Structure of ivory

Profiles with all orientations have been used to visualize the 3D structure of ivory from tusks of elephant, mammoth, walrus, hippopotamus, pig (bush, boar, and warthog), sperm whale, killer whale, and narwhal. Polished, forming, fractured, aged, and stained surfaces were prepared for microscopy using epi-illumination. Tusks have a minor peripheral component, the cementum, a soft derivative of the enamel layer, and a main core of dentine=ivory. The dentine is composed of a matrix of particles 5-20 microm in diameter in a ground substance containing dentinal tubules about 5 microm in diameter with a center to center spacing of 10-20 microm. Dentinal tubules may be straight (most) or curly (pigs). The main findings relate to the way that dentinal tubules align in sheets to form microlaminae in the length of the tusk. Microlaminae are sheets of laterally aligned dentinal tubules. They are axial but may be radial (most), angled to the forming face (pigs and hippopotamus canines), or radial but helical (narwhals). Within the microlaminae the dentinal tubules may be radial, angled to the axis (whales, walrus, and pigs), or may change their orientation from one microlamina to the next in helicoids (canines of hippopotamuses, incisors of proboscidea). In the nonbanded, featureless ivories from the hippopotamus incisors, the dentinal tubules form radial microlamina from which the arrangements in other ivories can be derived. In the canines of hippopotamuses and incisors of proboscidea, the dentinal tubule orientation changes incrementally from one microlamina to the next in a helicoid, a stack of dentinal tubules that change their orientation by 180 degrees anticlockwise. Dentinal tubules having different orientations are laid down concurrently, not layer by layer as in most examples of helicoidal architecture (e.g., insect cuticle). In proboscidean ivory, the microlaminae are radial, normal to the banding of growth layers marking the plane of deposition. They form radial segments with each 180 degrees turn in the orientation of their constituent dentinal tubules. Below the cementum they are almost complete 180 degrees helicoids, but nearer to the core they become narrower with the loss of radially oriented dentinal tubules. These truncated helicoidal patterns appear in longitudinal profile as VVVV feather patterns rather than intersection intersection intersection intersection, each V or intersection being the side view of a partial or complete helicoid. The Schreger pattern in proboscidean ivory consists of these helicoids divided tangentially into columns in the length of the tusk. Narwhals have the most abundant matrix particles with their radial/helical dentinal tubules having a twist opposite to that in the cementum.     

Late Pleistocene proboscidean ivory artifacts from the Hiscock site,NY

Here is described the corpus of proboscidean ivory artifacts – the fruit of 30 years excavating – within the Palaeo-American component at the Hiscock site, western New York State. Several types and varieties are newly recognized and unique for the North American archaeological record. They allow us to forge linkages between the Old and New Worlds.     

Stable isotope series from elephant ivory reveal lifetime histories of a true dietary generalist

Longitudinal studies have revealed how variation in resource use within consumer populations can impact their dynamics and functional significance in communities. Here, we investigate multi-decadal diet variations within individuals of a keystone megaherbivore species, the African elephant (Loxodonta africana), using serial stable isotope analysis of tusks from the Kruger National Park, South Africa. These records, representing the longest continuous diet histories documented for any extant species, reveal extensive seasonal and annual variations in isotopic--and hence dietary--niches of individuals, but little variation between them. Lack of niche distinction across individuals contrasts several recent studies, which found relatively high levels of individual niche specialization in various taxa. Our result is consistent with theory that individual mammal herbivores are nutritionally constrained to maintain broad diet niches. Individual diet specialization would also be a costly strategy for large-bodied taxa foraging over wide areas in spatio-temporally heterogeneous environments. High levels of within-individual diet variability occurred within and across seasons, and persisted despite an overall increase in inferred C(4) grass consumption through the twentieth century. We suggest that switching between C(3) browsing and C(4) grazing over extended time scales facilitates elephant survival through environmental change, and could even allow recovery of overused resources.     

The presence of open dentinal tubules affects the biological properties of dental pulp cells <Emphasis Type="Italic">Ex Vivo</Emphasis>

To investigate the effects of open dentinal tubules on the morphological and functional characteristics of dental pulp cells. Morphological changes in human dental pulp cells that were seeded onto dentin discs with open dentinal tubules were investigated on days 1, 2, 4, and 10 of culture using scanning electron microscopy and fluorescence microscopy. Samples collected on days 1, 3, 6, 8, and 10 of culture were evaluated for cell proliferation rate and alkaline phosphatase activity. Cultured human dental pulp cells developed a columnar or polygonal morphology and monopolar cytoplasmic processes that extended into the dentinal tubules. The cells formed a multilayer and secreted an extracellular matrix onto the cell surface. Scanning electron microscopy and fluorescence microscopy revealed polarized organization of odontoblasts. Cells seeded onto dentin discs proliferated minimally but showed high levels of ALP activity. Dental pulp cells seeded onto treated dentin discs develop an odontoblastlike phenotype, which may be a potential alternative for use in experimental research on dentinogenesis.     

