Scute’s polymorphism as a source of evolutionary development of the turtle shell

The scute mosaic (pholidosis) of the turtle shell is a complex correlated system of the modular type. Horny scutes are separate morphological elements partially closely connected with each other and partially relatively autonomous in development. The last feature causes high variability of scutes in the shape, size, rate and direction of growth, and provides the basis of transformation of the entire mosaic. In the evolution of turtles, the horny shell changed towards a decrease in the number of elements composing it. The process of oligomerization developed through reduction and fusion of scutes or their anlages. The traces of these transformations are observed in the ontogeny of living turtles. The scutes undergoing reduction display the following developmental deviations: (1) a decrease in size of the scute anlage, (2) the temporal shift in initiation to later embryonic stages, (3) absence of an anlage of a own furrow (the boundaries of the scute are formed by the furrows of neighboring scutes), and (4) a decrease in size of the zone and rate of the scute growth. The fusion of horny scutes follows two patterns: (1) fusion of scute anlages and (2) reduction of horny furrows separating scutes before. Secondary polymerization of the scute mosaic by the appearance of additional elements usually results from abnormal development and is infrequently fixed in evolution. The main mechanism of evolutionary changes in turtle pholidosis was heterochrony, i.e., the time shift in initiation and developmental rate of scutes. The heterotopies, i.e., changes in the position of scute anlages, played a minor role in the evolution of turtles; they usually caused only scute abnormalities, which was frequently asymmetrical.     

Unusual aetosaur armor from the Upper Triassic of West Texas,U.S.A.

We report two unusual aetosaur scutes from the Tecovas Member of the Dockum Formation, Chinle Group (upper Carnian) of Crosby County, Texas, U.S.A. Originally collected by University of Michigan expeditions in the 1920’s, these scutes cannot be assigned with certainty to any known species of aetosaurs known from the American Southwest. One of these scutes may be a cervical horn ofParatypothorax, and if so confirms the similarities of this aetosaur toDesmatosuchus in the possession of horned lateral spikes. The other is a paramedian scute of a new aetosaur taxon inadequately known at present to be assigned a formal name. These scutes indicate that aetosaurs were more diverse in the Chinle Group than currently known, but do not alter the robust Late Triassic biochronology based on aetosaurs.     

Proliferation in the epidermis of chelonians and growth of the horny scutes

The proliferation of the epidermis in soft skin, claws, and scutes of the carapace and plastron in the tortoise (Testudo hermanni) and the turtle (Chrysemys picta) were studied using autoradiographic and immunocytochemical methods. During the growing season, a basal keratinocyte in the epidermis of soft skin and claws takes 5-9 days to migrate into the corneous layer. In the tortoise, during fall/winter (resting season) a few alpha-keratin cells are produced in soft epidermis and hinge regions among scutes and occasional beta-keratin cells in the outer scute surface. When growth is resumed in spring (growing season), cell proliferation is intense, mainly around hinge regions and tips of marginal scutes. No scute shedding occurs and numerous beta-keratin cells are produced around the hinge regions, while alpha-keratin cells disappear. Beta-cells form a new thick corneous layer around the hinge regions, which constitute the growing rings of scutes. Beta-keratin cells produced in more central parts of scutes maintain a homogeneous thickness of the corneous layer along the whole scute surface. In the turtle, a more complicated process of scute growth occurs than in the tortoise. At the end of the growing season (late fall) the last keratinocytes formed beneath the old stratum corneum of the outer scale surface and hinge regions produce more alpha- than beta-keratin. These thin alpha-keratin cells form a scission layer below the old stratum corneum, which extends from the hinge regions toward the center of scutes and the tip of marginal scutes. In the resting season (fall/winter) most cells remain within the germinative layer of the carapace and plastron and a few alpha-cells move in 7-9 days into the corneous layer above hinge regions. In the following spring/summer (growing season) a new generation of beta-keratin cells is produced beneath the scission layer from the hinge region and more central part of the scutes. The epidermis of the inner surface of scutes and hinge regions contains most of the cells incorporating thymidine and histidine, while the remaining outer scute surface is less active. It takes 5-9 days for a newly produced beta-cell to migrate into the corneous layer. These cells form a new corneous layer that extends the whole scute surface underneath the maturing scission layer. The latter contains lipids and eventually flakes off, determining shedding of the above outer corneous layer in late spring or summer.     

