentas are classified as epitheliochorial, endotheliochorial or hemochorial. We review the occurrence of these placental types in the various orders of eutherian mammals within the framework of the four superorders identified by the techniques of molecular phylogenetics. The superorder Afrotheria diversified in ancient Africa and its living representatives include elephants, sea cows, hyraxes, aardvark, elephant shrews and tenrecs. Xenarthra, comprising armadillos, anteaters and sloths, diversified in South America. All placentas examined from members of these two oldest superorders are either endotheliochorial or hemochorial. The superorder Euarchontoglires includes two sister groups, Glires and Euarchonta. The former comprises rodents and lagomorphs, which typically have hemochorial placentas. The most primitive members of Euarchonta, the tree shrews, have endotheliochorial placentation. Flying lemurs and all higher primates have hemochorial placentas. However, the lemurs and lorises are exceptional among primates in having epitheliochorial placentation. Laurasiatheria, the last superorder to arise, includes several orders with epitheliochorial placentation. These comprise whales, camels, pigs, ruminants, horses and pangolins. In contrast, nearly all carnivores have endotheliochorial placentation, whilst bats have endotheliochorial or hemochorial placentas. Also included in Laurasiatheria are a number of insectivores that have many conserved morphological characters; none of these has epitheliochorial placentation. Consideration of placental type in relation to the findings of molecular phylogenetics suggests that the likely path of evolution in Afrotheria was from endotheliochorial to hemochorial placentation. This is also a likely scenario for Xenarthra and the bats. We argue that a definitive epitheliochorial placenta is a secondary specialization and that it evolved twice, once in the Laurasiatheria and once in the lemurs and lorises.

Carter, A.M., Enders, A.C., Kunzle, H., Oduor-Okelo, D., Vogel, P., 2004. Placentation in species of phylogenetic importance: The Afrotheria. Animal Reproduction Science 82-83, 35-48.
Abstract: Afrotheria, one of four mammalian superorders, comprises elephants, sea cows, hyraxes, aardvark, elephant shrews, tenrecs and golden moles. Their placentas either form an equatorial band or are discoid in shape. The interhemal region, separating fetal and maternal blood, is endotheliochorial in elephants, aardvark and possibly the sea cows, but hemochorial in the remaining orders. There is a secondary epitheliochorial placenta in elephant shrews while a similar structure in tenrecs erodes maternal tissues. Specialized hemophagous regions are a striking characteristic of some of these placentas yet absent in hyraxes, elephant shrews, and golden moles. It is possible that the common ancestor of the Afrotheria had an endotheliochorial placenta. Establishment of a hemochorial condition, as seen in rock hyraxes, elephant shrews, tenrecs, and golden moles, would be a more recent development. The elephant, manatee, and aardvark all have circumferential placentas. Thus the formation of a discoid placenta with a more or less extensive secondary placenta in elephant shrews and tenrecs would also be a derived state.

Carter, A.M., Enders, A.C., Kunzle, H., Oduor-Okelo, D., Vogel, P., 2004. Placentation in species of phylogenetic importance: the Afrotheria
705. Anim Reprod. Sci. 82-83, 35-48.
Abstract: Afrotheria, one of four mammalian superorders, comprises elephants, sea cows, hyraxes, aardvark, elephant shrews, tenrecs and golden moles. Their placentas either form an equatorial band or are discoid in shape. The interhemal region, separating fetal and maternal blood, is endotheliochorial in elephants, aardvark and possibly the sea cows, but hemochorial in the remaining orders. There is a secondary epitheliochorial placenta in elephant shrews while a similar structure in tenrecs erodes maternal tissues. Specialized hemophagous regions are a striking characteristic of some of these placentas yet absent in hyraxes, elephant shrews, and golden moles. It is possible that the common ancestor of the Afrotheria had an endotheliochorial placenta. Establishment of a hemochorial condition, as seen in rock hyraxes, elephant shrews, tenrecs, and golden moles, would be a more recent development. The elephant, manatee, and aardvark all have circumferential placentas. Thus the formation of a discoid placenta with a more or less extensive secondary placenta in elephant shrews and tenrecs would also be a derived state

Christiansen, P., 2004. Body size in proboscideans, with notes on elephant metabolism. Zoological Journal of the Linnean Society 140, 523-549.
Abstract: Mass estimates for a number of fossil proboscideans were computed using regression analyses on appendicular bones to body mass, for seven specimens of modern elephants, for which body masses had been recorded prior to death. The marked differences in physical proportions between extant Loxodonta and Elephas, implying substantial differences in body mass at any given shoulder height, were not present in their long bone parameters. Length and least circumferences proved to be the best parameters for prediction of body mass. Some extinct proboscideans, notably certain Mammuthus and Deinotherium, were much larger than extant elephants. Both the basal and the field metabolic rates of extant elephants are lower than predicted for a hypothetical mammal, in accordance with their body size and subsistence on low-quality foods. The feeding quantities often ascribed to extant wild elephants are exaggerated, and would in fact have sufficed to nourish much larger species.

Debruyne, R., 2004. How many species of elephants in Africa? Appraisal of an apparent controversy between molecular phylogenies. Cladistics-The International Journal Of The Willi Hennig Society 20, 80-81.
Abstract: Note: this is an abstract from the 22nd Annual Meeting of the Willi Hennig Society: The systematics of the African elephant, Loxodonta africana, has become an issue since the publication of the first mitochondrial sequence of a forest elephant (Barriel et al., 1999). The then acknowledged subspecies, L. africana cyclotis (Matschie, 1900), displayed a high level of divergence from the better-known savannah elephant (L. africana africana). Further molecular investigations have led authors to consider that forest and savnnah elephants consist of two distinct species (Comstock et al., 2002; Roca et al., 2001). They rely on an extensive sampling in Africa, though poorly documented for western populations and areas where both forms of elephants co-occur. They assayed the genetic diversity through nuclear markers, sequencing of four nuclear genes and microsatellites that converge to the recognition of two highly divergent clades they assume to match the systematic division. Conversely, the analysis of 593 bp of mtDNA from a broader sample produced results at odds (Eggert et al., 2002). The reciprocal monophyly of the two lneages is not retrieved and African elephants cluster in three main groups (western, central and south-eastern), whatever their systematics. However, these results are weakly supported. Here we address the validity of these inconsistent results by reanalyzing available data and sequencing 1960 bp of mtDNA for 43 African elephants (morphologically determined as 26 savannah and 17 forest elephants). The sequenced fragment spans cytochrome b gene, Thr and Pro tRNAs and Hyper-Variable Region 1 of the control region. Fifteen sequences of Hyper-Variable Region 2, Phe tRNA and 12S rRNA were also examined. Our phylogenetic analyses contradict both former studies. Firstly, they show that two unambiguously supported clades do exist within African elephants populations, Eggert's results (Eggert et al. 2002) being caused by an insufficient sampling of characters and a misleading attitude in the choice of the evolution model of their sequences. Secondly, this division into two main clades does not recover savannah and forest elephants as monophyletic taxa, so that they are not monophyletic species. While 21 sequences of savannah form a monophyletic group, the 5 remnants are apart within remote clades of forest elephant sequences. This conflicts with Roca et al.'s view (2001) that hybridization between the two forms can be neglected. Nevertheless, we confirm some former conclusions. Forest elephants display a stronger inner-structuration than savannah elephants. We acknowledge that some savannah elephants are mixed-up within forest elephant clades, but reciprocally, no forest elephant specimen depicted a typical savannah haplotype, so that the introgression of mitochondrial genome appears to be unidirectional. We assume that this extant phylogeographic pattern is the result of non-contemporary phenomena: (i) a prior isolation of forest elephants on the one hand and of savannah elephants on the other, leading to their molecular and morphological divergence, and (ii) secondary recurrent events of local admixture between the two forms, responsible for the geographic consistency of shared haplotypes between L. a. cyclotis and L. a. africana. Therefore we conclude that a high level of hybridization between the two forms of African elephant (with long-term effects on the genetic integrity of populations) precludes the recognition of two biological species.  Address: Museum Natl Hist Nat, Dept Hist Terre, CNRS, UMR 8569 8, F-75231 Paris, France

Debruyne, R., 2004. [Contribution of molecular phylogeny and morphometrics to the systematics of African elephants]
590. J. Soc. Biol. 198, 335-342.
Abstract: African elephants are conventionally classified as a single species: Loxodonta africana (Blumenbach 1797). However, the discovery in 1900 of a smaller form of the African elephant, spread throughout the equatorial belt of this land, has given rise to a debate over the relevance of a second species of elephant in Africa. The twentieth century has not provided any definite answer to this question. Actually, recent molecular analyses have sustained this issue by advocating either a division of forest elephants into a valid species, or their inclusion as a subspecies of L. africana. Our work initiated at the National Museum of Natural History of Paris provides new molecular (mitochondrial) and morphological (and morphometrical) evidence making it possible to propose a comprehensive phylogenetic hypothesis. It appears that there is no conclusive argument to keep forest elephants (cyclotis form) and savannah elephants (africana form) apart in two distinct species. A high level of mitochondrial introgression between the two forms, as well as a continuum in the morphology of the skulls of the two morphotypes rather suggests that, despite an ancient division, these two taxa freely interbreed wherever their ranges intersect. We thus adopt a conservative systematic position in considering these two forms as two subspecies, respectively: L. africana africana, the savannah elephant, and L. africana cyclotis, the forest elephant. We finally discuss the conservation topic in the light of this systematic framework

Elvin, M., 2004. The Retreat of the Elephants: An Environmental History of China. Yale University Press.
Abstract: Review from Nature 430, 505 - 506 (29 July 2004):                
Pity the poor elephants! Over more than 4,000 years they were gradually forced from living all over China to a few protected enclaves near the border with Burma. The main reason was the destruction of their habitat as humans cut down forests and introduced agriculture. Farmers found the dwindling elephant herds a nuisance, as crops were trampled and plundered. Others came to value elephants for military, transport and ceremonial purposes: their ivory was prized and their trunks became a gourmet delicacy. Elephant numbers shrank until they were little more than a memory for most Chinese. Mark Elvin uses the decline of the elephant as an allegory to illustrate the transformation of the Chinese environment to the end of pre-industrial times. Some of the same story can be seen in Africa today.

Elvin's book is not so much an environmental history of China as a collection of its fragments. With copious quotations from Chinese written sources of all kinds, he shows what happened in different places and why. Even if we can see from archaeology that comparable events took place elsewhere, only in China are there such written records, giving a unique account of how it felt to live through them. It was not always a pleasant or edifying process, and as usual the voices of those worst affected will never be heard.

In broad terms, the transformation of the Chinese environment, which was faster in some areas than others, had certain characteristics. First, deforestation made way for agriculture. There was then a bonanza as resources were exploited, species were lost and human numbers rose. This triggered the growth of towns, cities and states with social stratification, followed by increasing competition between them, with war as the spur and the environment sometimes used as a weapon. Better technology was mitigated by mismanagement of resources. Entrapment in limited local circumstances
left people vulnerable to change. Finally, there was a greater risk of social and economic collapse affecting society as a whole. Elvin shows the differences clearly in three areas: Jiaxing to the south of the Yangzi river; Guizhou in the south, where the Han people gradually displaced the indigenous Miao; and Zunhua in the mountainous northeast.

Everywhere, control of water was essential. 'Hydraulic despotism' may tell only part of the story, but communities and even states grew partly out of the need to manage this precious and sometimes capricious resource. The struggle to run irrigation systems, limit marine incursions, maintain banks and walls, undertake dredging, cope with floods and storms, and adapt to ever-changing weather patterns is as difficult today as it ever was. With huge populations dependent on particular systems, any change can become increasingly difficult to cope with.

The complexity of Chinese attempts to manage human effects on the environment is remarkable. Even more special are the Chinese beliefs and attitudes towards the environment that have existed over the millennia. Generalizations are bound to be misleading but, in general terms, the Chinese were driven, in Elvin's words, by a desire for rational mastery of the world. They had little hesitation in uprooting forests, redirecting and polluting rivers, destroying natural landscapes and giving political and military needs absolute priority. They had remarkable powers of organization, and ran projects far beyond European capacities at the time. But in doing so, the Chinese paid scant regard to the environment and unwittingly created many long-term problems.

On the other hand, the Chinese had a particularly sensitive respect for nature and natural beauty in all its forms. Even as forests were destroyed, individual trees were singled out for admiration. Heaven and Earth were closely linked, and the line between the natural and the supernatural was blurred. There was a confluence of matter leading to energy, and energy leading to life, each a product of Bright Force and Dark Force. Dragons and spirits were sometimes seen above the surface in thunder and lightning, and sometimes below it in earthquakes. They formed part of a living world that
sustained and punished humans. They even related the behaviour of the weather to human activity, so there was morality in meteorology.

In such a world, it was crucial to divine what the invisible forces felt or did. This could involve sacrificing animals or humans, or burning cracks in the shoulder blades of mammals or the undershells of turtles. In Shang times, such practices had political significance as the ruler was the intermediary between the visible and the invisible world. This was also true in other epochs when the apparatus of authority was given almost divine attributes.

It is as difficult for us to enter into this mental cosmology as into that of our own ancestors in pre-scientific times. Elvin shows that searching for observable and verifiable facts about the world, and putting them to use in programmes of thought, was almost entirely alien to the Chinese. As a result, the shock of change was more abrupt in China than it was in Europe, where the scientific revolution began earlier. Traces of the old thinking may have survived Mao Zedong and persist in fundamental ways today.

The Retreat of the Elephants is not an easy book to read. Some of the quotations seem scarcely relevant, and the whole text could have been usefully pruned. At the end there is an unilluminating venture into equations, as if sustainability could be reduced to an algorithm. Yet taken as a whole, the book is a fascinating, scholarly miscellany of stories, poetry and ideas from the history of the longest continuous civilization on Earth. The relationship of that civilization with its fragile and often tortured surroundings contains lessons for others - particularly at a time when industrial society in China, as elsewhere, is pressing harder than ever on the environment. This will be a source book, elephants and all, for generations to come.

Enders, A.C., Carter, A.M., 2004. What can comparative studies of placental structure tell us?--A review
739. Placenta 25 Suppl A , S3-S9.
Abstract: The diversity of placental structures in Eutherian mammals is such that drawing generalizations from the definitive forms is problematic. There are always areas of reduced interhaemal distance whether the placenta is epitheliochorial, synepitheliochorial, endotheliochorial or haemochorial. However, the thinning may be achieved by different means. The presence of a haemophagous area as an iron transport facilitator is generally associated with endotheliochorial placentae but is also found in sheep and goats (synepitheliochorial) and in tenrecs and hyaenas (haemochorial). Although similar chorioallantoic placentae are found within families, structure begins to diverge at the ordinal level and there is little correlation at the supraordinal level of phylogeny. Differences in formation and function of the yolk sac provide additional variation. There would appear to be considerable adaptive pressure for development or retention of the haemochorial type of chorioallantoic placenta. This type of placenta has several possible drawbacks including more ready passage of fetal cells to the maternal organism and, should the haemochorial condition be achieved early, oxidative stress. At any rate no animal larger than the human and gorilla has this type of placenta. The endotheliochorial condition is found in animals as large as the bears, manatee and elephants. In addition to the ungulates, the epitheliochorial condition is present in the largest animals with the longest gestation periods, the whales. Considering the length of time since the early stages of mammalian evolution, it is probable that few unmodified structural features are present in any currently surviving mammal. Nevertheless, more complete studies of divergent types of mammalian placenta should help our understanding of mammalian interrelationships as well as placental function

Greenwood, A.D., Englbrecht, C.C., MacPhee, R.D., 2004. Characterization of an endogenous retrovirus class in elephants and their relatives
667. BMC. Evol. Biol. 4, 38.
Abstract: BACKGROUND: Endogenous retrovirus-like elements (ERV-Ls, primed with tRNA leucine) are a diverse group of reiterated sequences related to foamy viruses and widely distributed among mammals. As shown in previous investigations, in many primates and rodents this class of elements has remained transpositionally active, as reflected by increased copy number and high sequence diversity within and among taxa. RESULTS: Here we examine whether proviral-like sequences may be suitable molecular probes for investigating the phylogeny of groups known to have high element diversity. As a test we characterized ERV-Ls occurring in a sample of extant members of superorder Uranotheria (Asian and African elephants, manatees, and hyraxes). The ERV-L complement in this group is even more diverse than previously suspected, and there is sequence evidence for active expansion, particularly in elephantids. Many of the elements characterized have protein coding potential suggestive of activity. CONCLUSIONS: In general, the evidence supports the hypothesis that the complement had a single origin within basal Uranotheria

Hoffmann, J.N., Montag, A.G., Dominy, N.J., 2004. Meissner corpuscles and somatosensory acuity: the prehensile appendages of primates and elephants
669. Anat. Rec. A Discov. Mol. Cell Evol. Biol. 281, 1138-1147.
Abstract: Meissner corpuscles (MCs) are specialized mechanoreceptors located exclusively in the papillae of glabrous skin. They are confined largely to cutaneous pads of the extremities and respond to transient, phasic, or vibratory stimuli. Though absent in most eutherian taxa, MCs are reported in all primates studied, being most developed in modern humans. The location of MCs between the internal ridges of the epidermis indicates they are well situated to detect friction or deformation at the external surface. Accordingly, MCs are hypothesized to provide primates generally with an enhanced tactile perception. However, the selective pressures favoring greater somatosensory acuity in primates are seldom considered. Interestingly, primate digital dexterity varies greatly. In general, dexterity improves with the extent to which foraging requires food manipulation or textural evaluation. This observation implies that MC density could vary accordingly. Here we report on the density of MCs in five anthropoid taxa selected to represent diverse dietary regimes. Results show that greater MC density correlates with the extent to which primates are frugivorous; however, locomotor and/or phylogenetic effects cannot be discounted

Repin, V.E., Taranov, O.S., Ryabchikova, Tikhonov, A.N., Pugachev, V.G., 2004. Sebaceous glands of the woolly mammoth, Mammothus primigenius Blum: histological evidence
651. Dokl. Biol. Sci. 398, 382-384.

Sarma, K.K., 2004. Extraction of decayed tusk in elephants. Indian Veterinary Journal 81, 812-814.
Abstract: Case history of dental pulp decay in eight male Asian elephants is discussed. Causes of injury and infection, pathological process and clinical signs are elaborated. Treatment of the cases by extraction of the decayed tusks, anaesthetic management, operative procedure, post operative care and the outcome of treatment has been discussed.

Shimoyama, M., Morimoto, S., Ozaki, Y., 2004. Non-destructive analysis of the two subspecies of African elephants, mammoth, hippopotamus, and sperm whale ivories by visible and short-wave near infrared spectroscopy and chemometrics
721. Analyst 129, 559-563.
Abstract: Visible (VIS) and short-wave near infrared (SW-NIR) spectroscopy was used for non-destructive analysis of ivories. VIS-SW-NIR (500-1000 nm) spectra were measured in situ for five kinds of ivories, that is two subspecies of African elephants, mammoth, hippopotamus, and sperm whale. Chemometrics analyses were carried out for the spectral data from 500 to 1000 nm region. The five kinds of ivories were clearly discriminated from each other on the scores plot of two principal components (PCs) obtained by principal component analysis (PCA). It was noteworthy that the ivories of the two subspecies of African elephants were discriminated by the scores of PC 1. The loadings plot for PC 1 showed that the discrimination relies on the intensity changes in bands due to collagenous proteins and water interacting with proteins. It was found that the scores plot of PC 2 is useful to distinguish between the ivories of the two subspecies of African elephants and the other ivories. We also developed a calibration model that predicted the specific gravity of five kinds of ivories from their VIS-SW-NIR spectral data using partial least squares (PLS)-1 regression. The correlation coefficient and root mean square error of cross validation (RMSECV) of this model were 0.960 and 0.037, respectively

Stahl, A.B., Stahl, P.W., 2004. Ivory production & consumption in Ghana in the early second millennium AD. Antiquity 78, 86-101.
Abstract: In the eighteenth to nineteenth century West Africa was the scene of the infamous Atlantic trade in ivory and slaves. The authors' researches show a different situation in the fourteenth to seventeenth centuries, when the people of Ghana were engaged in the indigenous procurement, manufacture and trade in ivory with neighbours across the Sahara. Address: Stahl AB, SUNY Binghamton, Dept Anthropol, Binghamton, NY 13902 USA.