Missing teeth: Discordances in the trade of hippo ivory between Africa and Hong Kong

As the global epicentre of wildlife trade, Hong Kong plays an important role in the preservation or demise of biodiversity, including species found continents away. If mismanaged, legal trade in threatened species can lead to unsustainable exploitation. Inaccurate and incomplete trade records from the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES) undermine the regulation of this trade. We examine the trade of hippo (Hippopotamus amphibius) teeth to illustrate the extent of mismatched data between key trading nations. More than 90% of global hippo teeth trade is imported to, and re‐exported from, Hong Kong. Of that imported, over 75% originated in Tanzania or Uganda, but there are notable disparities in declared trade volumes. In most transactions, Hong Kong declares more volume imported than the volume declared exported by Uganda. Overall, Hong Kong has reported the import of 3,176 kg more hippo teeth than declared exported by Tanzania. This indicates that actual trade levels may exceed internationally agreed quotas. In total, over 14,000 kg of hippo teeth is unaccounted for between Uganda and Hong Kong, representing more than 2,700 individual hippos—2% of the global population. This gross discordance in trade data undermines regulatory measures and challenges the persistence of hippo populations in Africa.     

Gross,histological and ultrastructural morphology of the aglomerular kidney in the lined seahorse Hippocampus erectus

Histologic evaluation of the renal system in the lined seahorse Hippocampus erectus reveals a cranial kidney with low to moderate cellularity, composed of a central dorsal aorta, endothelial lined capillary sinusoids, haematopoietic tissue, fine fibrovascular stroma, ganglia and no nephrons. In comparison, the caudal kidney is moderately to highly cellular with numerous highly convoluted epithelial lined tubules separated by interlacing haematopoietic tissue, no glomeruli, fine fibrovascular stroma, numerous capillary sinusoids, corpuscles of Stannius and clusters of endocrine cells adjacent to large calibre vessels. Ultrastructural evaluation of the renal tubules reveals minimal variability of the tubule epithelium throughout the length of the nephron and the majority of tubules are characterized by epithelial cells with few apical microvilli, elaborate basal membrane infolding, rare electron dense granules and abundant supporting collagenous matrix.     

Long-bone geometry in columnar-limbed animals: allometry of the proboscidean appendicular skeleton

Limb-bone allometry was investigated for 19 species of proboscideans, spanning almost the entire phylogenetic spectrum. More archaic proboscideans ('gompthotheres') have substantially thicker long-bone diaphyses relative to length than elephantids, as has been suggested previously, but contrary to previous suggestions it could not be confirmed that Mammuthus had more massive long-bone diaphyses on average than extant Elephas and Loxodonta . When correcting for phylogeny, the circumference of the limb bones to their length in proboscideans as a group generally scale with negative allometry, becoming stouter with increased length, as would be expected from limb mechanics. Few slopes were, however, statistically significantly negatively allometric. Correcting for phylogeny produced better correlations than traditional regression analyses, in contrast to most other studies where the reverse is the case. Intraspecific analyses of extant Elephas and Loxodonta , in addition to Mammuthus primigenius , Mammut americanum, and Gomphotherium productum , also resulted in negatively allometric regression slopes, frequently conforming to the theory of elastic similarity, as could be expected from the columnar posture of proboscideans. At present the reasons for the more massive limbs of gomphotheres s.l. are not fully understood. © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 149 , 423–436.     

