Divergence in the shell morphology of the land snail genus Aegista (Pulmonata: Bradybaenidae) under phylogenetic constraints

The role of natural selection in phenotypic evolution is central to evolutionary biology. Phenotypic evolution is affected by various factors other than adaptation, and recent focus has been placed on the effects of phylogenetic constraints and niche conservatism on phenotypic evolution. Here, we investigate the relationship between the shell morphology and habitat use of bradybaenid land snails of the genus Aegista and clarify the causes of the divergence in shell morphology among phylogenetically related species. The results of ancestral state reconstruction showed that arboreal species have evolved independently from ground‐dwelling species at least four times. A significant association was found between shell shape and habitat use, despite the existence of a certain degree of phylogenetic constraint between these traits. A principal component analysis showed that arboreal species tend to have a relatively high‐spired shell with a narrow umbilicus. By contrast, ground‐dwelling species have a low‐spired shell with a wide umbilicus. Although the latitude and elevation of the sampling locations showed no relationship with shell morphology, the geology of the sampling locations affected the shell size of arboreal species. The development of a well‐balanced shell shape is one effective method for reducing the cost of locomotion under the force of gravity in each life habitat, resulting in the divergence in shell morphology and the independent evolution of morphologically similar species among different lineages. The present study suggests that ecological divergence is probably the cause of shell morphology divergence in land snails. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 114 , 229–241.     

Evolutionary mechanisms of rib loss in anurans: a comparative developmental approach

ABSTRACT The presence of free ribs is presumed to be a primitive morphological character observed only in a few families of Recent anurans, whereas the absence of ribs has been considered to be a derived condition that is widespread within this order. A comparative study of rib development based on representatives of several anuran lineages (Alytes, Bombina, Bufo, Discoglossus, Hyla, Pelobates, Pelodytes, Rana, and Xenopus) reveals a previously undetected diversity of developmental features in the formation and interaction between neural arches and ribs. The absence of free ribs at premetamorphic or later stages is verified in some groups, but we present for the first time evidence of the existence of larval rib rudiments in others, both in the anterior (Rana, Hyla) and posterior (Bufo, Discoglossus, Pelobates) presacral regions. Heterochrony seems to have played a major role in the processes underlying rib reduction. The intracolumnar differences between anterior (V(2)-V(4)) and posterior (V(5)-V(8)) regions are based on perturbations in the timing of early differentiation. Furthermore, a clear shift in the relative timing of ossification among evolutionary lineages was detected. In this respect Xenopus has a highly derived condition. The use of the morphological character of "rib loss" in phylogenetic analyses must be reconsidered due to the different convergent developmental paths described here. The phylogenetic analysis of a "sequence units" matrix of rib development is compared with current anuran phylogenies. Some evolutionary information appears to be clearly present in the ontogenetic data of this "missing morphology," but its value for evolutionary inferences is rather limited.     

Genetic evidence for natural hybridization and apparent introgression between freshwater snail species (Viviparus ater and V. contectus)

We examined six allozyme markers and shell morphology to study gene flow between the naturally sympatric freshwater snail species Viviparus ater and V. contectus in Lake Garda, Italy, using offspring from experimental crosses and snails collected from natural populations. Hybrid offspring obtained from experimental crosses and hybrids collected from natural populations were heterozygous at four or five out of five diagnostic loci. Shell morphology was a poor predictor of the hybrid status of individual snails. Backcrossed offspring from experimentally bred hybrids and either of the parental species morphologically resembled the parental species. There was no evidence of segregation distortion at the six loci. Linkage analysis revealed one pair of linked loci (GPI and PNP). These cross experiments indicated a Mendelian type hybrid system in which gene introgression may occur. F₁ hybrids were found at four out of six sampling sites of Lake Garda, Italy. Local frequencies of F₁ hybrids ranged from 0% to 1.6% (estimated average = 0.74%) of the total population including both species. Alleles typical of V. ater were found at low frequencies in V. contectus at all six sampling sites. Alleles typical of V. contectus were found at low frequencies in V. ater at three out of six sampling sites. This is consistent with the hypothesis of introgression in both directions.     

Different thermal conditions of lakes affect host–parasite systems: A case study of Viviparus contectus (Millet, 1813) and digenean trematodes

1. Thermal disturbance of aquatic ecosystems directly and/or indirectly affects interspecific interactions, including parasitism. Both hosts and parasites respond differently to environmental changes, thus, predicting how host–parasite systems behave under the influence of disturbance remains a challenge. The aim of the study was to check how the differences in thermal conditions of lakes affect life-history traits of hosts and the level of parasitism, using a Viviparus contectus–digenean trematodes model.
2. Overall, we examined 480 individuals of V. contectus collected from a thermally polluted lake (TPL) and a natural lake (NL). Host features, including body size and fecundity, as well as the prevalence and species richness of digenean trematodes in snail populations were investigated.
3. We found that V. contectus from the TPL were significantly larger, heavier, and females were more fertile than snails collected from the NL. A total of 20.4% of the collected snails were infected with digenean larvae. The species richness of parasites was twice as high in the NL compared to the TPL (six and three species, respectively). A significant difference in the percentage of snails infected with parasites was identified between both types of lakes, with a higher prevalence of V. contectus in the NL (31.3%) compared to the TPL (7.3%).
4. These results indicate that host–parasite systems follow the environmental changes in lakes due to thermal pollution by increasing fertility and metabolism rate of viviparid hosts and by decreasing the prevalence and diversity of digenean trematodes.