Studies on the growth, structure and abundance of the Mediterranean spur-thighed tortoise, Tesudo graeca in field populations

The blackness and physical appearance of Mediterranean spur-thighed tortoises ( Testudo graeca L.) are recorded in field populations. The inter-relationships between carapace and plastron length, fourth vertebral scute width and other morphometric characters are related to growth. The rear fontanelle is larger in males and the plastron length correspondingly smaller. The relationship between carapace length ( y ) in mm and body mass ( x ) in g is strongly allometric and given by y = 16.63 x ^0–34. The maximum carapace lengths recorded for male and female T. graeca graeca L. in NW. Africa are 151 and 198 mm, and for larger T. graeca ibera Pallas in W. Turkey and NE. Greece 208 and 218 mm, respectively. The growth annuli of epidermal scutes are deposited yearly until external growth ceases. The number is used to assess the age of wild animals. Tortoises can be full-grown by 19 years but some, especially slower growing ones, deposit further annuli. The biggest mean growth increments occur up to 7 years with the largest in individuals over the 5–12 year period. The proportion of immature tortoises, adult sex ratios, size and body mass structure, age distribution, annual recruitment and survivorship until full-grown, and abundance (sighting frequency) are compared in northern and southern range populations of NW. African T. g. graeca and S. Anatolian and SE. European (Thrace and Macedonia) ones of T. g. ibera. These are discussed in relation to other tortoise species.     

Hemolivia mauritanica (Haemogregarinidae: Apicomplexa) infection in the tortoise Testudo graeca in the Near East with data on sporogonous development in the tick vector Hyalomna aegyptium

Testudo groeca tortoises were collected in the northern and southern Golan Heights (Israeli occupied territory of south Syria), and various locations in Israel and Palestine. Hyoalomma oegyptium ticks were found only on Golan Height tortoises, and only the tortoises and ticks from the northern Golan Heights were infected with Hemolivia mouritanica. Tortoises became infected after ingesting infected ticks. Male ticks carrying sporocysts, which remain attached to tortoises for extended durations, apparently served as the source for dissemination of new infections among tortoises. Sporogenesis followed the pattern observed in the two other known species of Hemolivia, though there was some evident variation in fine-structura detail. The sutural slit detected in the H. mouritanica mature sporocyst wall was reminiscent of the suture characteristic of Coccidia of heterothermic vertebrate hosts; it could be a common ancestral character for both hemogregarines     

Morphological and genetic variation in the endangered Sulawesi tortoise Indotestudo forstenii: evidence of distinct lineages?

Limited fieldwork and anecdotal evidence indicate that allopatric populations of the endangered Sulawesi tortoise (Indotestudo forstenii) differ in size and presence/absence of a nuchal scute, suggesting that these may constitute separate evolutionary lineages. We examined morphological and genetic variation to determine if the nuchal scute is correlated with size or genetic divergence. Our results indicate a strong correlation between size and nuchal scute such that turtles lacking a nuchal scute were larger than those possessing a nuchal scute. However, we found no correlation between genetic divergence and presence/absence of nuchal scutes, and thus no evidence of genetically differentiated lineages.     

Poxvirus in scaled quail and prevalences of poxvirus-like lesions in northern bobwhites and scaled quail from Texas

Prevalences of poxvirus-like lesions were determined for 177 northern bobwhites (Colinus virginianus) and 24 scaled quail (Callipepla squamata) trapped in southern Texas from 1976 to 1979 and for 190 northern bobwhites and 105 scaled quail shot at five locations in southern Texas from 1980 to 1981. None of the northern bobwhites trapped in 1976-1977 was infected, but 54% of the trapped scaled quail were infected; 17% of the northern bobwhites and 34% of the scaled quail shot in 1980-1981 had pox lesions, primarily on the wings. Prevalence was unrelated to sex or age of birds. For both species, prevalence was greatest during late spring and early summer. Histologic and electron microscopic examination confirmed poxvirus in two scaled quail, which constituted the first report of poxvirus in this species.     