Employing participatory surveys to monitor the illegal killing of elephants across diverse land uses in Laikipia–Samburu,Kenya

Levels and trends of illegal killing of elephants are measured by the Convention on International Trade in Endangered Species (CITES) Monitoring the Illegal Killing of Elephants (MIKE) programme in sites across Africa and Asia. In the mostly unprotected Laikipia–Samburu MIKE site in northern Kenya, elephant mortality data were collected using both standard law enforcement monitoring procedures, relying on patrolling, and participatory methods involving local communities. Qualitatively, traditional patrolling techniques were more successful in protected areas whereas participatory approaches provided more information outside protected areas, where elephant were most at risk from ivory poachers. A minimum of 35% of the 389 verified carcasses during 2001–2003 were illegally killed. In this baseline study, land uses ranked from highest to lowest by the proportion of illegally killed elephants (PIKE) were community conservation areas, government trust lands, forest reserves, private ranches, settlement areas and national reserves. PIKE trends derived from traditional and participatory data sources were similar across years and indicate elephants were at greater risk in regions outside government or privately patrolled areas. We suggest that PIKE is a useful index for comparing levels and trends in illegal killing of elephants, and that carcass ratios and presence/absence of tusks are useful proxy indicators of mortality and its causes.     

Origins of bone repair in the armour of fossil fish: response to a deep wound by cells depositing dentine instead of dermal bone

The outer armour of fossil jawless fishes (Heterostraci) is, predominantly, a bone with a superficial ornament of dentine tubercles surrounded by pores leading to flask-shaped crypts (ampullae). However, despite the extensive bone present in these early dermal skeletons, damage was repaired almost exclusively with dentine. Consolidation of bone, by dentine invading and filling the vascular spaces, was previously recognized in Psammolepis and other heterostracans but was associated with ageing and dermal shield wear (reparative). Here, we describe wound repair by deposition of dentine directly onto a bony scaffold of fragmented bone. An extensive wound response occurred from massive deposition of dentine (reactionary), traced from tubercle pulp cavities and surrounding ampullae. These structures may provide the cells to make reparative and reactionary dentine, as in mammalian teeth today in response to stimuli (functional wear or damage). We suggest in Psammolepis, repair involved mobilization of these cells in response to a local stimulatory mechanism, for example, predator damage. By comparison, almost no new bone is detected in repair of the Psammolepis shield. Dentine infilling bone vascular tissue spaces of both abraded dentine and wounded bone suggests that recruitment of this process has been evolutionarily conserved over 380 Myr and precedes osteogenic skeletal repair.     

The anatomy of Odobenocetops (Delphinoidea, Mammalia), the walrus-like dolphin from the Pliocene of Peru and its palaeobiological implications

The Pliocene Beds of the Pisco Formation on the southern coast of Peru have yielded three new skulls of the walrus-like odontocete, Odobenocetops (Delphinoidea, Cetacea). Two of the skulls are from a slightly younger horizon than the holotype of O. peruvianus and belong to a different species, O. leptodon . The holotype of O. leptodon bears a 135 cm long needle-like right tusk and a small 25 cm long left tusk, the apex of which was erupted. The third skull, referred to a female of O. peruvianus , bears two small tusks similar in size to the small left tusk of the holotype. The new specimens include periotic, tympanic and ear ossicles, which confirm referral to the Order Cetacea and the morphology of the humerus reinforces affinities of odobenocetopsids to monodontids. Because the anterodorsal edge of its orbit is slightly concave O. leptodon had reduced anterodorsal binocular vision, a condition compensated for by the probable presence of a small melon (and inferred echolocation). The head was bent ventrally when swimming in such a way that the long tusk was approximately parallel to the axis of the body. The extremely salient occipital condyles of Odobenocetops are indicative of great mobility of the neck, probably related to bottom-feeding. © 2002 The Linnean Society of London, Zoological Journal of the Linnean Society , 2002, 134 , 423–452.     

The histology and affinities of sinacanthid fishes: primitive gnathostomes from the Silurian of China

On the basis of well preserved specimens from the Lower Silurian of the Tarim Basin, Xinjiang Uygur Autonomous Region and Shiqian County, Guizhou Province, People's Republic of China we describe in detail the histological structure of sinacanthid spines, the only known remains of a group of fishes common in Silurian strata from China. The sinacanthids have previously been assigned either to the acanthodians or to the chondrichthyans. The spine structure is composed of an outer layer of atubular dentine and an inner layer of globular calcified cartilage, and the nature and distribution of these tissues indicates that the spines were formed as a result of interaction between the endoskeleton and dermoskeleton. The tissue distribution and style of growth described herein places the sinacanthids crownwards of the placoderms, and possibly within the total group Chondrichthyes. However, before they can be firmly placed within a phylogenetic scheme, further evidence is required both on the general anatomy of sinacanthids and on the nature of chondrichthyan apomorphies.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 144 , 379–386.     

