Frugivory and seed dispersal by chelonians: a review and synthesis

In recent years, it has become clear that frugivory and seed dispersal (FSD) by turtles and tortoises is much more common than previously thought. We here review published and unpublished records of chelonian FSD, and assess the role of chelonians as seed dispersers, from individual species to the community level. We first discuss the distribution of chelonian FSD and the characteristics of the fruit and/or seed species eaten and dispersed by chelonians. We then use the seed dispersal efficiency framework to explore the quantitative and qualitative components of seed dispersal by tortoises and turtles, embarking on a journey from when the fruits and/or seeds are consumed, to when and where they are deposited, and assess how efficient chelonians are as seed dispersers. We finally discuss chelonian FSD in the context of communities and of chelonians as megafauna. A substantial proportion of the world's aquatic and terrestrial turtles and a major part of testudinid tortoises (71 species in 12 families) include fruits and/or seeds in their diet; fruits of at least 588 plant species in 121 families are ingested and/or dispersed by chelonians. For some chelonians, overall or in certain seasons, fruit may even form the largest part of their diet. Contrary to seed dispersal by lizards, the other major reptilian frugivores, chelonian FSD is not an island phenomenon in terms of geographic distribution. Nevertheless, on islands tortoises are often among the largest native terrestrial vertebrates – or were until humans arrived. We synthesise our knowledge of chelonian FSD, and discuss the relevance of our findings for conservation and restoration, especially in relation to rewilding with large and giant tortoises.     

中国鲨类脑颅的研究

本文解剖观察了我国有代表性的鲨类脑颅共32种,分隶于8目14科24属。研究结果认为鲨类的脑颅共可分为9个式型和12个亚型。现存虎鲨目、须鲨目的鲸鲨科和扁鲨目吻软骨缺如,六鳃鲨目、须鲨目、角鲨目和锯鲨目均具一吻软骨,它们是一些古老和一些特化的类群。现存大多数种类均具3根吻软骨。在各不同分类阶元常有其不同的形态特征,可作为分类依据之一,亦可显示其亲缘关系。     

中国鳐类脑颅的研究

作者解剖观察了33种,隶于4目、7亚目、15科、19属的中国鳐类脑颅的形态。研究结果认为:锯鳐目和鳐目是原始类群,它们均具吻软骨,其中圆犂头鳐科和团扇鳐科是特化类群。电鳐目亦具吻软骨,它们是特化和退化类群。在较高等的鲼目则无吻软骨。依据鳐类不同的分类阶元,其脑颅亦各具有不同的式型。     

Patterns of sexual size dimorphism in Chelonia

Macroevolutionary patterns of sexual size dimorphism (SSD) indicate how sexual selection, natural selection, and genetic and developmental constraints mold sex differences in body size. One putative pattern, known as Rensch's rule, posits that, among species with female‐larger SSD, the relative degree of SSD declines with species' body size, whereas, among male‐larger SSD species, relative SSD increases with size. Using a dataset of 196 chelonian species from all fourteen families, we investigated the correlation in body size evolution between male and female Chelonia and the validity of Rensch's rule for the taxon and within its major clades. We conclude that male–female correlations in body size evolution are high, although these correlations differ among chelonian families. Overall, SSD scales isometrically with body size; Rensch's rule is valid for only one family, Testudinidae (tortoises). Because macroevolutionary patterns of SSD can vary markedly among clades, even in a taxon as morphologically conservative as Testudines, one must guard against inappropriately pooling clades in comparative studies of SSD. The results of the present study also indicate that regression models that assume the x‐variable (e.g. male body size) is measured without statistical error, although frequently reported, will result in erroneous conclusions about phylogenetic trends in sexual size dimorphism. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 108 , 396–413.     