Stringfield, C.E., Oh, P., Granich, R., Scott, J., Sun, B., Joseph, M., Flood, J., Sedgwick, C.J. Epidemiologic investigation of a Mycobacterium tuberculosis infection of multiple animal species in a metropolitan zoo. 2004 PROCEEDINGS AAZV, AAWV, WDA JOINT CONFERENCE.  46-48. 2004.
Ref Type: Conference Proceeding
Abstract: From 1997 to 2000, six cases of Mycobacterium tuberculosis (TB) infection were diagnosed in three species of animals at, or recently originating from, the Los Angeles Zoo. Restriction fragment length polymorphism (RFLP) analysis showed that five of six animal isolates shared an identical IS6110 pattern, with the sixth differing only by one additional band. A multiinstitutional epidemiologic investigation was conducted to identify and interrupt possible transmission among the animal cases, and to screen personnel for active TB infection and TB skin-test conversion.
Animal Cases
In April and October of 1994, Asian elephant (Elephas maximus) #1 and Asian elephant #2 arrived at the Los Angeles Zoo from a private elephant facility where they had lived together. They were housed together at the zoo until November of 1996 when elephant #2 was returned to the facility for several months before transfer to another zoo. In the spring of 1997, Elephant #1 (30 yr old) died of salmonellosis, with M. tuberculosis found in granulomatous lymph node lesions from the thoracic and abdominal cavities, and Elephant #2 (30 yr old) was found to have a positive trunk wash culture for M. tuberculosis. In July of 1998, one of a closed herd of three Rocky Mountain goats (Oreamnos americanus) consisting of a sire and two offspring, died of pulmonary M. tuberculosis at 6 yr of age. The goat's asymptomatic herdmates were screened and had negative chest radiographs and tracheal wash cultures, but one of the two goats was positive on tuberculin skin-test. In October of 1998, a clinically normal Black rhinocerus (Diceros bicornis) was diagnosed with Mycobacerium tuberculosis after a positive skin test and nasal wash culture. In the winter of 1998, the two remaining goats were evaluated again with negative chest radiographs and tracheal wash cultures. However, 1 yr later, both were humanely euthanatized at 8 and 12 yr of age due to clinical evidence of tuberculosis on chest radiographs (both animals), and active clinical signs in one (neither were able to be orally treated). In January of 2001, a rhino was humanely euthanatized after a protracted illness that was nonresponsive to aggressive treatment. The rhino was found to have severe multifocal hemosiderosis and atypical mycobacterial infection in her lungs, with no M. tuberculosis  cultured. This animal had been treated with oral Isoniazid and Rifampin for 1 yr, cultured routinely, and was never culture positive again.
 Epidemiologic Investigation
Investigators examined medical and location histories of the affected animals, animal handling practices, health-care procedures, and performed an infection control assessment of the animal compounds and health-care facilities (including measuring air flow in the compounds by smoke testing). We conducted a review of zoo employee medical records for evidence of TB symptoms, tuberculin skin-test results, and chest radiograph information. A list of current and former employees was cross-matched with reported TB cases in the California state registry from 1985 to 2000. As part of the annual occupational health screening in June of 2000, zoo employees underwent questioning regarding TB symptoms, received tuberculin skin tests, and completed a questionnaire on medical history, job type, and history of contact with the infected animals.
Epidemiologic Findings
No common cross-species contact outside the animal compounds and no contact with an infectious human were found. The distance at which the public was kept from the animals and the distance of the compounds from each other (the elephant compound was 27 meters from the rhino compound and the goat compound was 90 m from both) suggests that direct transmission was unlikely. No active TB cases in humans were found, and no matches were found in the database of reporte d cases. The RFLP analysis of this strain of M. tuberculosis matched that of three elephants with which #1 and #2 were housed at a private elephant facility from September of 1993-February of 1994.1 We hypothesize that elephants #1 and #2 were infected at the private facility and were shipped with latent M. tuberculosis infection in 1994, subsequently infecting the black rhino and Mountain goats at the Los Angeles Zoo.
Of interest, animal caretaking and animal contact were not associated with a positive tuberculin skin-test, while groundskeepers were found to have an increased risk of tuberculin skin-test conversion compared with other job categories. Employees attending the elephant necropsy and employees who trained elephants were more likely to have tuberculin skin-test conversion than those who did not.
Conclusion
This is the first documented human and veterinary epidemiologic investigation of Mycobacterium tuberculosis affecting multiple species in a zoo. ² No evidence of transmission from humans to animals or active infections in humans were found. Genotyping evidence strongly suggests transmission from one species to another, although no evidence of transmission was discovered. Human tuberculin skin-test conversions associated with the elephants were most likely due to lack of respiratory protection for these employees when the risk of TB infection was not known. The finding that groundskeepers and not animal handlers were associated with a higher risk of tuberculin skin-test conversion was surprising, and we hypothesized that this may have to do with groundskeepers as a group being more likely to have
been born outside of the United States.
Control measures to eliminate the spread of disease to people and animals were undertaken immediately and throughout this outbreak, and no further cases of M. tuberculosis have been diagnosed at the zoo in the past 3 yr despite ongoing surveillance. Four elephants and three rhinos that had direct contact with the infected animals remain TB negative by trunk and nasal wash culture methods as outlined by the USDA for elephant TB surveillance. Methods of indirect transmission in mammalian zoo species and causes of variability in infection and morbidity within and among species warrant further investigation. Ongoing vigilance, occupational health programs and infection control measures in potentially exposed animals are recommended to prevent ongoing transmission of M. tuberculosis in zoo settings.
Acknowledgments
The authors thank the Animal Care and Animal Health staff of the Los Angeles Zoo who cared so well for these animals, and the veterinarians (including consulting pathologists), technicians, and medical records staff who collected, analyzed, and organized the clinical data. We could not have performed this evaluation without Sue Thisdell, Safety Officer at the Los Angeles Zoo; Jothan Staley and Donna Workman-Malcom of the City of Los Angeles Occupational Health Services Division; Lee Borenstein, Elenor Lehnkering, Patrick Ryan, Jeanne Soukup, and Annette Nita of the Los Angeles County Department of Health Services; and Diana Whipple for her RFLP expertise.
LITERATURE CITED
1. Mikota, S.K., L. Peddie, J. Peddie, R. Isaza, F. Dunker, G. West, W. Lindsay, R.S.Larsen, M. D. Salman, D. Chatterjee, J. Payeur, D. Whipple, C. Thoen, D. Davis, C. Sedgwick, R.J. Montali, M. Ziccardi, J. Maslow. 2001. Epidemiology and diagnosis of Mycobacterium tuberculosis in captive asian elephants (Elephas maximus). J. Zoo Wildl. Med. 32: 1-16.
2. Oh, P., R. Granich, J. Scott, B. Sun, M. Joseph, C. Stringfield, S. Thisdell, J. Staley, D. Workman-Malcolm, L. Borenstein, E. Lehnkering, P. Ryan, J. Soukup, A.Nitta, J. Flood. 2002. Human exposure following Mycobacterium tuberculosis infection of multiple animal species in a metropolitan zoo. Emerging Infectious Diseases. 8 (11): 1290-1293.orte

Suarez, R.K., Darveau, C.A., Childress, J.J., 2004. Metabolic scaling: a many-splendoured thing
656. Comp Biochem. Physiol B Biochem. Mol. Biol. 139, 531-541.
Abstract: Animals at rest and during exercise display rates of aerobic metabolism, VO2, that represent mainly the sum of mitochondrial respiration rates in various organs. The relative contributions of these organs change with physiological state such that internal organs such as liver, kidney and brain account for most of the whole-body VO2 at rest, while locomotory muscles account for >90% of the maximum rate, VO2max, during maximal aerobic exercise. Mechanisms that regulate VO2 are complex and the relative importance of each step in a series, estimated by metabolic control analysis, depends upon the level of biological organization under consideration as well as physiological state. Despite this complexity, prominent single-cause models propose that metabolic rates are supply-limited and that the scaling of supply systems provides a sufficient explanation for the allometric scaling of metabolism. We argue that some assumptions, as well as current interpretations of the meaning (or consequences) of these constraints are flawed, i.e., elephants do not have lower mass-specific basal or maximal rates of aerobic metabolism because their mitochondria are more supply-limited than those of shrews. Animals do not violate the laws of physics, and the allometric scaling of supply systems would be expected, to some extent, to be matched by capacities for (and rates of) energy expenditure. But life is not so simple. Animals are so diverse that to do justice to metabolic scaling, it is also necessary to consider the scaling of energy expenditure. It is by doing so that models of metabolic scaling can be consistent with current paradigms in metabolic regulation and accommodate the range of inter- and intraspecific exponents found in nature. The "allometric cascade," a first attempt at such an accounting, was a source of great satisfaction to Peter Hochachka. It was the last door that he helped open to comparative physiologists before he said goodbye

Taranov, O., Ryabchikova, E., Tikhonov, A., Pugachev, V., Repin, V., 2004. Sebaceous glands of woolly mammoth (the histological evidence). Proceedings of the Russian Academy of Sciences, ser. biol. , 398, 139-141.

Tefera, M., 2004. Recent evidence of animal exploitation in the Axumite epoch, 1st-5th centuries AD
743. Trop. Anim Health Prod. 36, 105-116.
Abstract: This study reports evidence of animal exploitation during the Axumite era based on a survey of architectural features, rock art and artefacts recovered from the archaeological site at Axum, northern Ethiopia. Animals and agricultural tools were identified from materials not previously examined. Pottery, rock art and animal remains revealed a range of zoological species. Agricultural implements and sacrificial vessels also provided indirect evidence of animal exploitation. It is concluded that, in addition to plough-based agriculture, the hunting of large wild animals, such as elephants and lions, and the exploitation of domestic cattle, small ruminants and poultry were part of the Axumite subsistence regime. Although it is difficult to reconstruct an overall subsistence pattern based on this study alone, the physical and biological evidence suggests that the Axumites practised combined cultivation and animal herding. Further investigation is required to study the types and breeds of animals and their distribution in time and space

Clauss, M., Frey, R., Kiefer, B., Lechner-Doll, M., Loehlein, W., Polster, C., Rossner, G.E., Streich, W.J., 2003. The maximum attainable body size of herbivorous mammals: morphophysiological constraints on foregut, and adaptations of hindgut fermenters. Oecologia 136, 14-27.
Abstract: An oft-cited nutritional advantage of large body size is that larger animals have lower relative energy requirements and that, due to their increased gastrointestinal tract (GIT) capacity, they achieve longer ingesta passage rates, which allows them to use forage of lower quality. However, the fermentation of plant material cannot be optimized endlessly; there is a time when plant fibre is totally fermented, and another when energy losses due to methanogenic bacteria become punitive. Therefore, very large herbivores would need to evolve adaptations for a comparative acceleration of ingesta passage. To our knowledge, this phenomenon has not been emphasized in the literature to date. We propose that, among the extant herbivores, elephants, with their comparatively fast passage rate and low digestibility coefficients, are indicators of a trend that allowed even larger hindgut fermenting mammals to exist. The limited existing anatomical data on large hindgut fermenters suggests that both a relative shortening of the GIT, an increase in GIT diameter, and a reduced caecum might contribute to relatively faster ingesta passage; however, more anatomical data is needed to verify these hypotheses. The digestive physiology of large foregut fermenters presents a unique problem: ruminant-and nonruminant-forestomachs were designed to delay ingesta passage, and they limit food intake as a side effect. Therefore, with increasing body size and increasing absolute energy requirements, their relative capacity has to increase in order to compensate for this intake limitation. It seems that the foregut fermenting ungulates did not evolve species in which the intake-limiting effect of the foregut could be reduced, e.g. by special bypass structures, and hence this digestive model imposed an intrinsic body size limit. This limit will be lower the more the natural diet enhances the ingesta retention and hence the intake-limiting effect. Therefore, due to the mechanical characteristics of grass, grazing ruminants cannot become as big as the largest browsing ruminant. Ruminants are not absent from the very large body size classes because their digestive physiology offers no particular advantage, but because their digestive physiology itself intrinsically imposes a body size limit. We suggest that the decreasing ability for colonic water absorption in large grazing ruminants and the largest extant foregut fermenter, the hippopotamus, are an indication of this limit, and are the outcome of the competition of organs for the available space within the abdominal cavity. Our hypotheses are supported by the fossil record on extinct ruminant/tylopod species which did not, with the possible exception of the Sivatheriinae, surpass extant species in maximum body size. In contrast to foregut fermentation, the GIT design of hindgut fermenters allows adaptations for relative passage acceleration, which explains why very large extinct mammalian herbivores are thought to have been hindgut fermenters.  Institute of Animal Physiology, Physiological Chemistry and Animal Nutrition, Ludwig-Maximilians-University, Veterinaerstrasse 13, 80539, Munich, Germany. clauss@tiph.vetmed.uni-muenchen.de

Cristoffer, C., Peres, C.A., 2003. Elephants versus butterflies: the ecological role of large herbivores in the evolutionary history of two tropical worlds. Journal of Biogeography 9, 1357-1380.
Abstract: Aim Large herbivores have important effects upon Paleotropical ecosystems, but attain much lower biomass densities in the Neotropics. We assess how this difference in herbivore activity has generated different ecological and evolutionary trajectories in the New and Old World tropics. We also propose an explanation for how the greater biomass density in the Old World came about. Location Data were compiled primarily from moist tropical forests, although more of the relevant information to address most of our hypotheses was available from the mainland areas of Africa, Asia, and South America than elsewhere. Methods We gleaned data from published information and personal communication. We compared body masses and a variety of other types of information for the New- and Old-World tropics. We proposed that interhemispheric differences exist in a variety of processes, including herbivory, frugivory, and flower visitation. We erected hypotheses and evaluated them qualitatively, and, when information was available, tested them using simple ratios of species in various taxonomic and trophic categories. To make the comparisons more meaningful, we specified appropriate data selection criteria. Results A general pattern of differences emerges from this review.  Compared with Neotropical forests, the much greater biomass densities of large herbivores in Paleotropical forests are associated with a lesser diversity of small herbivores, different hunting methods used by indigenous humans, larger arboreal vertebrates, larger fruits, different patterns of fruit and flower dispersion in space and time, a lesser abundance of most types of reproductive plant parts, and other features. The existence of a species-rich fauna of large herbivores in the pre-Holocene Neotropical rain forest was not supported. Main conclusions: The potential for large herbivores to cause functional differences between the New and Old World tropical forests has been virtually unexplored, despite the well-known importance of large herbivores in the Old World tropics. The evaluations of our hypotheses suggest that the abundance of large herbivores in the Old World tropics has launched it onto a different evolutionary trajectory than that of the NewWorld tropics. The relevant evidence, although scanty, suggests that the interhemispheric ecological differences are not an artefact of recent megafaunal extinctions in the New World. Recent human activities have, however, reduced population sizes of large wild herbivores in the Old World, and increased population sizes of livestock. This has likely created a rather homogeneous, anthropogenic selection pressure that tends to erase the evolutionary differences between the two tropical worlds.

Debruyne, R., Barriel, V., Tassy, P., 2003. Mitochondrial cytochrome b of the Lyakhov mammoth (Proboscidea, Mammalia): new data and phylogenetic analyses of Elephantidae. Mol Phylogenet Evol 26, 421-434.
Abstract: The phylogenetic relationships between recent Elephantidae (Proboscidea, Mammalia), that is to say extant elephants (Asian and African) and extinct woolly mammoth, have remained unclear to date. The prevailing morphological scheme (mammoth grouped with Asian elephant) is either supported or questioned by the molecular results. Recently, the monophyly of woolly mammoths on mitochondrial grounds has been demonstrated (Thomas, et al., 2000), but it conflicts with previous studies (Barriel et al., 1999; Derenko et al., 1997). Here, we report the partial sequencing of two mitochondrial genes: 128 bp of 12S rDNA and 561 bp of cytochrome b for the Lyakhov mammoth, a 49,000-year-old Siberian individual. We use the most comprehensive sample of mammoth (11 sequences) to determine whether the sequences achieved by former studies were congruent or not. The monophyly of a major subset of mammoths sequences (including ours) is recovered. Such a result is assumed to be a good criterion for ascertaining the origin of ancient DNA. Our sequence is incongruent with that of Yang et al. (1996), though obtained for the same individual. As far as the latter sequence is concerned, a contamination by non-identified exogenous DNA is suspected. The robustness and reliability of the sister group relation between Mammuthus primigenius and Loxodonta africana are examined: down-weighting saturated substitutions has no impact on the topology; analyzing data partitions proves that the support of this clade can be assigned to the most conservative phylogenetic signal; insufficient taxonomic and/or characters sampling contributed to former discordant conclusions. We therefore assume the monophyly of "real mammoth sequences" and the (Mammuthus, Loxodonta) clade. Laboratoire de Paleontologie, CNRS UMR 8569, Museum National d'Histoire Naturelle, 8 Rue Buffon, F-75005 Paris, France. debruyne@mnhn.fr

Debruyne, R., Van Holt, A., Barriel, V., Tassy, P., 2003. Status of the so-called African pygmy elephant (Loxodonta pumilio (NOACK 1906)): phylogeny of cytochrome b and mitochondrial control region sequences. C R Biol 326, 687-697.
Abstract: Among the African elephants, it has been unanimously acknowledged that the forest elephants (cyclotis form) are peculiar, so that they have been elevated to the specific rank. The development of molecular analyses of extant Loxodonta has only focused on two forms yet: the savannah form (africana) and the forest form (cyclotis), disregarding the so-called pygmy elephants (pumilio or fransseni) the systematic status of which has been debated since their discovery. Therefore, we have sampled nine dwarfed-labelled specimens in collection and eight specimens of typical forest elephants that we compared to three savannah elephants and two Asian elephants. Because of the degraded nature of the nuclear DNA content in bone samples of old specimens, we assayed mitochondrial markers; 1961 bp of the mitochondrial genome were sequenced (over a continuous range spanning the cytochrome b gene, tRNA Thr, tRNA Pro, hypervariable region 1 and central conserved region of the control region). Pumilio and cyclotis are not sister-taxa: the phylogenetic analyses rather account for the inclusion of the so-called pygmy elephants within a monophyletic group of forest elephants sensu lato. The internal structure of this clade reveals to depend on isolation and remoteness between populations, characteristics that may have been extensively influenced by climatic variations during the Quaternary period. We conclude that the specific taxon Loxodonta pumilio (or Loxodonta fransseni) should be abandoned. FR 1541 CNRS, Service de systematique moleculaire, Museum national d'histoire naturelle, 43, rue Cuvier, 75005 Paris, France. debruyne@mnhn.fr