Ranavirus infection of free-ranging and captive box turtles and tortoises in the United States

Iridoviruses of the genus Ranavirus are well known for causing mass mortality events of fish and amphibians with sporadic reports of infection in reptiles. This article describes five instances of Ranavirus infection in chelonians between 2003 and 2005 in Georgia, Florida, New York, and Pennsylvania, USA. Affected species included captive Burmese star tortoises (Geochelone platynota), a free-ranging gopher tortoise (Gopherus polyphemus), free-ranging eastern box turtles (Terrapene carolina carolina), and a Florida box turtle (Terrepene carolina bauri). Evidence for Ranavirus infection was also found in archived material from previously unexplained mass mortality events of eastern box turtles from Georgia in 1991 and from Texas in 1998. Consistent lesions in affected animals included necrotizing stomatitis and/or esophagitis, fibrinous and necrotizing splenitis, and multicentric fibrinoid vasculitis. Intracytoplasmic inclusion bodies were rarely observed in affected tissues. A portion of the major capsid protein (MCP) gene was sequenced from each case in 2003-2005 and found to be identical to each other and to Frog virus 3 (FV3) across 420 base pairs. Ranavirus infections were also documented in sympatric species of amphibians at two locations with infected chelonians. The fragment profiles of HindIII-digested whole genomic DNA of Ranavirus, isolated from a dead Burmese star tortoise and a southern leopard frog (Rana utricularia) found nearby, were similar. The box turtle isolate had a low molecular weight fragment that was not seen in the digestion profiles for the other isolates. These results suggest that certain amphibians and chelonians are infected with a similar virus and that different viruses exist among different chelonians. Amphibians may serve as a reservoir host for susceptible chelonians. This report also demonstrated that significant disease associated with Ranavirus infections are likely more widespread in chelonians than previously suspected.     

Structure and Development of the Odontodes in an Armoured Catfish,Corydoras aeneus (Siluriformes,Callichthyidae)

Abstract In some living osteichthyans (e.g. the armoured catfishes) the postcranial dermal skeleton exhibits tooth-like structures (odontodes) similar to those present in the dermal skeleton of the ancient craniates. We have undertaken this work to compare odontode with tooth development, structure, attachment to a bony support and replacement. We studied the odontodes fixed on the scutes (i.e. postcranial dermal plates) in a growth series of Corydoras aeneus using light, scanning and transmission electron microscopy. Odontodes are constituted of a pulp cavity surrounded by a cone of dentine itself capped with hypermineralized substance. The pulp cavity is devoid of nerves and blood vessels and there are no odontoblastic processes in the dentine. The dentine cone is firmly attached to a circular bony protuberance of the scute surface, the pedicel or attachment bone, by means of a ligament. An odontode anlage develops as a small invagination of a dermal papilla projecting into the epidermis, the basal cell layer of which constitutes a dental epithelium. First, dentine is deposited, next the hypermineralized substance, then the ligament and attachment bone. Odontodes develop in two positions with regard to the scute surface: a primary position when new odontodes form at the posterior border of the enlarging scute; a secondary position when new odontodes replace old odontodes that have been shed during thickening of the scute. In this case, the ligament and part of the base of the dentine cone are resorbed but not the pedicel of attachment bone, which is covered by deposition of scute matrix after the odontode has been shed. Within the scute matrix, the embedded pedicels of successive generations of odontodes are preserved, forming piles in the scutes of adult specimens.     

Polymorphism of the 12S rRNA gene and phylogeography of the Central Asian tortoises Agrionemys horsfieldii gray, 1844

Based on intraspecific polymorphism of 12S rRNA gene, genetic variation of isolated populations of the Central Asian tortoise, Agrionemys horsfieldii, was for the first time investigated on a large part of the species distribution range, encompassing Uzbekistan, southern Kazakhstan, and northern and eastern Iran. In 59 tortoises, four haplotypes were discovered, including two (AH1 and AH2), described earlier. Haplotype AH1 was detected in 52 tortoises, inhabiting southern Kazakhstan and Uzbekistan. Haplotype AH2 was found in four tortoises from the border territory between Uzbekistan, Tajikistan, and Afghanistan. Two novel haplotypes, AH3 and AH4, were detected in the three tortoises from Iran. Based on nucleotide substitutions in the 12S rDNA sequence, the possible divergence time between the tortoises from different parts of the range was estimated. Possible pathways of the formation of modern intraspecific groups of A. horsfieldii are discussed.     