Chelonian mental glands

A survey of 69 of the 74 currently recognized chelonian genera revealed that 21 genera in three families (Emydidae, Platysternidae and Testudinidae) possess paired integumentary glands or gland vestiges in the anterior throat skin. These glands are here termed mental glands; they are holocrine and may be classified morphologically as follows: Class I mental glands are large, complex, multilobed, have specialized ducts, and are found only in the genus Gopherus: Class II mental glands are small, simple sacklike invaginations containing secretory cells or keratinizing cells. The structure of Class II glands varies from distinctive and saccular to shallow keratinized invaginations having no glandular tissue; they are found only in the families Platysternidae and Emydidae. Mental glands occur in 17 of the 22 genera in the subfamily Batagurinae (sensu McDowell, 64); only 2 of 9 genera in the subfamily Emydinae have these glands. The taxonomic occurrence of mental glands suggests that they are primitive structures. The loss of mental glands in most emydines is interpreted as a subfamilial trend toward integumentary simplification.     

Historical Burden In Systematics And The Interrelationships Of 'Parareptiles'

The interrelationships within the clade comprised of turtles, pareiasaurs, and procolophonid-like taxa are investigated via a cladistic analysis incorporating 56 characters. A single most parsimonious tree was found (80 steps, c. i. = 0·8) in which the successive outgroups to turtles are: pareiasaurs, Sclerosaurus, lanthanosuchids, procolophonoids (=Owenetta, Barasaurus and procolophonids), and nyctiphruretians (= nycteroleterids). Thus, as suggested recently by other workers (Reisz, in Fischman, 1993) turtles are the highly modified survivors of a radiation of poorly-known reptiles commonly called ‘parareptiles’. Pareiasaurs are united with turtles on the basis of twenty unambiguous derived features which are absent in other basal amniotes (=‘primitive reptiles’) and reptiliomorph amphibians: for example, the medially located choana, enlarged foramina palatinum posterius, blunt cultriform process, fully ossified medial wall of the prootic, opisthotic-squamosal suture, lateral flange of exoccipital, loss of ventral cranial fissure, thickened braincase floor, ‘pleurosphenoid’ ossification, reduced presacral count, acromion process, trochanter major, reduced fifth pedal digit, and presence of transverse processes on most caudals. Recent phylogenetic proposals linking turtles with captorhinids, with dicynodonts, and with procolophonoids are evaluated. None of the proposed traits supporting the first two hypotheses is compelling. The procolophonoid hypotheses is supported by only one synapomorphy (the slender stapes). All other synapomorphies proposed in favour of the above groupings either occur in many other primitive amniotes, or are not primitive for turtles, or are not primitive for the proposed chelonian sister-group. Nyctiphruretus and Lanthanosuchids and nycteroleterids, often considered to be seymouriamorph amphibians, are demonstrated unequivocally to be amniotes. The ‘rhipaeosaurs’, currently considered to be pareiasaur relatives, are shown to be a heterogenous assemblage of seymouriamorphs, therapsids and nycteroleterids. The phylogeny proposed here indicates that many of the traits of the earliest known turtle, Proganochelys, previously interpreted as unique specialisations, also occur in pareiasaurs and other near outgroups of turtles, and must instead represent the primitive chelonian condition: for example, the wide parietals and the short quadrate flange of the pterygoid. The sequence of acquisition of chelonian traits is discussed: many features once thought to be diagnostic of turtles actually characterize larger groupings of procolophonomorphs, and must have evolved long before the chelonian shell appeared. These traits include most of the chelonian-pareiasaur synapomorphies listed above, and many others which characterize more inclusive groupings found in this analysis. In putting Proganochelys much closer to the main line of chelonian evolution, in elucidating the sequence of acquisition of chelonian traits, and in reducing greatly the number of differences between turtles and their nearest relatives, this study helps bridge one of the major gaps in the fossil record. The failure of previous cladistic analyses to identify correctly the nearest relatives of turtles is attributed to biased character selection, caused by an over-reliance on cranial characters deemed ‘important’ by earlier workers, and by a tendency to shoehorn ‘parareptile’ taxa into phylogenies derived from analyses restricted to ‘mainstream’ groups such as synapsids, diapsids, turtles, and ‘captorhinomorphs’. Many of the synapomorphies that resolve turtle origins are postcranial, and the three nearest outgroups to turtles are all highly bizarre groups which were dismissed as ‘too specialized’ by early workers and continued to be inadequately assessed even by workers using a cladistic framework.     

Association of herpesvirus with fibropapillomatosis of the green turtle Chelonia mydas and the loggerhead turtle Caretta caretta in Florida.