Dubois, J.Y., Ursing, B.M., Kolkman, J.A., Beintema, J.J., 2003. Molecular evolution of mammalian ribonucleases 1. Mol Phylogenet Evol. 27, 453-463.
Abstract: There have been many studies on the chemistry of mammalian pancreatic ribonucleases (ribonucleases 1), but the functional biology of this family of homologous proteins is still largely unknown. Many studies have been performed on the molecular evolution and properties of this enzyme from species belonging to a large number of mammalian taxa, including paralogous gene products resulting from recent gene duplications. Novel ribonuclease 1 sequences were determined for three rodent species (gundi, brush-tailed porcupine, and squirrel), rabbit, a fruit bat, elephant, and aardvark, and the new sequences were used for deriving most parsimonious networks of ribonucleases from different mammalian orders, including earlier determined nucleotide sequences and also a larger set of protein sequences. Weak support for interordinal relationships were obtained, except for an Afrotheria clade containing elephant and aardvark. Results of current analyses and also those obtained 20 years ago on amino acid sequences confirm conclusions derived recently from larger data sets of other molecules. Several examples of recent gene duplications in ribonucleases 1 are discussed, with respect to illustrate the concepts of orthology and paralogy. Previously evidence was presented for extensive parallelism between sequence regions with attached carbohydrate (about one quarter of the molecule) of unrelated species with cecal digestion (pig and guinea pig). These features are also present in the sequences of elephant and fruit bat, species with cecal digestion, but with a very low ribonuclease content in their pancreas. Department of Biochemistry, University of Groningen, Groningen, The Netherlands. dubois@farm.rug.nl

Eggert, L.S., Eggert, J.A., Woodruff, D.S., 2003. Estimating population sizes for elusive animals: the forest elephants of Kakum National Park, Ghana. Mol Ecol 12, 1389-1402.
Abstract: African forest elephants are difficult to observe in the dense vegetation, and previous studies have relied upon indirect methods to estimate population sizes. Using multilocus genotyping of noninvasively collected samples, we performed a genetic survey of the forest elephant population at Kakum National Park, Ghana. We estimated population size, sex ratio and genetic variability from our data, then combined this information with field observations to divide the population into age groups. Our population size estimate was very close to that obtained using dung counts, the most commonly used indirect method of estimating the population sizes of forest elephant populations. As their habitat is fragmented by expanding human populations, management will be increasingly important to the persistence of forest elephant populations. The data that can be obtained from noninvasively collected samples will help managers plan for the conservation of this keystone species. Ecology, Behavior and Evolution Section, Division of Biological Sciences, University of California, San Diego 92093-0116, USA. lori_eggert@hotmail.com

Fernando, P., Vidya, T.N., Payne, J., Stuewe, M., Davison, G., Alfred, R.J., Andau, P., Bosi, E., Kilbourn, A., Melnick, D.J., 2003. DNA analysis indicates that Asian elephants are native to Borneo and are therefore a hgh priority for conservation. PLoS Biol. 1, E6 (Epub 2003 Aug 18).
Abstract: The origin of Borneo's elephants is controversial. Two competing hypotheses argue that they are either indigenous, tracing back to the Pleistocene, or were introduced, descending from elephants imported in the 16th-18th centuries. Taxonomically, they have either been classified as a unique subspecies or placed under the Indian or Sumatran subspecies. If shown to be a unique indigenous population, this would extend the natural species range of the Asian elephant by 1300 km, and therefore Borneo elephants would have much greater conservation importance than if they were a feral population. We compared DNA of Borneo elephants to that of elephants from across the range of the Asian elephant, using a fragment of mitochondrial DNA, including part of the hypervariable d-loop, and five autosomal microsatellite loci. We find that Borneo's elephants are genetically distinct, with molecular divergence indicative of a Pleistocene colonisation of Borneo and subsequent isolation. We reject the hypothesis that Borneo's elephants were introduced. The genetic divergence of Borneo elephants warrants their recognition as a separate evolutionary significant unit. Thus, interbreeding Borneo elephants with those from other populations would be contraindicated in ex situ conservation, and their genetic distinctiveness makes them one of the highest priority populations for Asian elephant conservation. Center for Environmental Research and Conservation, Columbia University, New York, New York, United States of America. pf133@columbia.edu

Fronicke, L., Wienberg, J., Stone, G., Adams, L., Stanyon, R., 2003. Towards the delineation of the ancestral eutherian genome organization: comparative genome maps of human and the African elephant (Loxodonta africana) generated by chromosome painting. Proc R Soc Lond B Biol Sci. 270, 1331-1340.
Abstract: This study presents a whole-genome comparison of human and a representative of the Afrotherian clade, the African elephant, generated by reciprocal Zoo-FISH. An analysis of Afrotheria genomes is of special interest, because recent DNA sequence comparisons identify them as the oldest placental mammalian clade. Complete sets of whole-chromosome specific painting probes for the African elephant and human were constructed by degenerate oligonucleotide-primed PCR amplification of flow-sorted chromosomes. Comparative genome maps are presented based on their hybridization patterns. These maps show that the elephant has a moderately rearranged chromosome complement when compared to humans. The human paint probes identified 53 evolutionary conserved segments on the 27 autosomal elephant chromosomes and the X chromosome. Reciprocal experiments with elephant probes delineated 68 conserved segments in the human genome. The comparison with a recent aardvark and elephant Zoo-FISH study delineates new chromosomal traits which link the two Afrotherian species phylogenetically. In the absence of any morphological evidence the chromosome painting data offer the first non-DNA sequence support for an Afrotherian clade. The comparative human and elephant genome maps provide new insights into the karyotype organization of the proto-afrotherian, the ancestor of extant placental mammals, which most probably consisted of 2n=46 chromosomes.  Erratum in: Proc R Soc Lond B Biol Sci. 2003 Dec 22;270(1533):2639.  Comparative Molecular Cytogenetics Section, National Cancer Institute-Genetics Branch, National Cancer Institute-Frederick, Nationa Institutes of Health, Building 560, Room 11-75, Frederick, MD 21702-1201, USA. froenickel@gmx.net

Martin, F., Niemitz, C., 2003. "Right-trunkers" and "left-trunkers": side preferences of trunk movements in wild Asian elephants (Elephas maximus). J. Comp Psychol. 117, 371-379.
Abstract: In this article, the side preferences of feeding-related trunk movements of free-ranging Asian elephants (Elephas maximus) were investigated for the first time. It is hypothesized that a functional asymmetry of the trunk is necessary to perform skillful feeding movements more efficiently. This might be connected with a corresponding hemispheric specialization. Video recordings of 41 wild elephants provided frequencies and durations of the following trunk-movement categories: object contact, retrieval, and reaching. In each category, individual side preferences were found. The strength of side preferences varied between the trunk-movement categories and the sexes. Mean durations of retrieval and reaching correlated negatively with the strength of side biases. Comparing the side preferences in the unpaired trunk with analogous phenomena in other unpaired grasping organs and in primate handedness. the authors discuss possible explanations for the evolution of asymmetries in unpaired grasping organs.

Meredith, M., 2003. Biography of an Endangered Species in Africa. PublicAffairs.
Abstract: Review from Publishers Weekly:  In this solid introduction to the world of elephants, Meredith covers all  the major topics including biology, social behavior, recent scientific  discoveries, ancient elephantology, the devastating  ivory trade, the truth  about elephant graveyards and the insistent threat  of extinction. Meredith demonstrates that human involvement in elephantine affairs has been disastrous to the pachyderm: the quest for ivory had caused the extinction of all Syrian herds by 500 B.C.; many ancient cultures took elephants to war; and Romans used the animals in their blood sports. Much of the book follows the history of the European exploitation of Africa's three treasures: gold, slaves and ivory. The quantities of murdered elephants and descriptions of killing methodologies are deeply affecting. Once Meredith's history reaches modern times, the shock of  population counts is astounding in comparison with the numbers of elephants that roamed free in the past.   Aristotle's treatise on the animals' anatomy,  behavior, diet and  reproduction was the beginning of a long line of nterest, but only recently  has science uncovered the answers to mysteries such as how separate herds coordinate movement over many miles. Meredith's primer on elephantine
matters will help turn a reader's casual interest into a fascination.

Morgan, B.J., Lee, P.C., 2003. Forest elephant (Loxodonta africana cyclotis) stature in the Reserve de Faune du Petit Loango, Gabon. Journal of Zoology 259, 337-344.
Abstract: The stature of forest elephants Loxodonta africana cyclotis was determined at the Petit Loango Reserve over 14 months from January to December 1998 and May to June 1999 using three measures: shoulder height, hind footprint length and boli diameter. The shoulder height of 53 identified elephants was measured using photogrammetric methods. The minimum estimated shoulder height was 69 cm from a young calf, and the tallest animal was 216 cm. Hind footprint length and boli diameter data were collected from unidentified individuals. The minimum footprint size was 12.5 cm and the largest 35.3 cm. Boli diameter ranged from 4.0 to 16.0 cm. A comparison of the size categories with those of savanna elephants in Amboseli National Park, Kenya, suggested a similar distribution of size, age and population structure, despite a marked difference in overall stature. These are the first data for measures of African forest elephant size compared to African savanna elephant size. Such data may add morphological evidence supporting recent genetic work suggesting that African forest elephants be re-classified as a distinct species from the African savanna elephant.

Murata, Y., Nikaido, M., Sasaki, T., Cao, Y., Fukumoto, Y., Hasegawa, M., Okada, N., 2003. Afrotherian phylogeny as inferred from complete mitochondrial genomes. Mol Phylogenet Evol 28, 253-260.
Abstract: Afrotheria is a huge assemblage of various mammals encompassing six orders that were once classified as distantly related groups. This superordinal relationship may have resulted from the break-up of Gondowanaland followed by the isolation of the African continent between 105 and 40 million years ago. Although the monophyly of Afrotheria is well supported by recent molecular studies, the interrelationships within afrotherian mammals remain unclarified. In this study, we determined the sequence of the complete mitochondrial genomes of hyrax, golden mole, and elephant shrew. These sequences were compared with those of other eutherians to analyze the phylogenetic relationships among afrotherians and, in particular, those among paenungulates. Our mitochondrial genome analysis supports the monophyly of Tethytheria.

Nikaido, M., Nishihara, H., Hukumoto, Y., Okada, N., 2003. Ancient SINEs from African endemic mammals. Mol Biol Evol 20, 522-527.
Abstract: Afrotheria is a newly recognized taxon comprising elephants, hyraxes, sea cows, aardvarks, golden moles, tenrecs, and elephant shrews, each of which originated in Africa. Although some members of this taxon were once classified into distantly related groups, recent molecular studies have demonstrated their close relationships. It was suggested that this group emerged as a result of physical isolation of the African continent during the successive breakup events of Gondowanaland. In this study, a novel family of SINEs, designated AfroSINEs, was isolated and characterized from the genomes of afrotherians. This SINE family is distributed exclusively among the afrotherian species, confirming their monophyletic relationships. Furthermore, a distinct subfamily, which shares a deletion in the middle region of the SINE, was identified. The distribution of this subfamily is apparently restricted to the genomes of hyraxes, elephants, and sea cows, suggesting monophyly of these three groups, which was previously proposed as Paenungulata. We characterized the structures of the AfroSINEs from all afrotherian representatives by PCR, and we discuss how they were generated as well as the phylogenetic relationships of their host species. Graduate School of Bioscience and Biotechnology, Tokyo Institute of Technology, Yokohama, Japan.

Shimoyama, M., Ninomiya, T., Ozaki, Y., 2003. Nondestructive discrimination of ivories and prediction of their specific gravity by Fourier-transform Raman spectroscopy and chemometrics. Analyst. 128, 950-953.
Abstract: Fourier-transform (FF) Raman spectroscopy and chemometrics were used for nondestructive analysis of ivories. The discrimination of five kinds of ivories, two subspecies of African elephant, mammoth, hippopotamus, and sperm whale, was investigated, and a calibration model for predicting their specific gravity was developed. FT-Raman spectra were measured in situ for them and chemometrics analyses were carried out for the 3050-350 cm(-1) region. The five kinds of ivories were clearly discriminated from each other on the scores plots of two or three principal components (PCs) obtained by principal component analysis (PCA). The loadings plot for PC 1 shows that the discrimination relies on the content ratio of organic collagenous protein and inorganic hydroxyapatite of ivories. The loadings plot for PC 2 shows that bands due to the CH3 and CH2 stretching modes of the protein also play a role in the discrimination. Using partial least squares regression (PLSR), we developed a calibration model that predicts the specific gravity of the ivories from the FT-Raman spectra. The correlation coefficient and root mean square error of cross validation (RMSECV) of this model were 0.980 and 0.024, respectively. Forensic Science Laboratory, Hyogo Prefectural Police Headquarters, Shimoyamate-dori, Chuo-ku, Kobe 650-8510, Japan.

West, J.B., Fu, Z., Gaeth, A.P., Short, R.V., 2003. Fetal lung development in the elephant reflects the adaptations required for snorkeling in adult life. Respir Physiol Neurobiol 138, 325-333.
Abstract: The adult elephant is unique among mammals in that the pleural membranes are thickened and the pleural cavity is obliterated by connective tissue. It has been suggested that this peculiar anatomy developed because the animal can snorkel at depth, and this behavior subjects the microvessels in the parietal pleura to a very large transmural pressure. To investigate the development of the parietal pleura, the thickness of the endothoracic fascia (ET) was measured in four fetal African elephants of approximate gestational age 111-130 days, and the appearances were compared with those in human, rabbit, rat and mouse fetuses of approximately the same stage of lung organogenesis. The mean thicknesses of ET in the elephant, human, rabbit, rat and mouse were 403, 53, 29, 27 and 37 microm, respectively. This very early development of a thick parietal pleura in the elephant fetus is consistent with the hypothesis of a long history of snorkeling in the elephant's putative aquatic ancestors. Department of Medicine, University of California San Diego, 9500 Gilman Drive, La Jolla, CA 92093-0623, USA. jwest@ucsd.edu

Yang, F., Alkalaeva, E.Z., Perelman, P.L., Pardini, A.T., Harrison, W.R., O'Brien, P.C., Fu, B., 2003. Reciprocal chromosome painting among human, aardvark, and elephant (superorder Afrotheria) reveals the likely eutherian ancestral karyotype. Proc Natl Acad Sci U S A 100, 1062-1066.
Abstract: The Afrotheria, a supraordinal grouping of mammals whose radiation is rooted in Africa, is strongly supported by DNA sequence data but not by their disparate anatomical features. We have used flow-sorted human, aardvark, and African elephant chromosome painting probes and applied reciprocal painting schemes to representatives of two of the Afrotherian orders, the Tubulidentata (aardvark) and Proboscidea (elephants), in an attempt to shed additional light on the evolutionary affinities of this enigmatic group of mammals. Although we have not yet found any unique cytogenetic signatures that support the monophyly of the Afrotheria, embedded within the aardvark genome we find the strongest evidence yet of a mammalian ancestral karyotype comprising 2n = 44. This karyotype includes nine chromosomes that show complete conserved synteny to those of man, six that show conservation as single chromosome arms or blocks in the human karyotype but that occur on two different chromosomes in the ancestor, and seven neighbor-joining combinations (i.e., the synteny is maintained in the majority of species of the orders studied so far, but which corresponds to two chromosomes in humans). The comparative chromosome maps presented between human and these Afrotherian species provide further insight into mammalian genome organization and comparative genomic data for the Afrotheria, one of the four major evolutionary clades postulated for the Eutheria.

Journal of Indian Veterinary Assocaition Kerala. Journal of Indian Veterinary Association Kerala 7[3], 1-64. 2002.
Ref Type: Journal (Full)

Alex, P.C., 2002. The Musth, the vicious and the rogue elephants - a review. Journal of Indian Veterinary Association Kerala 7, 26-27.

Anilkumar, K., 2002. Phylogeny of elephants. Journal of Indian Veterinary Association Kerala 7, 15-17.

Bist, S.S., 2002. The status of the domesticated elephants in India. Journal of Indian Veterinary Association Kerala 7, 4-7.

Chandrasekharan, K., 2002. Specific diseases of Asian elephants. Journal of Indian Veterinary Association Kerala 7, 31-34.
Abstract: The earliest writing describing the diseases of elephants in ancient literature said to be the works on "Gajasastra" (Elephantology) written in Sanskrit by authors like Gautama, Narada, Mrigacharma, Rajaputra and Vyasa. "Hasthyayurveda" a legendary book in Sanskrit written by a safe Palakapya deals with some diseases, treatment, desirable and undesirable points of selection, management practices and some mythological aspects on the origin of elephants. The earliest book in English dealing with diseases of elephants seems to be that of W. Gilchrist "A practical treatise on the treatment of diseases of elephants" published in 1848. Later Slym (1873), Sanderson (1878), Steel (1885), Evans (1910), Herpburn (1913), Milroy (1922), Ptaff (1940), Ferrier (1947), Utoke Gale (1974), Chandrasekharan (1979) and Panicker (1985) have documented their findings on the incidence, etiology and control of diseases of Asian elephants.

Eggert, L.S., Rasner, C.A., Woodruff, D.S., 2002. The evolution and phylogeography of the African elephant inferred from mitochondrial DNA sequence and nuclear microsatellite markers. Proc R Soc Lond B Biol Sci 269 , 1993-2006.
Abstract: Recent genetic results support the recognition of two African elephant species: Loxodonta africana, the savannah elephant, and Loxodonta cyclotis, the forest elephant. The study, however,#10; did not include the populations of West Africa, where the taxonomic affinities of elephants have been much debated. We examined mitochondrial cytochrome b control region sequences and four microsatellite#10; loci to investigate the genetic differences between the forest and savannah elephants of West and Central Africa. We then combined our data with published control region sequences from across Africa to#10; examine patterns at the continental level. Our analysis reveals several deeply divergent lineages that do not correspond with the currently recognized taxonomy: (i) the forest elephants of Central Africa;#10; (ii) the forest and savannah elephants of West Africa; and (iii) the savannah elephants of eastern, southern and Central Africa. We propose that the complex phylogeographic patterns we detect in African#10; elephants result from repeated continental-scale climatic changes over their five-to-six million year evolutionary history. Until there is consensus on the taxonomy, we suggest that the genetic and ecological#10; distinctness of these lineages should be an important factor in conservation management planning. #10;

Forsyth, I.A., Wallis, M., 2002. Growth hormone and prolactin--molecular and functional evolution. J Mammary Gland Biol Neoplasia 7, 291-312.
Abstract: Growth hormone, prolactin, the fish hormone, somatolactin, and related mammalian placental hormones, including placental lactogen, form a family of polypeptide hormones that share a common tertiary structure. They produce their biological effects by interacting with and dimerizing specific single transmembrane-domain receptors. The receptors belong to a superfamily of cytokine receptors with no intrinsic tyrosine kinase, which use the Jak-Stat cascade as a major signalling pathway. Hormones and receptors are thought to have arisen as a result of gene duplication and subsequent divergence early in vertebrate evolution. Mammalian growth hormone and prolactin show a slow basal evolutionary rate of change, but with episodes of accelerated evolution. These occurred for growth hormone during the evolution of the primates and artiodactyls and for prolactin in lineages leading to rodents, elephants, ruminants, and man. Placental lactogen has probably evolved independently on three occasions, from prolactin in rodents and ruminants and from growth hormone in man. Receptor sequences also show variable rates of evolution, corresponding partly, but not completely, with changes in the ligand. A principal biological role of growth hormone, the control of postnatal growth, has remained quite consistent throughout vertebrate evolution and is largely mediated by insulin-like growth factors. Prolactin has many and diverse roles. In relation to lactation, the relative roles of growth hormone and prolactin vary between species. Correlation between the molecular and functional evolution of these hormones is very incomplete, and it is likely that many important functional adaptations involved changes in regulatory elements, for example, altering tissue of origin or posttranscriptional processing, rather than change of the structures of the proteins themselves. The Babraham Institute, Babraham, Cambridge, United Kingdom. isabel.forsyth@bbsrc.ac.uk

Hashimoto, K., 2002. Historical accounts of elephants imported before the Meiji period. Journal of Veterinary Medicine, Japan 55, 776-780.

Murali, K., 2002. An introduction to Hastyayurveda. Journal of Indian Veterinary Association Kerala 7, 54,63-63.