Relationships among development, ecology, and morphology in the evolution of Echinoderm larvae and life cycles

The evolution of life cycles involves transitions between discrete states in one or more of the characters that comprise a developmental pattern. In this paper, we examine three of the major life cycle characters and the states for these characters. Using examples from echinoderms, we discuss the evolutionary transitions that have occurred in the type of morphogenesis, developmental habitat, and mode of nutrition during development. We evaluate the functional requirements associated with these transitions to infer the likelihood (frequency or rapidity) of change in a given character and of biases in the polarity of character state transitions. Using comparisons of closely related species, we evaluate the change between states in one character for dependence on the state of, or correlated changes in, other characters. Based on our analysis of congeneric species that differ in developmental habitat, we conclude that the transition between pelagic and benthic development is an ecological change that is independent of changes in morphogenesis and should be reversible. In contrast, the transition from feeding to nonfeeding development has been considered to be irreversible because it involves marked changes in larval morphology. We re-examine the transition between different modes of larval nutrition in light of recent studies that show that there exists a continuum of nutritional strategies between planktotrophy and lecithotrophy. This continuum is largely determined by variation in maternal investment and does not involve alterations in larval morphology. We suggest that the boundary between planktotrophy and lecithotrophy is frequently crossed and that this transition is reversible. Ecological changes represent the crossing of a functional threshold. Only after crossing the threshold, do larvae experience qualitatively different selective pressures that can lead to subsequent changes in morphology and development. Two different changes have occurred in the type of morphogenesis: the simplification of larval morphology that is associated with obligate (nonfeeding) lecithotrophy and the loss of the larval body plan in the evolution from indirect to direct development. It is the modification of morphology independent of the ecological changes that requires alterations in developmental processes, constrains evolutionary options, imposes irreversibility, and establishes the discrete nature of larval patterns in marine invertebrates.     

An exceptional amino acid replacement on the distal side of the iron atom in proboscidean myoglobin

Summary 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 functionally substitute for His at this position in vertebrate globins is Gln. Physiochemical studies imply that the slower rate of autooxidation of elephant myoglobin 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.     

Contribution of isotopic biogeochemistry (13C, 15N, 18O) to the paleoecology of mammoths (mammuthus primigenius)

Carbon isotopic abundances in Alaskan, Albertan and Russian mammoths indicate a diet of C₃‐plants. The relatively high nitrogen isotopic abundances suggest arid conditions in Alberta and Alaska during the last ice age, and similar conditions in Russia. Nitrogen isotopic abundances are higher in mammoths relative to coeval herbivores, which may be due to differences in protein content of their diet. Oxygen isotopic abundances are similar in mammoths and in modern mammals from the same areas. The characteristic isotopic signature of mammoth ivory allows distinction from elephant or marine mammal ivory.     

扬子鳄胚胎中脑视叶的组织发生

观察了16例不同时间扬子鳄胚胎中脑视叶的组织发生过程。胚胎孵育第6d,三个脑泡明显;孵育第9～10d,中脑泡可分细胞层和边缘层或纤维层,中脑水管未形成;孵育第18d,视叶隆起于中脑背侧,中脑水管形成,视叶分三层;孵育第24d,视叶分5层;孵育第34d,视叶分化为6层;孵育第51d,视叶分化为8层,与初生扬子鳄中脑视叶分层相同。     

Limb posture and gait in Numidotherium koholense, a primitive proboscidean from the Eocene of Algeria

The tapir-sized late Early Eocene proboscidean, Numidotherium koholense , is the oldest known member of the mammalian order to which the living elephants belong. Morphology of the limb skeleton is reviewed and newly discovered elements are described, with emphasis placed on features considered to have a bearing on limb posture and locomotion. Structure of the shoulder joint and the fact that the antebrachium is fused in a supinated position, indicates that forelimb posture would have been of the abducted or semi-sprawling type. The carpus, while exhibiting the typically proboscidean serial arrangement of carpals (no contact between unciform and lunar), also possesses a free os centrale. The presence of an os centrale and entepicondylar foramen in the humerus are unexpected eutherian plesiomorphies. The structure of the hip joint indicates an abducted hindlimb posture. In both the fore- and hindlimb, mechanisms exist at the major joints to cope with the complicated flexion-extension paths typical of a semi-sprawling gait. Both the carpus and tarsus are indicative of a plantigrade stance. This, in concert with an abducted limb posture, indicates that the gait in numidotheres contrasts markedly with the sub-unguligrade, parasagittal gait of elephantiform proboscideans. The recent suggestion that Proboscidea have their origins in a cursorially adapted phenacodontid-like stock receives no support from this analysis.