Polymorphism of the 12S rRNA gene and phylogeography of the Central Asian tortoises <Emphasis Type="Italic">Agrionemys horsfieldii</Emphasis> gray, 1844

Based on intraspecific polymorphism of 12S rRNA gene, genetic variation of isolated populations of the Central Asian tortoise, Agrionemys horsfieldii, was for the first time investigated on a large part of the species distribution range, encompassing Uzbekistan, southern Kazakhstan, and northern and eastern Iran. In 59 tortoises, four haplotypes were discovered, including two (AH1 and AH2), described earlier. Haplotype AH1 was detected in 52 tortoises, inhabiting southern Kazakhstan and Uzbekistan. Haplotype AH2 was found in four tortoises from the border territory between Uzbekistan, Tajikistan, and Afghanistan. Two novel haplotypes, AH3 and AH4, were detected in the three tortoises from Iran. Based on nucleotide substitutions in the 12S rDNA sequence, the possible divergence time between the tortoises from different parts of the range was estimated. Possible pathways of the formation of modern intraspecific groups of A. horsfieldii are discussed.     

Effects of the egg incubation environment on turtle carapace development

Developing organisms are often exposed to fluctuating environments that destabilize tissue-scale processes and induce abnormal phenotypes. This might be common in species that lay eggs in the external environment and with little parental care, such as many reptiles. In turtles, morphological development has provided striking examples of abnormal phenotypic patterns, though the influence of the environment remains unclear. To this end, we compared fluctuating asymmetry, as a proxy for developmental instability, in turtle hatchlings incubated in controlled laboratory and unstable natural conditions. Wild and laboratory hatchlings featured similar proportions of supernumerary scales (scutes) on the dorsal shell (carapace). Such abnormal scutes likely elevated shape asymmetry, which was highest in natural nests. Moreover, we tested the hypothesis that hot and dry environments cause abnormal scute formation by subjecting eggs to a range of hydric and thermal laboratory incubation regimes. Shape asymmetry was similar in hatchlings incubated at five constant temperatures (26–30°C). A hot (30°C) and severely Dry substrate yielded smaller hatchlings but scutes were not overtly affected. Our study suggests that changing nest environments contribute to fluctuating asymmetry in egg-laying reptiles, while clarifying the conditions at which turtle shell development remains buffered from the external environment.     

Development and fine structure of the bony scutes in Corydoras arcuatus (Siluriformes,callichthyidae)

The development and the structure of the bony scutes have been studied in a growth series of the armored catfish Corydoras arcuatus using light and electron microscopy. Fibroblast-like cell condensations appear in the dermis, in the posterior region of the caudal peduncle, and these will constitute the scute papillae. Collagen bundles of the preexisting dermis colonized by the papilla cells are remodeled and incorporated in the papilla to form, in addition to newly synthesized woven-fibered bony material, the initium of the scute. This process of formation differs from that described for the dermal papilla of an elasmoid scale. During growth, the osteoblasts surrounding the scute constitute the scute sac in which the scute grows. Parallel-fibered bone is deposited on both sides of the initium, and osteoblasts are incorporated within the scute matrix. The remodeling and incorporation of collagen bundles of the preexisting dermis is maintained during growth only in the deep, anterior region of the scute. The posterior region and the upper surface of the scute are close to the epidermal-dermal boundary. When growth slows down in the upper part of the scute, a characteristic, well-mineralized tissue, composed of thin vertical fibrils and granules and devoid of typical striated collagen fibrils, is deposited on the scute surface. A new term, hyaloine, is introduced for this nonosseous, highly mineralized layer constituting the upper part of the scute. Hyaloine shows thin electron-dense lines, which probably correspond to periodic growth arrests. The structure and localization of the hyaloine are compared to other well-mineralized, similar tissues found on the surface of the dermal skeleton in lower vertebrates. © 1993 Wiley-Liss, Inc.     

Salmonella from gopher tortoises (Gopherus polyphemus) in south Georgia

From 2002 to 2006, gopher tortoises (Gopherus polyphemus) were collected at Moody Air Force Base, Lowndes/Lanier counties, Georgia, USA, and opportunistically surveyed for the presence of Salmonella species. Four of 155 (2.6%) cloacal swabs collected from 80 tortoises were positive for the presence of Salmonella enterica, and the following serovars were identified: Give, Hartford, Javiana, and Luciana. Female tortoises (5%) were infected at a rate similar to male tortoises (5%). All isolates were obtained from adult tortoises (n = 73); subadults (n = 7) were all negative. Each isolated serovar is a potential human pathogen, suggesting appropriate precautions should be emphasized when handling these animals.     