Poulakakis, N., Theodorou, G.E., Zouros, E., Mylonas, M., 2002. Molecular phylogeny of the extinct pleistocene dwarf elephant Palaeoloxodon antiquus falconeri from Tilos Island, Dodekanisa, Greece. J Mol Evol 55, 364-374.
Abstract: A partial sequence of cytochrome b (228 bp) gene of mitochondrial DNA was successfully determined from rib bones of the dwarf elephant Palaeoloxodon antiquus falconeri BUSK, which were excavated from Charkadio cave of the island of Tilos, Dodekanisa, Greece. This is the first report of DNA sequence of a dwarf elephant. The sequences were used to examine the phylogenetic relationships among Elephantidae. Phylogenetic trees reconstructed by the neighbor-joining and maximum parsimony methods provided identical topologies. The results support the "Palaeoloxodon-Elephas" clade, which is consistent with previous morphological reports according to which Palaeoloxodon is more closely related to Elephas than to Loxodonta or Mammuthus.

Roca, A.L., Georgiadis, N., Pecon-Slattery, J., O'Brien, S.J., 2002. Genetic evidence for two species of elephant in Africa. Science 293, 1473-1477.
Abstract: Elephants from the tropical forests of Africa are morphologically distinct from savannah or bush elephants. Dart-biopsy samples from 195 free-ranging African elephants in 21 populations were examined for DNA sequence variation in four nuclear genes (1732 base pairs). Phylogenetic distinctions between African forest elephant and savannah elephant populations corresponded to 58% of the difference in the same genes between elephant genera Loxodonta (African) and Elephas (Asian). Large genetic distance, multiple genetically fixed nucleotide site differences, morphological and habitat distinctions, and extremely limited hybridization of gene flow between forest and savannah elephants support the recognition and conservation management of two African species: Loxodonta africana and Loxodonta cyclotis.

Schweitzer, M., Hill, C.L., Asara, J.M., Lane, W.S., Pincus, S.H., 2002. Identification of immunoreactive material in mammoth fossils. J Mol Evol 55, 696-705.
Abstract: Department of Microbiology, Montana State University, Bozeman, MT 59717, USA. schweitzer@montana.edu

The fossil record represents a history of life on this planet. Attempts to obtain molecular information from this record by analysis of nucleic acids found within fossils of extreme age have been unsuccessful or called into question. However, previous studies have demonstrated the long-term persistence of peptides within fossils and have used antibodies to extant proteins to demonstrate antigenic material. In this study we address two questions: Do immunogenic/antigenic materials persist in fossils? and; Can fossil material be used to raise antibodies that will cross-react with extant proteins? We have used material extracted from a well-preserved 100000-300000-year-old mammoth skull to produce antisera. The specificity of the antisera was tested by ELISA, western blotting, and immunohistochemistry. It was demonstrated that antisera reacted specifically with the fossils and not the surrounding sediments. Reactivity of antisera with modern proteins and tissues was also demonstrated, as was the ability to detect evolutionary relationships via antibody-antigen interactions. Mass spectrometry demonstrated the presence of amino acids and specific peptides within the fossil. Peptides were purified by anion-exchange chromatography and sequenced by tandem mass spectrometry. The collagen-derived peptides may have been the source of at least some of the immunologic reactivity, but the antisera identified molecules that were not observed by mass spectrometry, indicating that immunologic methods may have greater sensitivity. Although the presence of peptides and amino acids was demonstrated, the exact nature of the antigenic material was not fully clarified. This report demonstrates that antibodies may be used to obtain information from the fossil record.

Scigliano, E., 2002. Love, war, and circuses : the age-old relationship between elephants and humans. Houghton Mifflin, Boston.

Sethumadhavan, T.P., 2002. Who will win the battle? man or elephant? Journal of Indian Veterinary Association Kerala 7, 3.

Vogel, G., 2002. Ecology. African elephant species splits in two. Science 293, 1414.

Brody, R.H., Edwards, H.G.M., Pollard, A.M., 2001. Chemometric methods applied to the differentiation of Fourier-transform Raman spectra of ivories. Analytica Chimica Acta 427, 223-232.

Cozzi, B., Spagnoli, S., Bruno, L., 2001. An overview of the central nervous system of the elephant through a critical appraisal of the literature published in the XIX and XX centuries. Brain Res Bull 54, 219-227.
Abstract: The two species of elephants (Indian: Elephas maximus and African: Loxodonta africana) possess the largest brain among land mammals. Due to its size, the elephant brain is discussed in virtually every paper dealing with the evolution of the central nervous system of mammals and comparative brain size. Studies on the social habits of elephants also deal with the skills and the "intelligence" and brain size of these species. Yet most of the descriptions and conclusions reported in comparative studies rely on second-hand data derived from investigations performed several decades before, often dating as far back as the XIX century. Furthermore, many of the original papers actually describing gross and detailed features of the brain of elephants are either no longer available, are written in languages other than English, or are difficult to trace. The present study gives a short description of the anatomy of the central nervous system of elephants, with special attention to its distinctive features, reports all available literature on the subject, and briefly discusses its origins and rationale.

Fleischer, R.C., Perry, E.A., Muralidharan, K., Stevens, E.E., Wemmer, C.M., 2001. Phylogeography of the Asian elephant (Elephas maximus) based on mitochondrial DNA. Evolution 55, 1882-1892.
Abstract: Populations of the Asian elephant (Elephas maximus) have been reduced in size and become highly fragmented during the past 3000 to 4000 years. Historical records reveal elephant dispersal by humans via trade and war. How have these anthropogenic impacts affected genetic variation and structure of Asian elephant populations? We sequenced mitochondrial DNA (mtDNA) to assay genetic variation and phylogeography across much of the Asian elephant's range. Initially, we compared cytochrome b sequences (cyt b) between 9 Asian and 5 African elephants and used the fossil-based age  of their separation (_5 million years ago) to obtain a rate of about 0.013  (95% CI=0.011-0.018) corrected sequence divergence per million years. We also assessed variation in part of the mtDNA control region (CR) and adjacent tRNA genes in 57 Asian elephants from 7 countries (Sri Lanka,  India, Nepal, Myanmar, Thailand, Malaysia, and Indonesia). Asian elephants had typical levels of mtDNA variation, and coalescence analyses suggested their populations were growing in the late Pleistocene. Reconstructed  phylogenies revealed 2 major clades (A and B) differing on average by HKY85/GAMMA-corrected distances of 0.020 for cyt b and 0.050 for the CR segment (corresponding to a coalescence time based on our cyt b rate of   _1.2 million years). Individuals of both major clades existed in all locations, but Indonesia and Malaysia. Most elephants from Malaysia and all from Indonesia were in well-supported, basal clades within clade A, thus supporting their status as evolutionarily significant units (ESUs). The proportion of clade A individuals decreased to the north, which could result from retention and subsequent loss of ancient lineages in long-term stable populations or, perhaps more likely, via recent mixing of 2 expanding populations that were isolated in the mid-Pleistocene. The distribution of clade A individuals appeared to be impacted by human trade in elephants among Myanmar, Sri Lanka, and India, and the subspecies and  ESU statuses of Sri Lankan elephants were not supported by molecular data.

Greenwood, A.D., Lee, F., Capelli, C., DeSalle, R., Tikhonov, A., Marx, P.A., MacPhee, R.D., 2001. Evolution of endogenous retrovirus-like elements of the woolly mammoth (Mammuthus primigenius) and its relatives. Mol Biol Evol 18, 840-870.
Abstract: Endogenous retrovirus-like elements characterizable by a leucine tRNA primer (ERV-Ls) are reiterated genomic sequences known to be widespread in mammals, including humans. They may have arisen from an ancestral foamy virus-like element by successful germ line infection followed by copy number expansion. However, among mammals, only primates and rodents have thus far exhibited high copy number amplification and sequence diversification. Conventionally, empirical studies of proviral amplification and diversification have been limited to extant species, but taxa having good Quaternary fossil records could potentially be investigated using the techniques of "ancient" DNA research. To examine evolutionary parameters of ERV-Ls across both time and taxa, we characterized this proviral class in the extinct woolly mammoth (Mammuthus primigenius) and living elephants, as well as extant members of the larger clade to which they belong (Uranotheria, a group containing proboscideans, sirenians, hyraxes, and their extinct relatives). Ungulates and carnivores previously analyzed demonstrated low copy numbers of ERV-L sequences, and thus it was expected that uranotheres should as well. Here, we show that all uranothere taxa exhibit unexpectedly numerous and diverse ERV-L sequence complements, indicating active expansion within this group of lineages. Selection is the most parsimonious explanation for observed differences in ERV-L distribution and frequency, with relative success being reflected in the persistence of certain elements over a variety of sampled time depths (as can be observed by comparing sequences from fossil and extant elephantid samples).

Haynes, G. Elephant landscapes: human foragers in the world of mammoths, mastodons, and elephants. The World of Elephants-International Congress.  571-576. 2001.
Ref Type: Conference Proceeding
Abstract: Human groups able to subsist by opportunistic exploitation of proboscideans would be afforded abundant environmental clues to prey health and density, along with superior nutrients and other advantages such as information-rich trail networks.

Hecht, J. Telltale bones. New Scientist [2312], 14. 2001.
Ref Type: Magazine Article

Kirkman, S., ., Wallace, E.D., van Aarde, R.J., Potgieter, H.C., 2001. Steroidogenic correlates of pregnancy in the rock hyrax (Procavia capensis). Life Sciences 68, 2061-2072.
Abstract: In pregnant rock hyraxes (P. capensis) isolated leucocytes metabolize both [3H]pregnenolone and [3H]progesterone while whole blood, erythrocytes and an erythrocyte/leucocyte mixture only metabolized [3H]progesterone. Plasma displayed no tendency to metabolically convert any one of these two steroids. In whole blood, [3H]progesterone appears to be converted to 5alpha-pregnane-3,20-dione and a compound with chromatographic properties similar to that of 5alpha-pregnan-3alpha-ol-20-one. 5alpha-Pregnane-3,20-dione exhibited a high relative binding affinity for the uterine progesterone receptor (94%), but 5alpha-pregnan-3alpha-ol-20-one displayed very little affinity for the same receptor (0.4%). 5alpha-Pregnane-3,20-dione may therefore aid in the maintenance of pregnancy. Corpora lutea metabolized progesterone to 17alpha-hydroxyprogesterone, a compound exhibiting no progestational function because of its low relative binding affinity for the uterine progesterone receptor (2%). Progesterone appears to be the main product of the corpus luteum. However, 5alpha-pregnane-3,20-dione circulated at concentrations approximately 8.5 times higher than progesterone, probably due to the metabolic conversion of progesterone to 5alpha-pregnane-3,20-dione by the blood. We conclude that in the hyrax, progesterone, produced by the corpora lutea, enters the circulation, where it is reduced to 5alpha-pregnanes. 5alpha-Pregane-3,20-dione may then be transported to the uterus where it binds to the progesterone receptor to assist in the maintenance of pregnancy. This mechanism appears to be analogous to that of the African elephant (Loxodonta africana) which is phylogenetically related to the hyrax, except that in the elephant the 5alpha-reduced metabolites are produced by luteal tissue and not the blood.

Lister, A.M., Sher, A.V., 2001. The origin and evolution of the woolly mammoth. Science 294, 1094-1097.
Abstract: The mammoth lineage provides an example of rapid adaptive evolution in response to the changing environments of the Pleistocene. Using well-dated samples from across the mammoth's Eurasian range, we document geographical and chronological variation in adaptive morphology. This work illustrates an incremental (if mosaic) evolutionary sequence but also reveals a complex interplay of local morphological innovation, migration, and extirpation in the origin and evolution of a mammalian species. In particular, northeastern Siberia is identified as an area of successive allopatric innovations that apparently spread to Europe, where they contributed to a complex pattern of stasis, replacement, and transformation.

Mahasavangkul, S. Domestic Elephant Status and Management in Thailand. A Research Update on Elephants and Rhinos; Proceedings of the International Elephant and Rhino Research Symposium, Vienna, June 7-11, 2001.  71-82. 2001. Vienna, Austria, Schuling Verlag. 2001.
Ref Type: Conference Proceeding

McComb, K., Moss, C., Durant, S.M., Baker, L., Sayialel, S., 2001. Matriarchs as repositories of social knowledge in African elephants. Science 292, 491-494.
Abstract: Despite widespread interest in the evolution of social intelligence, little is known about how wild animals acquire and store information about social companions or whether individuals possessing enhanced social knowledge derive biological fitness benefits. Using playback experiments on African elephants (Loxodonta africana), we demonstrated that the possession of enhanced discriminatory abilities by the oldest individual in a group can influence the social knowledge of the group as a whole. These superior abilities for social discrimination may result in higher per capita reproductive success for female groups led by older individuals. Our findings imply that the removal of older, more experienced individuals, which are often targets for hunters because of their large size, could have serious consequences for endangered populations of advanced social mammals such as elephants and whales.

Miller, D.L., Dougherty, M.M., Decker, S.J., Bossart, G.D., 2001. Ultrastructure of the spermatozoa from a Florida manatee (Trichechus manatus latirostris). Anat Histol Embryol 30, 253-256.
Abstract: Semen was opportunistically collected from a free-ranging, 10-year-old, 275 cm (total length) Florida manatee (Trichechus manatus latirostris) during rehabilitation treatments. Ultrastructure of the spermatozoa was examined by scanning and transmission electron microscopy and differed slightly from that described for other mammals. Comparisons to the manatee's closest phylogenetic relatives, the elephant and hyrax, were made. The manatee spermatozoa had a similar acrosome but a distinct annulus and lacked the dense bodies observed in the neck of the elephant spermatozoa. Additionally, manatee spermatozoa lacked the lateral vacuoles observed in the nuclear chromatin from of the hyrax spermatozoa. These data add to our understanding of manatees and allow for comparative studies with other species that may be useful in phylogenetic and reproductive studies.

Murphy, W.J., Eizirik, E., Johnson, W.E., Zhang, Y.P., Ryder, O.A., O'Brien, S.J., 2001. Molecular phylogenetics and the origins of placental mammals. Nature 409, 614-618.
Abstract: The precise hierarchy of ancient divergence events that led to the present assemblage of modern placental mammals has been an area of controversy among morphologists, paleontologists and molecular evolutionists. Here we address the potential weaknesses of limited character and taxon sampling in a comprehensive molecular phylogenetic analysis of 64 species sampled across all extant orders of placental mammals. We examined sequence variation in 18 homologous gene segments (including nearly 10,000 base pairs) that were selected for maximal phylogenetic informativeness in resolving the hierarchy of early mammalian divergence. Phylogenetic analyses identify four primary superordinal clades: (I) Afrotheria (elephants, manatees, hyraxes, tenrecs, aardvark and elephant shrews); (II) Xenarthra (sloths, anteaters and armadillos); (III) Glires (rodents and lagomorphs), as a sister taxon to primates, flying lemurs and tree shrews; and (IV) the remaining orders of placental mammals (cetaceans, artiodactyles, perissodactyles, carnivores, pangolins, bats and core insectivores). Our results provide new insight into the pattern of the early placental mammal radiation.

Okayama, T., Sugardjito, J., Yusuf, I. Mitochondrial DNA Analysis of Sumatran Elephants. A Research Update on Elephants and Rhinos; Proceedings of the International Elephant and Rhino Research Symposium, Vienna, June 7-11, 2001.  278-281. 2001. Vienna, Austria, Schuling Verlag. 2001.
Ref Type: Conference Proceeding

Schulte, B.A. Examining Ideas on the Evolution of Musth. A Research Update on Elephants and Rhinos; Proceedings of the International Elephant and Rhino Research Symposium, Vienna, June 7-11, 2001.  287. 2001. Vienna, Austria, Schuling Verlag. 2001.
Ref Type: Conference Proceeding

van Dijk, M.A., Madsen, O., Catzeflis, F., Stanhope, M.J., De Jong, W.W., Pagel, M., 2001. Protein sequence signatures support the African clade of mammals. Proc Natl Acad Sci U S A 98, 188-189.

West, J.B., 2001. Snorkel breathing in the elephant explains the unique anatomy of its pleura. Respiratory Physiology 126, 1-8.
Abstract: It has been known for over 300 years that the anatomy of the elephant lung is unique among mammals in that the pleural cavity is obliterated by connective tissue. However no satisfactory explanation has been advanced. Recent studies suggest that the elephant has an aquatic ancestry and the trunk may have developed for snorkeling. In addition, the modern day elephant is the only mammal that can remain submerged far below the surface of the water while snorkeling. The resulting differences of pressures within the thorax mean that the small blood vessels of the pleura are in great danger of rupturing or causing severe edema. The same distribution of pressures occurs when the animal raises water inside its trunk prior to drinking although in this case the pressure differences are relatively short-lived. Evolution has provided a remarkable solution to this problem by replacing the normally delicate parietal and visceral pleurae by dense connective tissue, and separating the two pleurae by loose connective tissue to allow some sliding movement.

Whitehouse, A.M., Harley, E.H., 2001. Post-bottleneck genetic diversity of elephant populations in South Africa,revealed using microsatellite analysis. Mol Ecol 10, 2139-2149.
Abstract: Widespread hunting had fragmented and severely reduced elephant populations in South Africa by 1900. Elephant numbers increased during the 1900s, although rates of recovery of individual populations varied. The Kruger National Park elephant population increased rapidly, to more than 6000 by 1967, with recruitment boosted by immigration from Mozambique. The Addo Elephant National Park population was reduced to 11 elephants in 1931 and remains relatively small (n = 325). Loss of genetic variation is expected to occur whenever a population goes through a bottleneck, especially when post-bottleneck recovery is slow. Variation at nine polymorphic microsatellite loci was analysed for Kruger and Addo elephants, as well as museum specimens of Addo elephants shot prior to the population bottleneck. Significantly reduced genetic variation and heterozygosity were observed in Addo in comparison to Kruger (mean alleles/locus and H(E): Addo 1.89, 0.18; Kruger 3.89, 0.44). Two alleles not present in the current Addo population were observed in the museum specimens. Addo elephants represent a genetic subset of the Kruger population, with high levels of genetic differentiation resulting from rapid genetic drift. The Kruger population is low in genetic diversity in comparison to East African elephants, confirming this population also suffered an appreciable bottleneck.

Alexander, S., 2000. The Astonishing Elephant. Random House, New York.

Anderson, D., Wood, M. Web-based Search Templates on Elephants. Elephants: Cultural, Behavioral, and Ecological Perspectives; Program and Abstracts of the Workshop.  8. 2000. Davis, CA. 2000.
Ref Type: Conference Proceeding

Debuyst, R., Callens, F., Frechen, M., Dejehet, F., 2000. ESR study of elephant tooth enamel from the Karlich-Seeufer site in Germany. Appl Radiat Isot 52, 1327-1336.
Abstract: Enamel from 6 different positions in a well preserved elephant tooth from the Karlich-Seeufer site in Germany has been irradiated up to 32 kGy. The X-band (v = 9.5 GHz) ESR spectra of two subsamples have been decomposed into three real components with Maximum Likelihood Common Factor Analysis (MLCFA). One of these components due to orthorhombic CO2- radicals is predominant. Dose response curves for the contributions of these MLCFA components and for different heights in the ESR spectra have been obtained and fitted with different models. Depending on the model, the equivalent dose for the preferably used height at g = 1.9973, due to CO2-, ranges from 70 to 130 Gy. Due to a very low uranium and thorium content in both enamel and dentine (< or = approximately 10 ppb) and to an important external y-attenuation, the ages fluctuate between 300 and 575 ka.