Sea turtle fibropapillomatosis (FP) is a disease marked by proliferation of benign but debilitating cutaneous fibropapillomas and occasional visceral fibromas. Transmission experiments have implicated a chloroform-sensitive transforming agent present in filtered cell-free tumor homogenates in the etiology of FP. In this study, consensus primer PCR methodology was used to test the association of a chelonian herpesvirus with fibropapillomatosis. Fibropapilloma and skin samples were obtained from 17 green and 2 loggerhead turtles affected with FP stranded along the Florida coastline. Ninety-three cutaneous and visceral tumors from the 19 turtles, and 33 skin samples from 16 of the turtles, were tested. All turtles affected with FP had herpesvirus associated with their tumors as detected by PCR. Ninety-six percent (89/93) of the tumors, but only 9% (3/33) of the skin samples, from affected turtles contained detectable herpesvirus. The skin samples that contained herpesvirus were all within 2 cm of a fibropapilloma. Also, 1 of 11 scar tissue samples from sites where fibropapillomas had been removed 2 to 51 wk earlier from 5 green turtles contained detectable herpesvirus. None of 18 normal skin samples from 2 green and 2 loggerhead turtles stranded without FP contained herpesvirus. The data indicated that herpesvirus was detectable only within or close to tumors. To determine if the same virus infected both turtle species, partial nucleotide sequences of the herpesvirus DNA polymerase gene were determined from 6 loggerhead and 2 green turtle samples. The sequences predicted that herpesvirus of loggerhead turtles differed from those of green turtles by only 1 of 60 amino acids in the sequence examined, indicating that a chelonian herpesvirus exhibiting minor intratypic variation was the only herpesvirus present in tumors of both green and loggerhead turtles. The FP-associated herpesvirus resisted cultivation on chelonian cell lines which support the replication of other chelonian herpesviruses. These results lead to the conclusion that a chelonian herpesvirus is regularly associated with fibropapillomatosis and is not merely an incidental finding in affected turtles.     

The Prevalence of Pores and Canals in Leaf Cuticular Membranes. 2. Supplemental Studies

Evidence is presented for the existence of discrete, naturalcuticular pores concomitant with anticlinally-oriented transcuticularcanals found in the mature leaf surfaces of 26 out of 37 taxaamong 19 families. This investigation is an extension of earlierobservations made on 32 other taxa among 14 families. Dewaxed,chemically isolated, adaxial and abaxial cuticular membranesin conjunction with transverse leaf sections were examined usingordinary staining techniques. The ubiquitous pores occur randomlywith no evidence of clustering. Pore and canal diameters averageapprox. 1 µm. Canal lengths are directly related to cuticlethickness. No correlations were found between cuticle thicknessesand either pore frequencies or pore and canal diameters. Evidenceis provided by light microscopy photomicrographs. Leaf cuticles, cuticular membranes, cuticular pores, transcuticular canals, cuticular flanges     

Covariation between clutch size, egg weight and egg shape: comparative evidence for chelonians

The variation in reproductive variables is documented both within and across species of chelonians. At both the generic and family levels, egg weight and clutch size show positive, significant correlations with adult carapace length. There is a negative correlation across both genera and families between clutch size and egg weight after removing the effects of body size, suggesting an evolutionary trade-off between these two life-history characteristics. However, the trade-off is not complete, since clutch size is positively correlated with clutch weight after removing the effects of body size. Terrestrial species lay fewer and larger eggs for their size than freshwater or marine species, but this association is statistically confounded by the fact that chelonian families form ecological groups. There is no significant association between habitat and clutch weight after removing the effects of body size, nor between latitude and either egg weight or clutch size, but temperate species have a heavier clutch weight after removing the effects of body size. Larger species lay eggs that are more spherical, but after controlling for body size, egg shape is not associated with clutch size. The patterns of covariation between adult weight, egg weight and clutch size contrast with those reported for birds and mammals, and some reasons for these differences are discussed.     