Developmental structure of the vertebral column,fins, scutes and scales in bester sturgeon,a hybrid of beluga Huso huso and sterlet Acipenser ruthenus

In the larval bester, a hybrid sturgeon of beluga Huso huso and sterlet Acipenser ruthenus, development of cartilage around the notochord began 7 days post hatch (dph) (14·0 mm, total length, L_T). The vertebral cartilage develops in the following sequence: basidorsals and basiventrals, neural canals, neural spines and ribs. The development of ribs remained incomplete in the largest specimen (181 dph, 179 mm L_T) that was examined. Endoskeletal development of the fins began 4 dph for the dorsal and anal fins, 6 dph for the pectoral fin and 10 dph for the caudal and pelvic fins. Complete elements of all fins were observed by 91 dph and complete ossification of fin rays was observed by 122 dph in the double‐stained specimens. Observation of the histological sections, however, suggested that ossification occurred soon after the formation of the organic matrix in the fin rays. Dorsal scutes were first visible by 25 dph, followed by the lateral and ventral scutes, which were visible by 37 and 44 dph, respectively. The number of scutes was fixed at 44, 59 and 91 dph and ossification was complete by 59 (dorsal) and 91 dph (lateral and ventral scutes) in the double‐stained specimens. Ossification occurred soon after the formation of the scute organic matrix in the histological sections. Four types of scales were observed in the H. huso×A. ruthenus hybrid. Median predorsal, preanal and small scales on the anterior section of the head were visible by 59 dph. Scales on the caudal fin were visible by 91 dph and a variable assemblage of scales anterior to the anal fin was visible by 122 dph. Both the scutes and scales developed in a process that is similar to that of intramembranous ossification.     

Carapace and plastron sensitivity to touch and vibration in the tortoise (Testudo hermanni and T. graeca)

Neural impulses in response to tactile stimulation of the shell were recorded from afferent nerve fibres in tortoises ( T. graeca and T. hermanni). It was found that there is a mechanoreceptive innervation in the superficial layers of the shell which is sensitive to transient stimuli, particularly to vibration at frequencies up to 100 Hz. Receptive fields pertaining to single and small groups of individual afferent fibres were mapped: the fields were sharply circumscribed and distributed in relation to the scutes of the shell. The tactile innervation that was found would be consistent with a capacity for recognition and accurate localization of innocuous stimuli and may play a central role in courtship and mating behaviour.     

记甘肃晚侏罗世一蜥蜴新属

本文记述石龙子类 Scincomorpha 蜥蜴的—新属,甘肃拟贝氏蜥 Mimobefklesisaurus gansuensis gen. et sp. nov. 化石发现于甘肃省肃北县马宗山区晚侏罗世的赤金堡群.这是石龙子类化石在我国的第一次发现,也是目前所发现的这类动物在亚洲的最早的代表.     

Comparing aging precision of calcified structures in shovelnose sturgeon

Age estimates for population analysis must be precise. We assessed the usefulness of pectoral fin rays, sphenoids, opercula, and dorsal scutes of shovelnose sturgeonScaphirhynchus platorynchus (n = 30) as aging structures based on ease of collection, distinctness of annuli, and measures of precision both between and within readers. We also determined how age estimates from paired fin rays of individuals were related (n = 106). Pectoral fin rays generated the highest within‐reader precision (100% within 2 years) followed by sphenoids (58%), opercula (56%), and dorsal scutes (49%). Ages estimated by the pectoral fin ray also had higher between‐reader agreement (80% within 1 year) than did those from the operculum (60%), sphenoid (59%), or dorsal scute (56%). Likewise, age estimates from pectoral fin rays had the lowest mean coefficient of variation (8.2%) followed by sphenoids (9.9%), opercula (11.3%), and dorsal scutes (11.5%). Only the operculum produced biased estimates between readers. Ages from paired fin rays agreed poorly (36% exact, 30% within 1 year) although no aging bias occurred. The pectoral fin ray is typically used to age shovelnose sturgeon. Because uncertainty about accuracy and precision of age estimates from this structure remains, shovelnose sturgeon management objectives that result from age data should remain conservative.