Fernando, P., Pfrender, M.E., Encalada, S.E., Lande, R., 2000. Mitochondrial DNA variation, phylogeography and population structure of the Asian elephant. Heredity 84, 362-372.
Abstract: We report the first genetic analysis of free-ranging Asian elephants (Elephas maximus). We sampled 118 elephants from Sri Lanka, Bhutan/North India, and Laos/Vietnam by extracting DNA from dung, PCR amplifying and sequencing 630 nucleotides of mtDNA, including part of the variable left domain of the control region. Comparison with African elephant (Loxodonta africana) sequences indicated a relatively slow molecular clock in the Proboscidea with a sequence divergence of _1%/million years. Genetic diversity within Asian elephants was low, suggesting a small long-term effective population size. 17 haplotypes were identified within Asian elephants, which clustered into 2 well-differentiated assemblages with an estimated Pliocene divergence of 2.5-3.5 million years ago. The 2 assemblages showed incomplete geographical partitioning, suggesting allopatric divergence and secondary admixture. On the mainland, little genetic differentiation was observed between elephant populations of Bhutan and India or Laos and Vietnam. A significant difference in haplotype frequencies but relatively weak subdivision was observed between the Bhutan-India and Laos-Vietnam regions. Significant genetic differentiation was observed between the mainland and Sri Lanka and between northern, mid-latitude and southern regions in Sri Lanka.

Hall, T., 2000. To the elephant graveyard. John Murray, London.

Hart, L.A. The Mahout-Elephant Relationship: Ancient and Modern Versions. Elephants: Cultural, Behavioral, and Ecological Perspectives; Program and Abstracts of the Workshop.  13. 2000. Davis, CA. 2000.
Ref Type: Conference Proceeding

Shelton, J. Dancing and Dying: The Display of Elephants in Ancient Roman Arenas. Elephants: Cultural, Behavioral, and Ecological Perspectives; Program and Abstracts of the Workshop.  21. 2000. Davis, CA. 2000.
Ref Type: Conference Proceeding

Thomas, M.G., Hagelberg, E., Jones, H.B., Yang, Z., Lister, A.M., 2000. Molecular and morphological evidence on the phylogeny of the Elephantidae. Proceedings of the Royal Society of London SeriesB, Biological-Sciences 267:1561, 2493-2500.
Abstract: The African and Asian elephants and the mammoth diverged ca. 4-6 million years ago and their phylogenetic relationship has been controversial. Morphological studies have suggested a mammoth-Asian elephant relationship, while molecular studies have produced conflicting results. We obtained cytochrome b sequences of up to 545 base pairs from five mammoths, 14 Asian and eight African elephants. A high degree of polymorphism is detected within species. With a dugong sequence used as the outgroup, parsimony and maximum-likelihood analyses support a mammoth-African elephant clade. As the dugong is a very distant outgroup, we employ likelihood analysis to root the tree with a molecular clock, and use bootstrap and Bayesian analyses to quantify the relative support for different topologies. The analyses support the mammoth-African elephant relationship, although other trees cannot be rejected. Ancestral polymorphisms may have resulted in gene trees differing from the species phylogeny. Examination of morphological data, especially from primitive fossil members, indicates that some supposed synapomorphies between the mammoth and Asian elephant are variable, others convergent or autapomorphous. A mammoth-African elephant relationship is not excluded. Our results highlight the need, in both morphological and molecular phylogenetics, for multiple markers and close attention to within-taxon variation and outgroup selection.

Victor, S., Nayak, V.M., 2000. Evolutionary anticipation of the human heart. Ann R Coll Surg Engl 82, 297-302.
Abstract: We have studied the comparative anatomy of hearts from fish, frog, turtle, snake, crocodile, birds (duck, chicken, quail), mammals (elephant, dolphin, sheep, goat, ox, baboon, wallaby, mouse, rabbit, possum, echidna) and man. The findings were analysed with respect to the mechanism of evolution of the heart.

Wallis, M., 2000. Episodic evolution of protein hormones: molecular evolution of pituitary prolactin. J Mol Evol 50, 465-473.
Abstract: Previous studies have shown that pituitary growth hormone displays an episodic pattern of evolution, with a slow underlying evolutionary rate and occasional sustained bursts of rapid change. The present study establishes that pituitary prolactin shows a similar pattern. During much of tetrapod evolution the sequence of prolactin has been strongly conserved, showing a slow basal rate of change (approx 0.27x10(9) substitutions/amino acid site/year). This rate has increased substantially ( approximately 12- to 38-fold) on at least four occasions during eutherian evolution, during the evolution of primates, artiodactyles, rodents, and elephants. That these increases are real and not a consequence of inadvertent comparison of paralogous genes is shown (for at least the first three groups) by the fact that they are confined to mature protein coding sequence and not apparent in sequences coding for signal peptides or when synonymous substitutions are examined. Sequences of teleost prolactins differ markedly from those of tetrapods and lungfish, but during the course of teleost evolution the rate of change of prolactin has been less variable than that of growth hormone. It is concluded that the evolutionary pattern seen for prolactin shows long periods of near-stasis interrupted by occasional bursts of rapid change, resembling the pattern seen for growth hormone in general but not in detail. The most likely basis for these bursts appears to be adaptive evolution though the biological changes involved are relatively small.

Whitehouse, A.M., Hall-Martin, A.J., 2000. Elephants in Addo Elephant National Park, South Africa: Reconstruction of the population's history. Oryx 34, 46-55.
Abstract: The history of the Addo elephant population in South Africa, from the creation of the Addo Elephant National Park (AENP) in 1931 to the present (every elephant currently living within the park is known), was reconstructed. Photographic records were used as a primary source of historical evidence, in conjunction with all documentation on the population. Elephants can be identified in photographs taken throughout their life by study of the facial wrinkle patterns and blood vessel patterns in their ears. These characteristics are unique for each elephant and do not change during the individual's life. The life histories of individual elephants were traced: dates of birth and death were estimated and, wherever possible, the identity of the individual's mother was ascertained. An annual register of elephants living within the population, from 1931 to the present, was compiled, and maternal family trees constructed. Preliminary demographic analyses for the period 1976-98 are presented. The quantity and quality of photographs taken during these years enabled thorough investigation of the life histories of all elephants. Prior to 1976, insufficient photographs were available to provide reliable data on the exact birth dates and mothers' identities for every calf born. However, data on annual recruitment and mortality are considered sufficiently reliable for use in analyses of the population's growth and recovery.

Barriel, V., Thuet, E., Tassy, P., 1999. Molecular phylogeny of Elephantidae. Extreme divergence of the extant forest African elephant. C R Acad Sci III 322, 447-454.
Abstract: A phylogenetic study of the Elephantidae (Proboscidea, Mammalia) is based on the cytochrome b mitochondrial gene: 31 terminals, that is, all known sequences, one non-elephantid proboscidean, the extinct American mastodon, and four outgroups. The data set includes 11 new sequences with the first published sequence of the forest African elephant, L. a. cyclotis. The analyses of extant taxa only and of both extant and extinct taxa show that L. a. cyclotis is highly divergent from L. a. africana. It is as divergent from L. a. africana as Loxodonta is divergent from Elephas. Southern L. a. africana form a clade. The continental subspecies E. m. indicus is paraphyletic with individuals from India and Thailand closer to E. m. maximus (Sri-Lanka). Members of Mammuthus primigenius are more closely related to Loxodonta although they do not form a clade; two specimens of M. primigenius are closer to L. a. africana making the genus Loxodonta paraphyletic. The latter conclusion may be partly due to unequal length of the various polymorphic mammoth sequences.

Chatkupt, T.T., Sollod, A.E., Sarobol, S., 1999. Elephants in Thailand: determinants of health and welfare in working populations
531. J. Appl. Anim Welf. Sci. 2, 187-203.
Abstract: The Asian elephant (Elephas maximus) has played a prominent role in Thai history and society. However, in the face of modernization, elephant handlers have been struggling to justify their continued ownership. As a result, working elephants may still encounter situations in which their health and welfare are jeopardized. This study developed both a survey instrument and a visual assessment to describe and evaluate the health and living conditions of elephants encountered in a variety of work and living situations. These situations were found to be significantly associated with whether or not an elephant received proper husbandry or was in good body condition. These results may prove valuable in predicting the welfare of elephants according to work and living situations

Gaeth, A.P., Short, R.V., Renfree, M.B., 1999. The developing renal, reproductive, and respiratory systems of the African elephant suggest an aquatic ancestry. Proc Natl Acad Sci U S A 96, 5555-5558.
Abstract: The early embryology of the elephant has never been studied before. We have obtained a rare series of African elephant (Loxodonta africana) embryos and fetuses ranging in weight from 0.04 to 18.5 g, estimated gestational ages 58-166 days (duration of gestation is approximately 660 days). Nephrostomes, a feature of aquatic vertebrates, were found in the mesonephric kidneys at all stages of development whereas they have never been recorded in the mesonephric kidneys of other viviparous mammals. The trunk was well developed even in the earliest fetus. The testes were intra-abdominal, and there was no evidence of a gubernaculum, pampiniform plexus, processus vaginalis, or a scrotum, confirming that the elephant, like the dugong, is one of the few primary testicond mammals. The paleontological evidence suggests that the elephant's ancestors were aquatic, and recent immunological and molecular evidence shows an extremely close affinity between present-day elephants and the aquatic Sirenia (dugong and manatees). The evidence from our embryological study of the elephant also suggests that it evolved from an aquatic mammal.

Greenwood, A.D., Capelli, C., Possnert, G., Paabo, S., 1999. Nuclear DNA sequences from late Pleistocene megafauna. Mol Biol Evol 16, 1466-1473.
Abstract: We report the retrieval and characterization of multi- and single-copy nuclear DNA sequences from Alaskan and Siberian mammoths (Mammuthus primigenius). In addition, a nuclear copy of a mitochondrial gene was recovered. Furthermore, a 13,000-year-old ground sloth and a 33,000-year-old cave bear yielded multicopy nuclear DNA sequences. Thus, multicopy and single-copy genes can be analyzed from Pleistocene faunal remains. The results also show that under some circumstances, nucleotide sequence differences between alleles found within one individual can be distinguished from DNA sequence variation caused by postmortem DNA damage. The nuclear sequences retrieved from the mammoths suggest that mammoths were more similar to Asian elephants than to African elephants.

Groning, K., Saller, M., 1999. Elephants - A Cultural and Natural History. Konemann, Germany.

Rasmussen, L.E.L., 1999. Evolution of chemical signals in the Asian elephant, Elephas maximus: behavioural and ecological influences. Journal of Biosciences 24, 241-251.
Abstract: In antiquity, the Asian elephant, Elephas maximus, gradually spread southward and eastward to become a successfully surviving, ecologically dominant megaherbivore in the tropical environment of south-east Asia. The changing physical environment forced dynamic fluxes in its social structure and altered its metabolism. Such events shaped the production and ultimately the stability of certain chemicals released by body effluvia. Some of these chemicals took on significance as chemical signals and/or pheromones. This article demonstrates by experimental and observational evidence, and hypothesizes based on speculative reasoning, how and why specific chemical signals evolved in the modern Asian elephant. Evidence, including the functional criteria required by elephant social structure and ecology, is presented for the hypothesis that the recently identified female-emitted, male-received sex pheromone, (Z)-7-dodecenyl acetate evolved first as a chemical signal. Subsequently, the cohesiveness and harmony of small, matriarchally-led female groups were strengthened by a female-to-female chemical signal, recently defined behaviourally. The looser societal structure of freer, roaming males also became bounded by chemical signals; for the males, breath and temporal gland emissions, as well as urinary ones function in chemical signaling. Basic knowledge about elephant chemical signals is now linking chemical information to behaviour and beginning to demonstrate how these signals affect elephant social structure and enable the species to cope with environmental changes.

Rasmussen, L.E.L., Schulte, B.A., 1999. Ecological and biochemical constraints on pheromonal signaling systems in Asian elephants and their evolutionary implications. In: Johnston, R.E., Muller-Schwarze, D., Sorenson, P.W. (Eds.), Advances in Chemical Communication in Vertebrates 8. Kluwer/Academic/ Plenum Press, pp. 49-62.
Abstract: The Asian elephant is an unusual example of how intraspecies chemical communication helps maintain societal cohesiveness within familial and herd units. The amount of multi-directional chemical communication is surprising, because long-lived elephants have a highly organized society, are capable of trans-generational passage of information, possess a sophisticated vocalization system, and are capable of complex learning and tool use. This paper discusses the ecological, behavioral, and biochemical aspects of chemical signals in elephants from an evolutionary perspective. Diverse bodily emissions are utilized as intraspecies chemical signals (including pheromones), often with imposed biochemical constraints. In this chapter, chemosignals released from the temporal gland secretions and breath of male Asian elephants in musth and a urinary female-to-male preovulatory pheromone are utilized as examples of these concepts. Furthermore, specific behavioral and biochemical studies with (Z)-7-dodecenyl acetate (a female-to-male urinary sex pheromone) demonstrate that social context significantly influences responsivity (demonstrated by field studies in Myanmar) and that additional biochemical requirements, perhaps lipocalin-like proteins, may be required for full bioactivity. The remarkable convergent evolution of (Z)-7-dodecenyl acetate, both structurally and functionally, in elephants and Lepidoptera, allows the use in elephant studies of effective biochemical tools developed for insect investigations. This convergence of chemical signaling systems of elephants and insects has several interesting implications.

Stone, R., 1999. Paleontology. Siberian mammoth find raises hopes, questions. Science 286, 867-877.

Waddell, P.J., Cao, Y., Hauf, J., Hasegawa, M., 1999. Using novel phylogenetic methods to evaluate mammalian mtDNA, including amino acid-invariant sites-LogDet plus site stripping, to detect internal conflicts in the data, with special reference to the positions of hedgehog, armadillo, and elephant. Syst Biol 48, 31-53.
Abstract: We look at the higher-order phylogeny of mammals, analyzing in detail the complete mtDNA sequences of more than 40 species. We test the support for several proposed superordinal relationships. To this end, we apply a number of recently programmed methods and approaches, plus better-established methods. New pairwise tests show highly significant evidence that amino acid frequencies are changing among nearly all the genomes studied when unvaried sites are ignored. LogDet amino acid distances, with modifications to take into account invariant sites, are combined with bootstrapping and the Neighbor Joining algorithm to account for these violations of standard models. To weight the more slowlyevolving sites, we exclude the more rapidly evolving sites from the data by using "site stripping". This leads to changing optimal trees with nearly all methods. The bootstrap support for many hypotheses varies widely between methods, and few hypotheses can claim unanimous support from these data. Rather, we uncover good evidence that many of the earlier branching patterns in the placental subtree could be incorrect, including the placement of the root. The tRNA genes, for example, favor a split between the group hedgehog, rodents, and primates versus all other sequenced placentals. Such a grouping is not ruled out by the amino acid sequence data. A grouping of all rodents plus rabbit, the old Glires hypothesis, is also feasible with stripped amino acid data, and rodent monophyly is also common. The elephant sequence allows confident rejection of the older taxon Ferungulata (Simpson, 1945). In its place, the new taxa Scrotifera and Fereuungulata are defined. A new likelihood ratio test is used to detect differences between the optimal tree for tRNA versus that for amino acids. While not clearly significant as made, some results indicate the test is tending towards significance with more general models of evolution. Individual placement tests suggest alternative positions for hedgehog and elephant. Congruence arguments to support elephant and armadillo together are striking, suggesting a superordinal group composed of Xenarthra and African endemic mammals, which in turn may be near the root of the placental subtree. Thus, while casting doubt on some recent conclusions, the analyses are also unveiling some interesting new possibilities.

Zecchini, A., 1999. Life and death of species. Reconstituted animals? Courrier de la Nature 177, 22-27.
Abstract: This article gives an account of a breeding project, started in 1921, to reconstitute the aurochs, which became extinct in 1627, from existing breeds of cattle, and of a project to reconstitute the quagga (Equus quagga), which became extinct in 1921, from crosses among zebra species. The possibility of reconstituting the mammoth by obtaining semen from a mammoth preserved in the permafrost of Siberia and using it to fertilize elephant ova, followed by repeated backcrossing of hybrids to the mammoth, using mammoth semen, is discussed.

Daniel, J.C., 1998. The Asian elephant: a natural history. Natraj Publishers, Dehra Dun,  India.

Noro, M., Masuda, R., Dubrovo, I.A., Yoshida, M.C., Kato, M., 1998. Molecular phylogenetic inference of the woolly mammoth Mammuthus primigenius, based on complete sequences of mitochondrial cytochrome b and 12S ribosomal RNA genes. J Mol Evol 46, 314-326.
Abstract: Complete sequences of cytochrome b (1,137 bases) and 12S ribosomal RNA (961 bases) genes in mitochondrial DNA were successfully determined from the woolly mammoth (Mammuthus primigenius), African elephant (Loxodonta africana), and Asian elephant (Elephas maximus). From these sequence data, phylogenetic relationships among three genera were examined. Molecular phylogenetic trees reconstructed by the neighbor-joining and the maximum parsimony methods provided an identical topology both for cytochrome b and 12S rRNA genes. These results support the "Mammuthus-Loxodonta" clade, which is contrary to some previous morphological reports that Mammuthus is more closely related to Elephas than to Loxodonta.

Shoshani, J., 1998. Understanding proboscidean evolution: a formidable task. Trends in Ecology and Evolution 13, 480-487.

Stanhope, M.J., Madsen, O., Waddell, V.G., Cleven, G.C., De Jong, W.W., Springer, M.S., 1998. Highly congruent molecular support for a diverse superordinal clade of endemic African mammals. Mol Phylogenet Evol 9, 501-508.
Abstract: A solution to higher level mammalian phylogeny is going to depend on the congruent establishment of superordinal groupings followed by a linking together of these clades. We present congruent and convincing evidence from four disparate nuclear protein coding genes and from a tandem alignment of the 12S-16S mitochondrial region, for a superordinal clade of endemic African mammals that includes elephant shrews, aardvarks, golden mole, elephants, sirenians, and hyraxes. Because of strong support for golden mole as part of this clade, the Insectivora are rendered paraphyletic or polyphyletic, with constrained monophyly of the insectivores judged significantly worse in the vast majority of tests. Branching arrangement within this clade remains highly uncertain; however, a tandem alignment of the protein coding genes suggests elephant shrew is the earliest African lineage. None of the individual data sets or combinations of data sets support the widely held view of a mirorder Tethytheria (Sirenia/Proboscidea), although only a tandem alignment of protein coding and mitochondrial loci significantly rejects this association. The majority of the data sets and analyses provide strong support for Caviomorpha as part of a monophyletic Rodentia.

Kania, S.A., Richman, L.K., Kennedy, M., Montali, R.J., Potgleter, L.N.D., 1997. The isolation, detection, and cross-reactivity of Asian elephant IgG for the development of serological diagnostic tests. Journal of Veterinary Allergy and Clinical Immunology 5, 125-128.
Abstract: Asian elephant (Elephas maximus) IgG was isolated and purified using a recombinant protein A and proteins G affinity matrix and DEAE cibacron blue chromatography. Rabbits were inoculated with elephant IgG to produce anti-Asian elephant IgG. Using an ELISA, it was determined that the anti-Asian elephant sera has strong reactivity with Asian elephant IgG and African elephant (Loxodonta africana) IgG, moderate reactivity with manatee (Trichechus manatus latirostris) IgG and weaker reactivity with IgG from hyrax (Procavia capensis) and black rhinoceros (Diceros bicornis). Commercially available antisera produced against cattle, deer, and rabbit IgG react weakly with Asian elephant IgG.

Madsen, O., Deen, P.M., Pesole, G., Saccone, C., De Jong, W.W., 1997. Molecular evolution of mammalian aquaporin-2: further evidence that elephant shrew and aardvark join the paenungulate clade. Mol Biol Evol 14, 363-371.

Maliarchuk, B.A., 1997. Restriction analysis of the Enmynveem mammoth. Dokl Akad Nauk 353, 423-425.

Meng, J., Shoshani, J., Ketten, D., 1997. Evolutionary evidence for infrasonic sound and hearing in proboscideans. J. Vert. Paleo. 17, 64A-65A.