First insights into the function of the sawshark rostrum through examination of rostral tooth microwear

Potential roles of the rostrum of sawsharks (Pristiophoridae), including predation and self‐defence, were assessed through a variety of inferential methods. Comparison of microwear on the surface of the rostral teeth of sawsharks and sawfishes (Pristidae) show that microwear patterns are alike and suggest that the elongate rostra in these two elasmobranch families are used for a similar purpose (predation). Raman spectroscopy indicates that the rostral teeth of both sawsharks and sawfishes are composed of hydroxyapatite, but differ in their collagen content. Sawfishes possess collagen throughout their rostral teeth whereas collagen is present only in the centre of the rostral teeth of sawsharks, which may relate to differences in ecological use. The ratio of rostrum length to total length in the common sawshark Pristiophorus cirratus was found to be similar to the largetooth sawfish Pristis pristis but not the knifetooth sawfish Anoxypristis cuspidata. Analysis of the stomach contents of P. cirratus indicates that the diet consists of demersal fishes and crustaceans, with shrimp from the family Pandalidae being the most important dietary component. No prey item showed evidence of wounds inflicted by the rostral teeth. In light of the similarities in microwear patterns, rostral tooth chemistry and diet with sawfishes, it is hypothesised that sawsharks use their rostrum in a similar manner for predation (sensing and capturing prey) and possibly for self‐defence.     

The sternal pore areas of geophilomorph centipedes (Chilopoda: Geophilomorpha)

Most geophilomorph centipedes have segmental clusters of exocrine glands whose opening pores are arranged in more or less well-defined sternal pore areas. We describe here the cuticular structures forming and/or accompanying the gland openings on the sternites and the shape of the pore areas along the body axis in representatives of most geophilomorph families. The cuticular ring around the pore may exhibit either of two forms. In Himantariidae ( Himantarium ) and in Dignathodontidae ( Henia ) the ring looks like a continuous ribbon with a visible suture, whereas in the representatives of the remaining families no suture is seen. As to the distribution of the pores on ventral surface of the body, we record the presence of pores on the last leg-bearing segment of Clinopodes flavidus , whereas that segment was described as poreless in all geophilomorphs. We also provide a taxonomic survey of shape and distribution of pore areas in the individual families, where the pore areas may take very different shapes that we regard as transformational homologues. As for the segmental distribution of sternal pore areas, there is a considerable amount of complexity along the trunk of geophilomorph centipedes, in contrast to the apparently uniform trunk structure.     

Venipuncture techniques in chelonian species

Blood collection is one of the most common diagnostic procedures in chelonian species. Knowledge and case-dependent selection of appropriate sampling sites are necessary to obtain a diagnostic blood sample.     

湖南爬行动物区系与地理区划

本文刊出了湖南已发现的88种爬行动物的名录。报道一种湖南龟类新记录。根据多年的调查资料,分析阐述了湖南爬行动物区系和爬行动物地理区划。     

How does the curvature of the upper beak bone reflect the overlying rhinotheca morphology?

The beak has independently been evolved accompanied by the edentulism in many tetrapod linages, including extant Testudinata and Aves, and its form and function have been greatly diversified. The beak is formed by beak bones and the overlying keratinous cover, although their profiles are different from each other. Therefore, it is difficult to reliably reconstruct the entire profile of the beak in extinct taxa, whose keratinous tissues are rarely preserved. For elucidation of the morphological relationship between beak bone and overlying keratinous cover, we compared the curvature distribution of the culminal profiles of the upper beak bone and the overlying keratinous cover (rhinotheca) with each other using CT‐scan, in 66 extant testudinatan and avian specimens (Aves: 33 genera, 24 families; Testudinata: 12 genera seven families). In both, rhinotheca and beak bone, the curvature of the profile was nearly constant rostral to a certain point, which was defined as the transition point, and the transition points of the rhinotheca and beak bone were close to each other. The profiles of the rhinotheca and beak bone rostral to their transition point were different in curvature and length. However, the ratio between the curvatures of rhinotheca and the beak bone strongly correlated with the arc angle of the rostral culminal profiles of the beak bone. The upper beak profile in extinct taxa is expected to be reconstructed more reliably using the abovementioned relationship between the beak bone and the rhinotheca.     