Ozawa, T., Hayashi, S., Mikhelson, V.M., 1997. Phylogenetic position of mammoth and Steller's sea cow within Tethytheria demonstrated by mitochondrial DNA sequences. J Mol Evol 44, 406-413.
Abstract: Here we report DNA sequences from mitochondrial cytochrome b gene segments (1,005 base pairs per species) for the extinct woolly mammoth (Mammuthus primigenius) and Steller's sea cow (Hydrodamalis gigas) and the extant Asian elephant (Elephas maximus), the Western Indian manatee (Trichechus manatus), and the hyrax (Procavia capensis). These molecular data have allowed us to construct the phylogeny for the Tethytheria. Our molecular data resolve the trichotomy between the two species of living elephants and the mammoth and confirm that the mammoth was more closely related to the Asian elephant than to the African elephant. Our data also suggest that the sea cow-dugong divergence was likely as ancient as the dugong-manatee split, and it appears to have been much earlier (22 million years ago) than had been previously estimated (4-8 million years ago) by immunological comparison.

Rasmussen, L.E., Lee, T.D., Zhang, A., Roelofs, W.L., Daves, G.D.Jr., 1997. Purification, identification, concentration and bioactivity of (Z)-7-dodecen-1-yl acetate: sex pheromone of the female Asian elephant, Elephas maximus. Chemical Senses 22, 417-437.
Abstract: In their natural ecosystems, adult male and female Asian elephants, Elephas maximus, live separately. For several weeks prior to ovulation, female elephants release a substance in their urine which elicits a high frequency of non-habituating chemosensory responses, especially flehmen responses, from male elephants. These responses occur prior to, and are an integral part of, mating. Using bioassay-guided fractionation, quantitatively dependent on these chemosensory responses, a specific sex pheromone was isolated and purified by an alternating series of organic and/or aqueous extractions, column chromatography, gas chromatography and high-performance liquid chromatography. Using primarily 1H-proton nuclear magnetic resonance (NMR) spectrometry and gas chromatography-mass spectrometry (GC-MS) of the urine-derived pheromone and its dimethyl disulfide derivative, we determined the structure of the active compound to be (Z)-7-dodecen-1-yl acetate (Z7-12:Ac). Concentrations of Z7-12:Ac in the female urine increased from non-detectable during the luteal phase to 0.48 microgram/ml (0.002 mM) early in the follicular phase and to 33.0 micrograms/ml (0.146 mM) just prior to ovulation. Bioassays with commercially available authentic synthetic Z7-12:Ac, using 10 Asian male elephants at several locations in the US, demonstrated quantitatively elevated chemosensory responses that were robust during successive tests, and several mating-associated behaviors. Bioassays with Z7-12:Ac with adult male elephants dwelling in more natural social situations in forest camps in Myanmar revealed some differing contextual pre-mating behavioral components. The remarkable convergent evolution of this compound suggests that compounds identified in mammalian exudates that are also present in pheromone blends of insects should be re-evaluated as potential mammalian chemosignals.

Rasmussen, L.E.L., Lee, T.D., Zhang, A., Roelofs, W.L., Daves, G.D., 1997. Purification, Identification, Concentration and Bioactivity of (Z)-7-Dodecen-1-yl Acetate: Sex Pheromone of the Female Asian Elephant, Elephas maximus. Chemical Senses 22, 417-437.
Abstract: In their natural ecosystems, adult male and female Asian elephants, Elephas maximus, live separately. For several weeks prior to ovulation, female elephants release a substance in their urine which elicits a high frequency of non-habituating chemosensory responses, especially flehmen responses, from male elephants. These responses occur prior to, and are an integral part of, mating. Using bioassay-guided fractionation, quantitatively dependent on these chemosensory responses, a specific sex pheromone was isolated and purified by an alternating series of organic and/or aqueous extractions, column chromatography, gas chromatography and high-performance liquid chromatography. Using primarily ¹H-proton nuclear magnetic resonance (NMR) spectrometry and gas chromatography-mass spectrometry (GC-MS) of the urine-derived pheromone and its dimethyl disulfide derivative, we determined the structure of the active compound to be (Z)-7-dodecen-1-yl acetate (Z7-12:Ac). Concentrations of Z7-12:Ac in the female urine increased from non-detectable during the luteal phase to 0.48 ug/ml (0.002 mM) early in the follicular phase and to 33.0 ug/ml (0.146mM) just prior to ovulation. Bioassays with commercially available authentic synthetic Z7-12:Ac, using 10 Asian male elephants at several locations in the US, demonstrated quantitatively elevated chemosensory responses that were robust during successive tests, and several mating-associated behaviors. Bioassays with Z7-12:Ac with adult male elephants dwelling in more natural social situations in forest camps in Myanmar revealed some differing contextual pre-matching behavioral components. The remarkable convergent evolution of this compound suggests that compounds identified in mammalian exudates that are also present in pheromone blends of insects should be re-evaluated as potential mammalian chemosignals.

Shoshani, J., 1997. Origins and evolution. In: Eltringham, M.A. (Ed.), The illustrated encyclopedia of elephants. Salamander Books Ltd., London, pp. 12-29.

Yang, H., Golenberg, E.M., Shoshani, J., 1997. Proboscidean DNA from museum and fossil specimens: an assessment of ancient DNA extraction and amplification techniques. Biochem Genet 35, 165-179.

Lavergne, A., Douzery, E., Stichler, T., Catzeflis, F.M., Springer, M.S., 1996. Interordinal mammalian relationships: evidence for paenungulate monophyly is provided by complete mitochondrial 12S rRNA sequences. Mol Phylogenet Evol 6, 245-158.
Abstract: The complete mitochondrial 12S rRNA sequences of 5 placental mammals belonging to the 3 orders Sirenia, Proboscidea, and Hyracoidea are reported together with phylogenetic analyses (distance and parsimony) of a total of 51 mammalian orthologues. This 12S rRNA database now includes the 2 extant proboscideans (the African and Asiatic elephants Loxodonta africana and Elephas maximus), 2 of the 3 extant sirenian genera (the sea cow Dugong dugon and the West Indian manatee Trichechus manatus), and 2 of the 3 extant hyracoid genera (the rock and tree hyraxes Procavia capensis and Dendrohyrax dorsalis). The monophyly of the 3 orders Sirenia, Proboscidea, and Hyracoidea is supported by all kinds of analysis. There are 23 and 3 diagnostic substitutions shared by the 2 proboscideans and the 2 hyracoids, respectively, but none by the 2 sirenians. The 2 proboscideans exhibit the fastest rates of 12S rRNA evolution among the 11 placental orders studied. Based on various taxonomic sampling methods among eutherian orders and marsupial outgroups, the most strongly supported clade in our comparisons clusters together the 3 orders Sirenia, Proboscidea, and Hyracoidea in the superorder Paenungulata. Within paenungulates, the grouping of sirenians and proboscideans within the mirorder Tethytheria is observed. This branching pattern is supported by all analyses by high bootstrap percentages (BPs) and decay indices. When only one species is selected per order or suborder, the taxonomic sampling leads to a relative variation in bootstrap Paernungulata (92-99%). When each order or suborder is represented by two species, this relative variation decreased to 10% for Tethytheria (78-87%) and 3% for Paenungulata (96-99%). Two nearly exclusive synapomorphies for paenungulates are identified in the form of one transitional compensatory change, but none were detected for tethytherians. Such a robust and reliable resolution of the paenungulate node implies a long history of the common ancestors, allowing time for synapomorphies to accumulate. This observation suggests a Late Cretaceous/Early Paleocene origin for the Paenungulata.

Maliarchuk, B.A., Derevenko, M.V., Lapinskii, A.G., Solovenchuk, L.L., 1996. The use of the polymerase chain reaction in analyzing ancient DNAs (exemplified by that of the Enmynveem mammoth). Izv Akad Nauk Ser Biol 6, 681-686.
Abstract: DNA was isolated from the Enmynveem mammoth muscles, and the control region and cytochrome b gene of the mitochondrial genome were analyzed by polymerase chain reaction (PCR). The mammoth DNA was amplified by both the classical PCR (two primer system) and the single-primer PCR (spPCR) resulting in DNA fragments up to 1600 bp long. Restriction analysis of the mitochondrial cytochrome b gene was carried out. Cytochromes b genes in three Elephantidae genera were compared.

Poole, J., 1996. Coming of Age with Elephants: a Memoir. Hodder and Stoughton, New York.

Porter, C.A., Goodman, M., Stanhope, M.J., 1996. Evidence on mammalian phylogeny from sequences of exon 28 of the von Willebrand factor gene. Mol Phylogenet Evol 5, 89-101.
Abstract: Phylogenetic relationships among 27 extant mammalian species (representing 15 placental orders) were studied using sequences of exon 28 of the gene encoding von Willebrand Factor (vWF), a glycoprotein which functions in blood clotting. Analysis of sequences coding for vWF revealed evidence for several subordinal and superordinal groupings, but the earliest branching sequence of placental mammals was left largely unresolved. Strong support was found for a monophyletic clade consisting of elephants, sea cows, hyraxes, aardvarks, and elephant shrews. This systematic placement of the elephant shrews agrees strongly with two other molecular data sets (interphotoreceptor retinoid binding protein and alpha-lens crystallins) and is consistent with analysis of fossil elephant shrews recently discovered in north Africa. Evidence from vWF sequences agrees with a number of previous molecular and morphological studies in providing strong support for the monophyly of both bats and rodents. The orders Primates, Proboscidea, Carnivora, Perissodactyla, and Artiodactyla were represented by more than one species which joined in each case to form a monophyletic order.

Stanhope, M.J., Smith, M.R., Waddell, V.G., Porter, C.A., Shivji, M.S., Goodman, M., 1996. Mammalian evolution and the interphotoreceptor retinoid binding protein (IRBP)gene: convincing evidence for several superordinal clades. J Mol Evol 43, 83-92.
Abstract: Phylogenetic relationships of 25 mammalian species representing 17 of the 18 eutherian orders were examined using DNA sequences from a 1.2-kb region of the 5' end of exon 1 of the single-copy nuclear gene known as interphotoreceptor retinoid binding protein (IRBP). A wide variety of methods of analysis of the DNA sequence, and of the translated products, all supported a five-order clade consisting of elephant shrew (Macroscelidea)/aardvark (Tubulidentata)/and the paenungulates (hyracoids, sirenians, and elephants), with bootstrap support in all cases of 100%. The Paenungulata was also strongly supported by these IRBP data. In the majority of analyses this monophyletic five-order grouping was thefirst branch off the tree after the Edentata. These results are highly congruent with two other recent sources of molecular data. Another superordinal grouping, with similar 100% bootstrap support in all of the same wide-ranging types of analyses, was Artiodactyla/Cetacea. Other superordinal affinities, suggested by the analyses, but with less convincing support, included a Perissodactyla/Artiodactyla/Cetacea clade, an Insectivora/Chiroptera clade, and Glires (an association of rodents and lagomorphs).

Yang, H., Golenberg, E.M., Shoshani, J., 1996. Phylogenetic resolution within the Elephantidae using fossil DNA sequence from the American mastodon (Mammut americanum) as an outgroup. Proc Natl Acad Sci U S A 93, 1190-1194.
Abstract: DNA was extracted from the extinct American mastodon, the extinct woolly mammoth, and the modern Asian and African elephants to test the traditional morphologically based phylogeny within Elephantidae. Phylogenetic analyses of the aligned sequences of the mitochondrial gene cytochrome b support a monophyletic Asian elephant-woolly mammoth clade when the American mastodon is used as an outgroup. Previous molecular studies were unable to resolve the relationships of the woolly mammoth, Asian elephant, and African elephant because the sequences appear to have evolved at heterogeneous rates and inappropriate outgroups were used for analysis. The results demonstrate the usefulness of fossil molecular data from appropriate sister taxa for resolving phylogenies of highly derived or early radiating lineages. Erratum in: Proc Natl Acad Sci U S A 1996 Apr 30;93(9):4519

Barua, P., Bist, S.S., 1995. Changing Patterns in the Distribution and Movement of Wild Elephants in North Bengal. In: Daniel, J.C. (Ed.), A Week with Elephants; Proceedings of the International Seminar on Asian Elephants. Bombay Natural History Society; Oxford University Press, Bombay, India, pp. 66-84.

Boomker, J., Bain, O., Chabaud, A., Kriek, N.P.J., 1995. Stephanofilaria thelazioides n. sp. (Nematoda: Filariidae) from a hippopotamus and its affinities with the species parasitic in the African black rhinoceros. Systematic Parasitology 32, 205-210.
Abstract: Stephanofilaria thelazioides sp. nov. is described and figured from an ulcerated skin lesion on a hippopotamus Hippopotamus amphibius from the Kruger National Park, South Africa. This nematode is closely related to S. dinniki, a parasite of the black rhinoceros Diceros bicornis in Africa, but differs from it in the number of cuticular spines surrounding the mouth, the arrangement of the cloacal papillae and the measurements of the spicules, gubernaculum and microfilariae. Species of the genus Stephanofilaria possess spines on the head which have been derived by modification of the sensory papillae. S. thelazioides is the most primitive species of the genus and has the least modified arrangement of these papillae, with 6 bifid internal labial spines, 4 bifid external labial spines and 4 cephalic papillae. The genus appears to have diversified in various mammals which have in common a thick skin, such as rhinoceroses, elephants, buffaloes and now the hippopotamus. It appears to have become adapted secondarily to domestic bovines, initially in Asia and subsequently in North America.

Hauf, J., Baur, A., Chalwatzis, N., Zimmermann, F.K., Joger, U., Lazarev, P.A., 1995. Selective amplification of a mammoth mitochondrial cytochrome b fragment using an elephant-specific primer. Curr Genet 27, 486-487.

Jayewardene, J., 1995. An Overview of Elephant Conservation in Sri Lanka. In: Daniel, J.C. (Ed.), A Week with Elephants; Proceedings of the International Seminar on Asian Elephants. Bombay Natural History Society; Oxford University Press, Bombay, India, pp. 217-224.

Krishnamurthy, V., Wemmer, C. Veterinary Care of Asian Timber Elephants in India: Historical Accounts and Current Observations.  534. 1995. Bombay, India, Bombay Natural History Society; Oxford University Press.
Ref Type: Abstract

Krishnamurthy, V., Wemmer, C., 1995. Timber Elephant Management in the Madras Presidency of India (1844-1947). In: Daniel, J.C. (Ed.), A Week with Elephants; Proceedings of the International Seminar on Asian Elephants. Bombay Natural History Society; Oxford University Press, Bombay, India, pp. 456-472.

Masson, J.M., McCarthy, S., 1995. When elephants weep : the emotional lives of animals. Delacorte, New York.

Mezhzherin, S.V., Morozov-Leonov, S.I., 1995. The genetic differentiation of mammalian taxa: their assessment by biochemical genetic markers. Zh Obshch Biol 56, 71-96.
Abstract: A review of data on genetic differentiation of mammalian taxa has been made on the basis of estimating the percent of fixed gene differences (PFD). The results substantiate the existence of evident differences in the scale of genetic divergence between taxa in different mammalian orders. Among smaller mammals (marsupials, insectivores, chiropterans, myomorph and sciuromorph rodents, african mole rats, and elephant shrews) interspecific differences within a genus involve the average of 25-40% of investigated loci. At the genetic level the value is 50-60%, whereas at the familial level the differences are beyond the resolution capacity of the method (PFD = 60-80%). Orders of larger mammals can be divided into two subgroups. One of them that includes carnivores, artiodactylans, and hystricomorph rodents is characterized by PFD values of 10-14%, 30-50%, and 69-70% at respective levels. The other subgroup composed of proboscideans, primates, pinnipeds, and toothed whales, has a low level of
genetic divergence expressed by PFD values of 0-3%, 7-36%, and 50-60% at species, generic and familial levels, respectively. Insufficiency of data on baleen whales and Perissodactyla does not allow to cluster them ultimately with any of these groups. There are three possible, but not necessarily alternative, causes for the observed differences in genetic divergence: 1) over-ranking of genera in larger mammals; 2) different paleontological age of orders; 3) unequal rates of molecular evolution.

Queralt, R., Adroer, R., Oliva, R., Winkfein, R.J., Retief, J.D., Dixon, G.H., 1995. Evolution of protamine P1 genes in mammals. Journal of Molecular Evolution 40, 601-607.
Abstract: A polymerase chain reaction-based approach was used to amplify and sequence the protamine genes of the rat, guineapig (order Rodentia), cat, bear (Carnivora), elephant (Proboscidea), horse (Perissodactyla), camel, elk, deer, moose and gazelle (Artiodactyla). The predicted amino acid sequences for these genes, together with previously reported amino acid sequences of protamine genes of humans, mice, pigs, cattle, fowls, quails and opossums, resulted in a data set of 25 P1 genes and 30 P1 amino acid sequences. A bootstrapped DNA parsimony tree of the set of protamine sequences was constructed. The results showed that protamines were amongst the most rapidly diverging proteins studied. In spite of the large differences, there were conserved motifs that were also common to birds. The C-terminus appeared to be the most variable region. The molecular evolution of P1 genes was in agreement with the expected species evolution.

Wemmer, C., 1995. Gaonbura Sahib - A.J.W. Milroy of Assam. In: Daniel, J.C. (Ed.), A Week with Elephants; Proceedings of the International Seminar on Asian Elephants. Bombay Natural History Society; Oxford University Press, Bombay, India, pp. 483-496.

Zhao, X., Vyas, K., Nguyen, B.D., Rajarathnam, K., La Mar, G.N., Li, T., Phillips, G.N., Jr., Eich, R.F., Olson, J.S., Ling, J., 1995. A double mutant of sperm whale myoglobin mimics the structure and function of elephant myoglobin. J Biol Chem 270, 20763-20764.
Abstract: The functional, spectral, and structural properties of elephant myoglobin and the L29F/H64Q mutant of sperm whale myoglobin have been compared in detail by conventional kinetic techniques, infrared and resonance Raman spectroscopy, 1H NMR, and x-ray crystallography. There is a striking correspondence between the properties of the naturally occurring elephant protein and those of the sperm whale double mutant, both of which are quite distinct from those of native sperm whale myoglobin and the single H64Q mutant. These results and the recent crystal structure determination by Bisig et al. (Bisig, D. A., Di Iorio, E. E., Diederichs, K., Winterhalter, K. H., and Piontek, K. (1995) J. Biol. Chem. 270, 20754-20762) confirm that a Phe residue is present at position 29 (B10) in elephant myoglobin, and not a Leu residue as is reported in the published amino acid sequence. The single Gln64(E7) substitution lowers oxygen affinity approximately 5-fold and increases the rate of autooxidation 3-fold. These unfavorable effects are reversed by the Phe29(B10) replacement in both elephant myoglobin and the sperm whale double mutant. The latter, genetically engineered protein was originally constructed to be a blood substitute prototype with moderately low O2 affinity, large rate constants, and increased resistance to autooxidation. Thus, the same distal pocket combination that we designed rationally on the basis of proposed mechanisms for ligand binding and autooxidation is also found in nature.

Georgiadis, N., Bischof, L., Templeton, A., Patton, J., Karesh, W., Western, D., 1994. Structure and history of African elephant populations: I. Eastern and southern Africa. J Hered 85, 100-104.
Abstract: Patterns of restriction site variation within mitochondrial DNA (mtDNA) of 270 individuals were used to examine the current structure of savanna elephant populations and to infer historical patterns of gene flow across eastern and southern Africa. Elephants have a complex population structure characterized by marked subdivision at the continental level (Fst = 0.39; 95% confidence interval 0.19-0.58), and isolation by distance at the regional level. However, phylogeographic analysis revealed evidence of protracted gene flow across the continent. First, one relatively derived haplotype was found at all sampling locations. Second, haplotypes representing exceptionally divergent (up to 8.3%) mitochondrial clades were found to coexist at distant (> 2,000 km) sampling locations. In the few other species characterized by sympatric individuals bearing such divergent haplotypes, all such individuals were found to coexist within limited geographical regions. Accordingly, pronounced mitochondrial divergence within populations is often attributed to ancestral isolation in allopatry, followed by secondary contact. The patterns within elephants do not accord with ancestral isolation in allopatry. Given the exceptional mobility of elephants, a geographical barrier is unlikely to have obstructed gene flow between regions for long enough to produce the observed mitochondrial divergence. Rather, the patterns are consistent with the more parsimonious hypothesis, based on neutral coalescent theory, that gene flow has maintained a sufficiently large effective population size (> 50,000 females) for representatives of clades that diverged at least 4 million years ago to have persisted by chance within a population that was subdivided, but not strictly isolated in allopatry.

Hagelberg, E., Thomas, M.G., Cook, C.E.Jr., Sher, A.V., Baryshnikov, G.F., Lister, A.M., 1994. DNA from ancient mammoth bones. Nature 370, 333-334.

Hart, L.A., 1994. The Asian elephants-driver partnership: the drivers'perspective. Applied Animal Behaviour Science 40, 297-312.

Hoss, M., Paabo, S., Vereshchagin, N.K., 1994. Mammoth DNA sequences. Nature 370, 333.

Jayewardene, J., 1994. The Elephant in Sri Lanka. WHT Publications Ltd., Colombo, Sri Lanka.

Armbrusters, P., Lande, R., 1993. A population viability analysis for African elephant (Loxodonta africana): How big should reserves be? Conservation Biology 7, 602-610.
Abstract: We present an age-structured, density-dependent model of elephant population dynamics in a fluctuating environment drawing primarily upon the life history parameters obtained from studies in semi-arid land at Tsavo National Park, Kenya. Density regulation occurs by changes in the age of first reproduction and calving interval. We model environmental stochasticity with drought events affecting sex- and age-specific survivorships. Results indicate a maximum population growth rate of 3% per year and an equilibrium elephant density of 3.1/square mile. Analysis of the demographic results and their sensitivity to changes in juvenile survivorship and drought frequencies, supported by genetic considerations, suggests that in semi-arid regions a minimum reserve size of 1000 square mile is necessary to attain a 99% probability of population persistence for 1000 years. The effect of age-independent culling on population viability is also analyzed.

Garutt, V.E., Aver'ianov, A.O., Vartanian, S.L., 1993. The systematic position of the holocene population of the mammoth, Mammuthus primigenius (Blumenbach, 1799), of the Vrangel Island (Northeast Siberia). Dokl Akad Nauk 332, 799-801.

Ma, D.P., Zharkikh, A., Graur, D., VandeBerg, J.L., Li, W.H., 1993. Structure and evolution of opossum, guinea pig, and porcupine cytochrome b genes. J Mol Evol 36, 327-334.
Abstract: We have sequenced the mitochondrial cytochrome b gene from the guinea pig, the African porcupine, and a South American opossum. A phylogenetic analysis, which includes 22 eutherian and four other vertebrate cytochrome b sequences, indicates that the guinea pig and the porcupine constitute a natural clade (Hystricomorpha) that is not a sister group to the clade of mice and rats (Myomorpha). Therefore, the hypothesis that the Rodentia is paraphyletic receives additional support. The artiodactyls, the perissodactyls, and the cetaceans form a group that is separated from the primates and the rodents. The 26 sequences are used to study the structure/function relationships in cytochrome b, whose function is electron transport. Most of the amino acid residues involved in the two reaction centers are well conserved in evolution. The four histidines that are believed to ligate the two hemes are invariant among the 26 sequences, but their nearby residues are not well conserved in
evolution. The eight transmembrane domains represent some of the most divergent regions in the cytochrome b sequence. The rate of nonsynonymous substitution is considerably faster in the human and elephant lineages than in other eutherian lineages; the faster rate might be due to coevolution between cytochrome b and cytochrome c.

 1992. Elephants. Majestic creatures of the wild. Rodale Press, Emmaus, Pennsylvania.

Karunakaran, R., 1992. The Riddle of Ganesha. Book Quest, Bombay, India.

Schaedler, J.M., Krook, L., Wootton, J.A., Hover, B., Brodsky, B., Naresh, M.D., Gillette, D.D., Madsen, D.B., Horne, R.H., Minor, R.R., 1992. Studies of collagen in bone and dentin matrix of a Columbian mammoth (late Pleistocene) of central Utah. Matrix 12, 297-307.
Abstract: A Columbian mammoth, Mammuthus columbi, was excavated at an elevation of 9000 feet in Huntington Canyon, Emery County, Utah. Radiocarbon dates on the skeleton indicated death approximately 11,200 years ago. The skeleton was removed from postglacial, Late Quaternary, lake sediments deposited as glacial runoff approximately 9500 years ago. The bones and teeth were especially well preserved in a saturated lake bed. After excavation the bones and teeth were preserved by controlled desiccation, without hardeners, over a period of 9 months. Microradiography, light and electron microscopy, medium and high angle X-ray diffraction, amino acid analysis and cyanogen bromide peptide mapping were undertaken to evaluate the packing, organization, and preservation of collagen in bone and dentin of this mammoth. Microradiography and light microscopy showed that the bone consisted of especially well preserved compact and trabecular bone, and electron microscopy of demineralized bone and tusk showed that the matrix consisted of lamellae of densely packed cylindrical collagen fibrils. Cell remnants with intact nuclei, with or without a nucleolus, as well as variable lengths of plasma membrane were occasionally present on the surface of bony trabecula. Remnants of odontoblast processes were present in some dentin tubules. High and low angle X-ray diffraction demonstrated that the demineralized matrix contained native collagen molecules and amino acid analysis showed that the composition was comparable to that of type I collagen. Cyanogen bromide peptide mapping indicated that the major peptides of type I collagen were present and had the same electrophoretic mobility as that of type I collagen of demineralized Asian elephant bone and rat tail tendon. Abstract truncated at 250 words.

Wylie, K.C., 1992. Elephants used as war machines. In: Shoshani, J. (Ed.), Elephants. Majestic creatures of the wild. Rodale Press, Emmaus, Pennsylvania, pp. 146-148.

Haynes, G., 1991. Mammoths, mastodonts, and elephants : biology, behavior, and the fossil record. Cambridge University Pres, Cambridge ; New York.

Irwin, D.M., Kocher, T.D., Wilson, A.C., 1991. Evolution of the cytochrome b gene of mammals. J Mol Evol 32, 128-144.
Abstract: With the polymerase chain reaction (PCR) and versatile primers that amplify the whole cytochrome b gene (approximately 1140 bp), we obtained 17 complete gene sequences representing three orders of hoofed mammals (ungulates) and dolphins (cetaceans). The fossil record of some ungulate lineages allowed estimation of the evolutionary rates for various components of the cytochrome b DNA and amino acid sequences. The relative rates of substitution at first, second, and third positions within codons are in the ratio 10 to 1 to at least 33. For deep divergences (greater than 5 million years) it appears that both replacements and silent transversions in this mitochondrial gene can be used for phylogenetic inference. Phylogenetic findings include the association of (1) cetaceans, artiodactyls, and perissodactyls to the exclusion of elephants and humans, (2) pronghorn and fallow deer to the exclusion of bovids (i.e., cow, sheep, and goat), (3) sheep and goat to the exclusion of other pecorans (i.e., cow, giraffe, deer, and pronghorn), and (4) advanced ruminants to the exclusion of the chevrotain and other artiodactyls. Comparisons of these cytochrome b sequences support current structure-function models for this membrane-spanning protein. That part of the outer surface which includes the Qo redox center is more constrained than the remainder of the molecule, namely, the transmembrane segments and the surface that protrudes into the mitochondrial matrix. Many of the amino acid replacements within the transmembrane segments are exchanges between hydrophobic residues (especially leucine, isoleucine, and valine). Replacement changes at first and second positions of codons approximate a negative binomial distribution, similar to other protein-coding sequences. At four-fold degenerate positions of codons, the nucleotide substitutions approximate a Poisson distribution, implying that the underlying mutational spectrum is random with respect to position.

Norton, B., 1991. The African elephant :last days of Eden. Voyageur Press, Stillwater, MN, U.S.A.

Nowak, R.M., 1991. Order Proboscidea. Walker's Mammals of the World. The Johns Hopkins University Press, Baltimore, pp. 1277-1286.

Sondaar, P.Y., 1991. Island mammals of the past. Sci Prog 75, 249-264.
Abstract: The effect of insularity on mammal evolution is approached by considering endemic fossil mammals of islands in the Mediterranean. It is shown that in general the same mammals will arrive on the islands by sweepstake dispersal. Among the large mammals this includes elephants, hippopotamids and deer: all mammals known for their good swimming ability. The evolutionary changes these mammals underwent follow parallel patterns on different islands and can be explained as an adaptational process to an island environment lacking large carnivores.

Altevogt, R., Thenius, E., Kurt, F., Grzimek, B., 1990. Proboscideans. Grzimek's Encyclopedia. McGraw-Hill, Inc., pp. 460-520.

Fischer, M.S., 1990. The unique ear of elephants and manatees (Mammalia): A phylogenetic paradox. C. R. Acad. Sci. Ser. III Sci. Vie 311, 157-162.

Rasmussen, D.T., Gagnon, M., Simons, E.L., 1990. Taxeopody in the carpus and tarsus of Oligocene Pliohyracidae (Mammalia:Hyracoidea) and the phyletic position of hyraxes. Proc Natl Acad Sci U S A 87, 4688-4691.
Abstract: Recent hyracoids and elephants share a taxeopode arrangement of tarsal and carpal bones, a condition in which bones are aligned with minimal interlocking between adjacent elements. Taxeopody has often been interpreted as a synapomorphy reflecting a close phyletic link between Hyracoidea and Proboscidea, but recently it has been suggested [Fischer, M. S. (1986) Cour. Forschungsinst. Senckenberg 84, 1-132] that hyracoid taxeopody is an independent acquisition resulting from selection favoring increased midcarpal and midtarsal rotation and that Hyracoidea is actually allied with Perissodactyla. As a test of this hypothesis, isolated carpal and tarsal bones of primitive Oligocene hyracoids from the Fayum, Egypt, have been examined to determine whether these indicate a taxeopode or diplarthral carpus and tarsus. Four complete astragali from the Fayum, representing at least three taxa, show a single, slightly convex articular surface on the head for articulation with the navicular and lack a facet for the cuboid. Two complete magna representing two species have a single proximal facet for articulation with the lunar, and they lack a facet for the scaphoid. Thus, both the carpus and tarsus of Fayum hyracoids are taxeopode. Taxeopody in hyracoids cannot be attributed to selection for carpal and tarsal rotation in climbers because the Oligocene, Miocene, and Recent species show great diversity in body size and probably locomotor specializations, despite relative uniformity of structure in the carpus and tarsus. The shared taxeopody of hyracoids and proboscideans, along with other osteological characters and similarities in hemoglobin, eye lens proteins, and other molecules, all suggest that Hyracoidea belongs within Paenungulata.

Yu, L.P., La Mar, G.N., Mizukami, H., 1990. Rearrangement of the distal pocket accompanying E7 His----Gln substitution in elephant carbonmonoxy- and oxymyoglobin: ¹H NMR identification of a new aromatic residue in the heme pocket. Biochemistry 29, 2578-2585.
Abstract: Two-dimensional 1H NMR methods have been used to assign side- chain resonances for the residues in the distal heme pocket of elephant carbonmonoxymyoglobin (MbCO) and oxymyoglobin (MbO2). It is shown that, while the other residues in the heme pocket are minimally perturbed, the Phe CD4 residue in elephant MbCO and MbO2 resonates considerably upfield compared to the corresponding residue in sperm whale MbCO. The new NOE connectivities to Val E11 and heme-induced ring current calculations indicate that Phe CD4 has been inserted into the distal heme pocket by reorienting the aromatic side chain and moving the CD corner closer to the heme. The C zeta H proton of the Phe CD4 was found to move toward the iron of the heme by approximately 4 A relative to the position of sperm whale MbCO, requiring minimally a 3-A movement of the CD helical backbone. The significantly altered distal conformation in elephant myoglobin, rather than the single distal E7 substitution, forms a plausible basis for its altered functional properties of lower autoxidation rate, higher redox potential, and increased affinity for CO ligand. These results demonstrate that one-to-one interpretation of amino acid residue substitution (E7 His----Gln) is oversimplified and that conformational changes of substituted proteins which are not readily predicted have to be considered for interpretation of their functional properties

Chong, T.S., Ohta, H., Nakashima, Y., Iida, T., Saisho, H., 1989. ESR dating of elephant teeth and radiation dose rate estimation in soil. Int J Rad Appl Instrum [A] 40, 1199-1202.
Abstract: Chemical analysis of 238U, 232Th and 40K in the dentine as well as enamel of elephant tooth fossil has been carried out in order to estimate the internal absorbed dose rate of the specimens, which was estimated to be (39 +/- 4) mrad/y on the assumption of early uptake model of radionuclides. The external radiation dose rate in the soil including the contribution from cosmic rays was also estimated to be (175 +/- 18) mrad/y with the help of gamma-ray spectroscopic techniques of the soil samples in which the specimens were buried. The 60Co gamma-ray equivalent accumulated dose of (2 +/- 0.2) x 10(4) rad for the tooth enamel gave "ESR age" of (9 +/- 2) x 10(4) y, which falls in the geologically estimated range between 3 x 10(4) and 30 x 10(4) y before the present.

Sukumar, R., 1989. The historical background. The Asian Elephant: Ecology and Management. Cambridge University Press, Cambridge, pp. 1-9.

Hallett, H.S., 1988. A Thousand Miles on an Elephant in the Shan States. White Lotus Co., Ltd., Bangkok, Thailand.
Abstract: Note: This book, first published in 1890, resulted from Hallett's thorough fact-finding trip through northern Thailandin 1876 in search of the best route for a railway by which British goods would be transported from Burma to Thailand. It presents an excellent oerview of the topography, economy, peoples and customs, legends and local histories of nothern Thailand in the late 19th century.

Moss, C., 1988. Elephant memories. Fawcett Columbine (Ballantine), New York.

Terkel, A.S., 1988. 1987 -- the year of the elephant in Ramat-Gan. International Zoo News 35, 13-18.

 1986. New species of bacteria in the genus Kurthia--Kurthia sibirica sp. nov. Mikrobiologiia 55, 831-835.
Abstract: Six aerobic gram-positive nonspore-forming bacterial strains belonging to the Kurthia genus were isolated from the Magadan (Susuman) mammoth found in the permafrost of the East Siberia. The strains are a phenotypically homogeneous group different from the two known species (K. zopfii and K. gibsonii) in requiring more vitamins, the absence of growth in a medium with 7% NaCl, and a low level of DNA-DNA hybridization (not more than 45%). Moreover, the strains differ from K. zopfii in the synthesis of a yellow pigment, the activity of phosphatase, and the absence of coccoid forms; the bacteria differ from K. gibsonii in the absence of growth at a temperature above 40 degrees C. The organisms are referred to as Kurthia sibirica sp. nov. The type strain 13-2 has been deposited in the All-Union Collection of Microorganisms as strain VKB B-1549.

Kleinschmidt, T., Czelusniak, J., Goodman, M., Braunitzer, G., 1986. Paenungulata: a comparison of the hemoglobin sequences from elephant, hyrax, and manatee. Molecular Biology and Evolution 3, 427-435.
Abstract: Inspection of the amino acid differences among hemoglobin sequences of a wide range of mammalian species suggested that at alpha 19, alpha 110, alpha 111, beta 23, beta 44, and beta 56, synapomorphies group manatee (Trichechus inungius, Sirenia), Indian and African elephant (Elephas maximus and Loxodonta africana, Proboscidea), and rock hyrax (Procavia habessinica, Hyracoidea) into a monophyletic clade. Results obtained by parsimony analysis provide evidence for this grouping--and thus support for the genealogical validity of Simpson's superorder Paenungulata, which contains as the extant orders Proboscidea, Sirenia, and Hyracoidea. All of the 39 most, or nearly most, parsimonious of 10,395 trees constructed from a tandemly combined alpha- and beta-hemoglobin sequence for 103 vertebrate species (of which 79 were mammals from 16 extant orders), depicted Paenungulata as one of the most anciently separated branches of Eutheria. It was found on examining thousands of alternative trees that to not group Proboscidea, Hyracoidea, and Sirenia in a monophyletic clade required at least four additional substitutions.

Mariappa, D., 1986. Anatomy and Histology of the Indian Elephant. Indira Publishing House, Oak Park, MI.
Abstract: This book deals with all aspects of gross anatomy of the Indian elephant. The chapter on histology covers 35 organs. Apart from detailed information on the anatomy and histology of the Indian elephant, the book provides information on many other species of mammals to cater to the needs of veterinarians and comparative anatomists. contemporary thoughts on phylogeny of elephants are also discussed in the introduction.

Yu, N.T., Thompson, H.M., Mizukami, H., Gersonde, K., 1986. The cobalt-nitrosyl stretching vibration as a sensitive resonance Raman probe for distal histidine-nitrosyl interaction in monomeric hemoglobins. Eur. J Biochem. 159, 129-132.
Abstract: The Co-NO stretching vibration has been assigned in the resonance Raman spectra of various cobalt-substituted monomeric hemoglobins by employing isotope-labeling of nitrosyl (14N16O, 15N16O, 14N18O). Monomeric hemoglobins with a distal histidine (sperm whale myoglobin and leghemoglobin) exhibit this vibration at 573-575 cm-1, whereas hemoglobins without distal histidine (elephant myoglobin and insect hemoglobin from Chironomus thummi thummi, CTT III) show this vibration in the range of 553-558 cm- 1. The Fe-NO stretching vibration which occurs in the range of 554-556 cm-1 does not reflect the distal histidine-ligand interaction. Therefore, the Co-NO moiety which is isoelectronic with the Fe-O2 moiety is a good monitor for distal effects on the exogenous ligand of hemoglobins, especially due to the fact that in hemoglobins with distal histidine the Fe-O2 stretching vibration (567-572 cm-1) is similar to the Co-NO stretching vibration

Johnson, P.H., Olson, C.B., Goodman, M., 1985. Isolation and characterization of deoxyribonucleic acid from tissue of the woolly mammoth, Mammuthus primigenius. Comp Biochem Physiol B 81, 1045-1051.
Abstract: DNA was isolated from tissue samples of several mammoth specimens, radiocarbon dated between 10,000 and 53,000 years old. The DNA was purified by chromatography on hydroxyapatite at 60 degrees C and was characterized as a heterogeneous population of fragments ranging in size from 3000 to 200 base pairs. Thermal denaturation analysis demonstrated that approximately 25% of the DNA had a base composition similar to Asian elephant DNA calculated as 36% G + C. Preliminary analysis by nucleic acid hybridization indicated that only a small fraction of DNA isolated from mammoth tissue (2-5%) was homologous to DNA of Asian elephant, a close living relative of the mammoth. Our results provide the first definitive isolation and characterization of DNA from ancient tissue and suggest a purification strategy that will lead to preparations of DNA from mammoth tissue significantly enriched in elephant-related DNA sequences.

Douglas-Hamilton, I., 1984. African elephant. In: Mason, I.L. (Ed.), Evolution of domesticated animals. Longman, London, pp. 193-198.

Mikhailov, M.I., Kuznetsov, S.V., Zhdanov, V.M., 1984. Electron microscopy of the intestinal content of a mammoth. The Lancet 2, 111-112.

Olivier, R.C.D., 1984. Asian elephant. In: Mason, I.L. (Ed.), Evolution of domesticated animals. Longman, London, pp. 185-193.