Plasmodesmatal pores in the torus of bordered pit membranes affect cavitation resistance of conifer xylem

The pit membrane in bordered pits of conifer tracheids is characterized by a porous margo and central thickening (torus), which is traditionally considered to function as an impermeable safety valve against air-seeding. However, electron microscopy based on 33 conifer species, including five families and 19 genera, reveals that pores occur in the torus of 13 of the species studied. The pores have a plasmodesmatal origin with an average diameter of 51 nm and grouped arrangement. Evidence for embolism spreading via pores in tori is supported by the pore sizes, which correspond relatively well with the pressure inducing cavitation. Predictions based on earlier correlations between pit structure and cavitation resistance were only weakly supported for species with punctured tori. Moreover, species with punctured tori are significantly less resistant to cavitation than species with non-punctured tori. Nevertheless, absolute pore diameters must be treated with caution and correlations between theoretical and measured air-seeding pressures are weak. Because most pores appear not to traverse the torus but are limited to one torus pad, only complete pores would trigger air-seeding. Embolism spreading through a leaky torus is not universal across gymnosperms and unlikely to represent the only air-seeding mechanism.     

Redescription of the clam shrimp Lynceus gracilicornis (Packard) (Branchiopoda, Conchostraca, Lynceidae) from Florida, with notes on its biology

Clam shrimps of the family 1.ynccidac differ markedly from all other families of the Conchostraca. The biology and morphology of the genus Lynceus are poorly known. In North America the genus is represented by four species: L. brachyurus, L. mucronatus, L. brevifrons and L. grarcilicornis. The last species, previously known from two localities in Texas, is reported from an ephemeral pond in north Florida. Amoung the characters distinguishing L. gracilicornis from North American congeners arc dimorphic male claspers and a broad, straight rostral margin. Females are distinct in having a smoothly rounded, distal rostra1 margin. External and internal morphology are described using light and scanning electron microscopy. Scanning electron microscopy reveals the presence of frontal setose sensory fields and a small rostral pit in both sexe5. Numerous setal types are described from the claspers and posterior thoracopods. Internal structures include a large anterior hepatopancreas, C-shaped gut and large paired gonads ventro-lateral to the gut.     

Raja koreana, a new species of skate (Elasmobranchii, Rajoidei) from Korea

A new rajid species,Raja koreana, is described from a single adult female specimen, 735 mm in total length, collected off the southwestern coast of the Korean Peninsula. AlthoughR. koreana is included in the group of species characterized by the scapulocoracoid lacking an anterior bridge and having the postventral fenestra expanded, it is unique among the latter in possessing: pectoral girdle propterygium not extending to snout tip; rostral shaft of neurocranium narrow and thick, unsegmented base with filamentous cartilage; snout fleshy; pores of ampullae of Lorenzini densely distributed over much of ventral surface to behind cloaca; most thorns on tail directed anteriorly; tail short; a pair of longitudinally elongated black blotches on middle of dorsal surface of disc when fresh; a pair of black blotches (grayish at center) posteriorly on pectoral fins; ventral surface of dise uniformly blackish-brown, except for areas around pores.     

Development of the turtle carapace: Implications for the evolution of a novel bauplan

The chelonian carapace is composed of the endochondral ribs and vertebrae associated with a specialized dermis. The ribs are found in an aberrant position compared to those of all other tetrapods; they are superficial and dorsal to the limb girdles. This morphological arrangement, which constitutes the unique chelonian Bauplan, is examined from a developmental perspective. Embryos of Chelydra serpentina were studied during stages of carapace development. Tissue morphology, autoradiography, and indirect immunofluorescent localization of adhesion molecules indicate that the outgrowth of the embryonic carapace occurs as the result of an epithelial–mesenchymal interaction in the body wall. A carapacial ridge composed of mesenchyme of the dermis and overlying ectoderm is formed dorsal to the ectodermal boundary between somitic and lateral plate mesoderm. It is the anlage of the carapace margin, in which the ribs will eventually terminate. The ectoderm of the carapacial ridge is thickened into a pseudostratified columnar epithelium, which overlies a condensation in the mesenchyme of the dermis. Patterns of cell proliferation and the distribution of N-CAM and fibronectin in the carapacial ridge are consistent with patterns seen in other structures initiated by epithelial–mesenchymal interactions such as feathers and limb buds. Based on an analogy to this developmental mechanism in the development of the limb skeleton, a further analogy with the evolution of the limbs from lateral fin folds is used to form a hypothesis on the evolution of the carapace from elements of the primitive reptilian integument.