Baker, C.M.A., Manwell, C., 1983. Man and elephant.  The "dare theory" of domestication and the origin of breeds. Zeit. Tierzucht. Zuchtungscbiol. 100, 55-75.

Gibbs, S.J., Heller, R.M., Sloan, M., James, A.E., Jr., 1983. External root resorption in mastodon molar. Oral Surg Oral Med Oral Pathol 55, 542.

Kleinschmidt, T., Braunitzer, G., 1983. The primary structure of hemoglobins of the rock hyrax (Procavia habessinica, Hyracooidea): insertion of glutamine in the alpha chains. Hoppe. Seylers. Z. Physiol. Chem. 364, 1303-1313.
Abstract: The chromatography of the hemoglobin of the rock hyrax (Procavia habessinica) gives two components (73% HbI and 27% HbII). The amino-acid analysis and the sequences of the globin chains elucidated with the phenylthiohydantoin method, did not show any differences between the alpha I and alpha II or beta I and beta II chains, respectively. The different chromatographical behaviour cannot be explained. After chain separation by chromatography on CM-52 cellulose, all four primary structures were elucidated automatically in a sequenator on the chains and the tryptic peptides. In 20% of the beta I chains the N-terminal valine was blocked by acetyl. The alignment was performed by homology with the chains of human adult hemoglobin. The alpha chain of the rock hyrax has 142 amino-acid residues, i.e. one residue more than normal mammalian alpha chains, caused by an insertion of glutamine in the GH region supposed between positions 115 and 116. A comparison of human and hyrax hemoglobins shows an exchange of 21 amino-acid residues in the alpha chains and of 24 in the beta chains. Some substitutions in alpha 1 beta 1 contacts and in the surrounding of the heme are not supposed to effect the function of the hemoglobin. The phylogenetic relationship between the rock hyrax and the Indian elephant (Elephas maximus) on the one hand and with some Perissodactyla on the other, is discussed. Up to now the exchanges of alpha 110(G17)Ala leads to Ser and beta 56(D7)Gly leads to His have only been found in hyrax and elephant. This indicates a certain relationship between Hyracoidea and Proboscidea.

Valente, A., 1983. Hair structure of the woolly mammoth, Mammuthus primigenius and the modern elephants, Elephas maximus and Loxodonta africanus. Journal of Zoology (Lond) 199, 271-274.
Abstract: The structure of overhairs from a Wooly mammoth, Mammuthus primigenius, 10-13 thousand years old,is compared with that of its living relatives, Elephas maximus and Loxodonta africana.  The hair profile, cross-sectional appearance, wholemount and cuticular scale cast of the hairs of the three species were examined and a selected array of photographs representing the hair structure of each species is presented.  In general there is little variation between the three species in the gross structure of the overhairs.

Eltringham, S.K., 1982. Elephants. Blandfort Books,Ltd., United Kingdom.

De Jong, W.W., Zweers, A., Goodman, M., 1981. Relationship of aardvark to elephants, hyraxes and sea cows from alpha-crystallin sequences. Nature 292, 538-540.

Romero-Herrera, A.E., Goodman, M., Dene, H., Bartnicki, D.E., Mizukami, H., 1981. An exceptional amino acid replacement on the distal side of the iron atom in proboscidean myoglobin. Journal of Molecular Evolution 17, 140-147.
Abstract: Amino acid sequence determination of elephant myoglobin revealed the presence of the unusual substitution E7 His -- Gln.  Stereochemical analyses suggest that the most suitable residue which can functional substitute for His at this position in vertebrate globins in Gln.  Physiological studies imply that the slower rate of autooxidation of elephant is the result of this substitution which may confer some selective advantage on the species.  Comparative sequence data of paenungulate myoglobins suggest that the His -- Gln mutation probably occurred in an ancestor of Elephantinae.

Barnhart, M.I., Barmatoski, S.P., Goodman, M., Romero-Herrera, A.E., Lande, M.A., Birk, D.E., Shoshani, J., Prychodko, W., Lerman, E.J., Mikhelson, V.M., 1980. Tissue vestiges of an ancient Magadan mammoth calf. Scan Electron Microsc (Pt 2), 163-170.

Dene, H., Goodman, M., Romero-Herrera, A.E., 1980. The amino acid sequence of elephant (Elephas maximus) myoglobin and the phylogeny of the Proboscidae. Proc. Roy. Soc. Lond. [B] Biol. Sci. 207, 111-127.
Abstract: The complete amino acid sequence of skeletal myoglobin from the Asian elephant (Elephas maximus) is reported.  The functional significance of variations seen when this sequence is compared with that of sperm whale myoglobin is explored in the light of the crystallographic model available for the latter molecule.  The phylogenetic implications of the elephant myoglobin amino acid sequence are evaluated by using the maximum parsimony technique.  A similar analysis is also presented which incorporates all of the proteins sequenced from the elephant.  These results are discussed with respect to current views on proboscidean phylogeny.

Johnson, D.L., 1980. Problems in the Land Vertebrate Zoogeography of Certain Islands and the Swimming Powers of Elephants. Journal of Biogeography 7, 383-398.
Abstract: The presence of fossil elephants on certain islands off California, in the Mediterranean, in Indonesia, and off China has led to two widely accepted assumptions: (1) that elephants, being poor swimmers, could not have swum to the islands and therefore must have walked to them, which indicates that (2) land bridges once joined the islands to the mainland. These two assumptions have profoundly influenced various insular biogeographic and geologic reconstructions on and around these islands. New evidence, however, shows unequivocally that living elephants are excellent distance swimmers. They swim in a lunging, porpoise-like fashion while using their trunk as a snorkel. Elephant swimming speeds have been measured up to 2.70 km/h, and maximum distances estimated at 48 km. Their chief motives for swimming seem to be that they can see the islands and smell food on them. Because elephants are excellent distance swimmers, we must re-appraise the origin of land vertebrates on all the islands that held elephants, as well as reappraise the geologic reconstructions that assumed land bridges once connected these islands to the mainland.

Prager, E.M., Wilson, A.C., Lowenstein, J.M., Sarich, V.M., 1980. Mammoth albumin. Science 209, 287-289.
Abstract: Serum albumin was detected immunologically in muscle from a mammoth that died about 40,000 years ago. Rabbits injected with ground mammoth muscle produced antibodies that react strongly with elephant albumin, weakly with sea cow albumin, and still more weakly or not at all with other mammalian albumins. Since elephant albumin elicited antibodies with the same specificity, some of the surviving mammoth albumin molecules evidently have antigenic sites identical to those on native elephant albumin. Much of the mammoth albumin has, however, undergone postmortem change. The small amount of soluble albumin extractable from mammoth muscle is heterogeneous in size, charge, and antigenic properties.

Gillespie, J.M., 1979. Mammoth hair: stability of alpha-keratin structure and constituent proteins. Science 170, 1100-1102.

Kingdon, J., 1979. East African mammals. An atlas of evolution in Africa III. Large mammals. Academic Press, London.

Kirkwood, T.B., Holliday, R., 1979. The evolution of aging and longevity. Proc R Soc Lond B Biol Sci 205, 531-546.
Abstract: Aging is not adaptive since it reduces reproductive potential, and the argument that it evolved to provide offspring with living space is hard to sustain for most species. An alternative theory is based on the recognition that the force of natural selection declines with age, since in most environments individuals die from predation, disease or starvation. Aging could therefore be the combined result of late-expressed deleterious genes which are beyond the reach of effective negative selection. However, this argument is circular, since the concept of 'late expression' itself implies the prior existence of adult age-related physiological processes. Organisms that do not age are essentially in a steady state in which chronologically young and old individuals are physiologically the same. In this situation the synthesis of macromolecules must be sufficiently accurate to prevent error feedback and the development of lethal 'error catastrophes'. This involves the expenditure of energy, which is required for both kinetic proof-reading and other accuracy promoting devices. It may be selectively advantageous for higher organisms to adopt an energy saving strategy of reduced accuracy in somatic cells to accelerate development and reproduction, but the consequence will be eventual deterioration and death. This 'disposable soma' theory of the evolution of aging also proposes that a high level of accuracy is maintained in immortal germ line cells, or alternatively, that any defective germ cells are eliminated. The evolution of an increase in longevity in mammals may be due to a concomitant reduction in the rates of growth and reproduction and an increase in the accuracy of synthesis of macromolecules. The theory can be tested by measuring accuracy in germ line and somatic cells and also by comparing somatic cells from mammals with different longevities.

Kreps, E.M., Chirkovskaia, E.V., Pomazanskaia, L.F., Avrova, N.F., Levitina, M.V., 1979. Brain lipids of a mammoth, Elephas primigenius, which died more than 40,000 years ago. Zh Evol Biokhim Fiziol 15, 227-238.
Abstract: Studies have been made on the brain lipids of the 6--7-month mammoth which remained in the eternal ice for more than 40.000 years. Thin layer chromatography of chloroformmethanol extract of the brain lipids shows that all glycerophospholipids in the brain were destroyed. On the contrary, sphingophospholipid sphingomyelin yielded the evident spot which was identified by specific reactions and by comparison with sphingomyelin from the brain of rat. Sphingomyelin content was evaluated. Using gas-liquid chromatography, fatty acid composition of sphingomyelin was investigated. It was found to be close to that in contemporary mammals. Other shingolipids -- cerebrosides, sulfatides, gangliosides -- persisted (probably, only partially) and were studied quantitatively. Relative content of cerebrosides with normal fatty acids and hydrooxyacids was determined. Studies were also made on fatty acid composition of cerebrosides, sulfatides and gangliosides, as well as on the composition of spingosine bases of gangliosides. Free cholesterol was found in the brain of the mammoth. Other sterols were not detected. With respect to quantitative evaluation of the preserved lipids, it should be mentioned that on the one hand, the brain underwent dehydration which increased lipid content per a unit of "wet" weight, whereas on the other one lipids were partially degraded, this process decreasing their content.

Subbotin, V.M., Tum, Iu.V., 1978. Results of a microscopic study of several remains of the internal organs of Shandrin's mammoth. Arkh Anat Gistol Embriol 74, 85-89.
Abstract: The survived portion of an organ which topographically can be considered as a part of the lung, as well as a section of muscular tissue, perhaps that of the heart, has been studied. Under weak systems, the portion resembles the lung, spaces resembling alveoles, alveolar ducts and alveolar saccules are seen. "Alveolar" walls are stained dark grey by different methods; "alveoles" contain yellow granules, ferric components are not detected in them. The intestinal wall and that of the large vessel in the mesenteric cavity preserves a fibrillar carcass, fibrillar ultrastructure corresponding to collagen.

De Jong, W.W., Gleaves, J.T., Boulter, D., 1977. Evolutionary changes of alpha-crystallin and the phylogeny of mammalian orders. J Mol Evol 10, 123-135.
Abstract: The sequences of the A chains of the eye lens protein alpha-crystallin from seventeen mammalian species were compared. They showed a generally slow rate of evolution, but with marked variations in different lineages. Most substitutions have occurred in the C-terminal part of the chain, which probably forms part of the surface of the alpha-crystallin aggregate. The ancestral sequence method of Dayhoff revealed interesting indications about the phylogenetic relationships between the eleven mammalian orders that were represented by the investigated species. Most evident was the divergence of marsupial and placental orders. A notable resemblance between the hyrax and elephant sequences was observed, setting them apart from the ungulates, including whale. Primates, rodents, lagomorphs, insectivores and tupaiids seem to derive from a common stem group. These phylogenetic inferences are discussed in relation to current paleontological and taxonomical opinions, and compared to evidence from other protein sequence data.

De Jong, W.W., Nuy-Terwindt, E.C., Versteeg, M., 1977. Primary structures of alpha crystallin A chains of elephant, whale, hyrax and rhinoceros. Biochim. Biophys. Acta 491, 573-580.
Abstract: As part of a study of the evolutionary development of the eye lens protein alpha-crystallin the 173 residue A chain of this protein has been studied in elephant, whale, hyrax and rhinoceros.  The primary sturctures were inferred mainly from amino acid compositions of peptides obtained by enzymatic digestions and CNBr cleavage.  The positions of substitutions, as compared to known bovine A chain, were confirmed by Edman degradation.  In accordance with the previously observed slow rate of evolution of the A chain only a small number of substitutions were found among these species.  Elephant and hyrax share a number of unique substitutions, strongly indicating a common ancestry of these two species within the mammalian class.

Hanks, J., 1977. Comparative aspects of reproduction in the male hyrax and elephant. In: Calaby, J.H., Tyndale-Biscoe, C.H. (Eds.), Reproduction and evolution. Australian Academy of Science, pp. 155-164.

Douglas-Hamilton, I., Douglas-Hamilton, O., 1975. Among the elephants. Viking Press, New York.

Gombe, S., Heap, R.B., Sale, J.B., 1975. Endocrinology of pregnancy in the hyrax; plasma progesterone concentration and erythrocyte metabolism. Proc. Physiol. Soc. September, 13P-14P.

Heap, R.B., Gombe, S., Sale, J.B., 1975. Pregnancy in the hyrax and erthrocyte metabolism of progesterone. Nature 257, 809-811.

Moss, C., 1975. Portraits in the wild. University of Chicago Press, Chicago.

Riemann, G.P., Burridge, M.J., Behymer, D.E., Franti, C.E., 1975. Toxoplasma gondii antibodies in free-living African mammals. J Wildl Dis 11, 529-533.
Abstract: Twelve species of free-living African mammals from Kenya, Tanzania, Uganda and Zambia were tested for antibodies to Toxoplasma gondii using the indirect hemagglutination test. Of 157 animals sampled, 20 (13%) were seropositive. T. gondii antibodies were detected in Burchell's zebra, (Equus burchelli), hippopotamus (Hippopotamus amphibius), African elephant (Loxodonta africana), defassa waterbuck (Kobus defassa), lion (Panthera leo), and rock hyrax (Procavia capensis), The highest titers were found in elephants, two having titers of 1:4096 and one of 1:8192. These results are discussed in relation to the maintenance of T. gondii among African wildlife.

Scullard, H.H., 1974. The Elephant in the Greek and Roman World. Cornell University Press, Ithaca, NY.

Fawcett, D.W., Neaves, W.B., Flores, M.N., 1973. Comparative observations on intertubular lymphatics and the organization of the interstitial tissue of the mammalian testis. Biology of Reproduction 9, 500-532.

Maglio, V.J., 1972. Evolution of mastication in the Elephantidae. Evolution 26, 638-658.

Sikes, S.K., 1971. The Natural History of the African Elephant. American Elsevier Publishing Company, Inc., New York.

Aguirre, E., 1969. Evolutionary history of the elephants. Science, New Series 164, 1366-1376.
Abstract: Elephants which are among the most popular and decorative of animals, stand as a witness of prehistory, having been a part of the environment of our ancestors. The dinosaur was not contemporary with early man, as many films and stories insist, but the mammoth was. Although prehistoric or extinct elephants are frequently referred to as mammoths, such a designation is not always correct. The true mammoth is but one of many species of extinct elephants; furthermore, it belongs to one of a few genera, which include four or five species that have affinities with the woolly elephants. These different genera and species are grouped by zoologists into a family, Elephanttidae. Because this family originated by the beginning of the Pleistocne period, elephants can be considered contemporary with man. Anthropologists and prehistorians have often attempted to establish a chronology of sites of fossil man through correlations based upon the species of elephants associated with them but the systematics of the Elephantidae is quite confused. The documented monograph of Osborn established 10 genera and some 59 species of elephants; to these Garutt added two more genera. However many taxonomists have recognized only one genus and no more than five or six valid species. In the museum collection from most major sites, there are many samples with dubious identifications and many intermediate forms labeled either with two names or with a composite or new name. It has been assumed that many different species have lived contemporaneously in a single area, as was the case for the sample excavated in the railway trench of San Paolo, Italy, in the first years of this century. Explanations of the phylogeny of elephants have had one feature in common: the patterns for the phyletic trees have agreed with with the fashionable evolutionary theories of the particular period. Thus all the trees are dichotomic and linear form 1881 to 1888, fairly dichotomic form 1888 to 1912 and polyphyletic until 1923. After 1940 dichotomic patterns are again found. A review of the evolutionary history of the Proboscidae before the appearance of the elephants may help us to understand the significance of the evolving character in the latter. For Proboscidae since the Old Tertiary period, two major characteristics have been defined: the anterior teeth are missing except for one or two pairs of tusks; and there is an increasing number of rows of cusps, with every new transversal row appearing behind the other and elongating the molar teeth.

Hooijer, D.A., 1967. Indo-Australian insular elephants. Genetica 38, 143-162.

Nirmalan, G., Nair, S.G., Simon, K.J., 1967. Hematology of the Indian elephant (Elephas maximus). Canadian Journal of Physiology and Pharmacology 45, 985-991.
Abstract: Blood samples from 11 baby elephants, 14 tuskers, and 16 females comprising 5 pregnant and 11 nonpregnant nonlactating elephants (Elephas maximus) were analyzed to assess the hematological values and the influence of age, sex and pregnancy on these. The results obtained show that, compared with other mammals, the erythrocyte count in the blood of elephants is low and the mean corpuscular volume (m.c.v.), mean corpuscular hemoglobin (m.c.h.), and erythrocyte sedimentation rate (e.s.r.) are high. The very high values for m.c.v. and m.c.h. and the low erythrocyte count seen in elephants suggest that in the evolution of blood through they are more advanced than reptiles and birds in having eliminated the nuclei from the erythrocytes, they are still in the primitive state compared with the other mammals inasmuch as their erythrocytes have not attained the efficiency in the transportation of blood gasses that results from a reduction in size to facilitate numerical increase.  A low erythrocyte count and packed cell volume, a high e.s.r., an increase in neutrophils, and a decrease in the percentage of lymphocytes and eosinophils characterize pregnancy.  In baby elephants there is an increase in the total leucocyte and lymphocyte counts and a decrease in eosinophils.

Buettner-Janusch, J., Buettner-Janusch, V., Sale, J.B., 1964. Plasma proteins and haemoglobins of the African elephant and the hyrax. Nature 201, 510-511.

Carrington, R., 1959. Elephants:  A short account of their natural history, evolution and influence on mankind. Basic Books, Inc., New York.

Carrington, R., 1957. Mermaids and mastodons. Chalto & Windus, London.

Dolch, E.W., Dolch, M.P., 1956. Elephant Stories. Garrard Publishing Company, Champaign, Illinois.
Abstract: Note: This book is part of a Basic Vocabulary Series

Deraniyagala, P.E.P., 1955. Some Extinct Elephants, Their Relatives and the Two Living Species. Ceylon National Museums publication, Government Press, National Museum, Colombo (Sri Lanka).

Deranyagala, P.E.P., 1955. Some extinct elephants, their relatives and the two living species. Ceylon National Museum Publ., Colombo.

Weitz, B., 1953. Serological relationships of hyrax and elephant. Nature 171, 261.

Morrison-Scott, T.C.S., 1947. A revision of our knowledge of African elephants' teeth, with notes on Forest and "Pygmy" elephants. Procedings of the Zoological Society of London 117, 505-527.

Wislocki, G.B., 1940. The topography of the hypophysis of the elephant, manatee, and hyrax. Anatomical Record 77, 427-445.

Nilakantha of Rajamangalam, 1931. The elephant lore of the Hindus.  The elephant-sport (Matanga-lila) of Nilakantha.  Translated from original Sanskrit with introduction, notes and glossary by Franklin Edgerton. Yale University Press, New Haven.

Goodwin, G.G., 1925. The first living elephant in America. Journal of Mammalogy 6, 256-263.

Bishop, C.W., 1921. The elephant and its ivory in ancient China. Journal of the American Oriental Society 41, 290-306.

 1839. The Elephant (as he exists in a wild state and as he has been made subservient, in peace and war, to the purposes of man). Harper and Brothers, New York.
Abstract: Note: This work was originally published by the British Society for the Diffusion of Useful Knowledge

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