     

Conchometrics, systematics and distribution of Melanopsis (Mollusca: Gastropoda) in the Levant

In this conchometric study, the systematics and distribution of the freshwater gastropod Melanopsis in the Levant are described. Of the ten species found, three are widespread, two have narrow distributions and five are known only from their type locality. Five smooth‐shelled species are recognized (buccinoidea, ammonis, dircaena, khabourensis and meiostoma). Within M. buccinoidea, Jordan Valley populations may belong to a separate subspecies. M. ammonis is clearly differentiated from buccinoidea of the nearby Jordan Valley, but less so from buccinoidea of more distant sites; differences between ammonis and meiostoma are significant but not diagnostic. Five species have ribbed shells. Within costata, four subspecies are recognized (in the northern Orontes, upper Jordan, Sea of Galilee, and in southernmost parts of the Levant). Throughout the Levant, M. buccinoidea frequently hybridizes with M. costata and fossil evidence suggests that these species have been hybridizing for the last 1.5 Myr. M. saulcyi differs from M. costata in its narrower shell and shorter, bumpier ribs and frequently hybridizes with M. buccinoidea throughout the Levant. Specimens from Homs differ from those of the Jordan Valley in their higher figurativity index, fewer ribs and lower rib density. M. germaini differs from M. costata in its more numerous ribs, M. pachya in its shorter ribs, and M. infracincta in its bumpy shell in which each rib has huge tubercles, with a pronounced ridge flanking the columella. Our conclusion that there are ten species in the Levant differs from previous studies that suggested only two subspecies of one species (or superspecies). This difference could stem from (1) our use of nonstandard as well as standard conchometrics, (2) a reappraisal of the importance of the shell vs. the radula in intrageneric systematics, and (3) differences of opinion on the subspecies concept. © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society, 2005, 144 , 229?260.     

Vicariance and convergence in Magellanic and New Zealand long‐looped brachiopod clades (Pan‐Brachiopoda: Terebratelloidea)

Phylogenetic reconstructions using parsimony, maximum likelihood (ML), and Bayesian relaxed molecular clock analyses of ～2850 nucleotides of nuclear‐encoded small and large subunit ribosomal DNAs (SSU and LSU rDNAs) from 14 long‐looped (terebratelloid) ingroup and four short‐looped (terebratulidine) outgroup brachiopod taxa, together with ML analyses of ～663 nucleotides of mitochondrial cytochrome oxidase subunit 1 (cox1) from 12 terebratelloid taxa, show that deep divergence separates taxa endemic to waters in the vicinity of New Zealand from those with a Magellanic distribution around South America and Antarctica. This deep divergence also separates Magellanic Terebratella dorsata from New Zealand Terebratella sanguinea, showing that they are not congeneric. Instead, they belong to separate ‘Magellanic’ (MAG) and ‘New Zealand’ (NZ) clades that first diverged about 82 Mya (95% highest posterior density, 48–120 Mya), correlating with separation between the NZ and Antarctic tectonic plates. Sequence analyses also reveal (1) that the Antarctic endemic taxa Magellania fragilis and Magellania joubini are not congeneric with Magellania venosa, suggesting that their previous placement in Aerothyris should be restored, and (2) that divergence between Antarctic and NZ species of the terebratulide Liothyrella occurred much later than plate separation, perhaps because of continuing gene flow caused by long‐lived larvae. The topology of the rDNA and cox1 gene trees implies that radial ornament of the shell (‘ribbing’) has been gained (and/or lost) independently within the MAG and NZ clades. Radial ribs are widespread in articulate brachiopods throughout the Phanerozoic, but no comparisons of brachiopod rib morphology and morphogenesis have been published. Our comparisons of transverse shell mid‐sections in the scanning electron microscope reveal no obvious evidence of differences in morphology between independently gained ribs. We also consider several ways in which ribs may affect fitness, including effects on hydrodynamics. Only scanty and inconclusive evidence is available, but we suggest that effects (if any) are likely to be of small magnitude; adaptive value of brachiopod shell ribs remains to be demonstrated. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162 , 631–645.     

16层螺旋CT三维重建与DR平片检查对外伤性肋骨骨折的诊断价值比较研究

目的:研究16层螺旋电子计算机断层扫描(CT)三维重建与数字化摄影(DR)平片检查对外伤性肋骨骨折的诊断价值,为临床诊治提供参考。方法:将2017年6月至2018年6月期间于本院接受诊治的82例外伤性肋骨骨折患者作为研究对象,所有患者均接受16层螺旋CT三维重建与DR平片检查,观察并记录患者的骨折发生部位,并比较两种诊断方法对外伤性肋骨骨折诊断的准确率、灵敏度、特异性、阳性预测值、阴性预测值以及漏诊情况。结果:82例外伤性肋骨骨折患者经影像学与临床诊断明确发生骨折179处,多发性骨折发生率为62.20%,单发性骨折发生率为37.80%,骨折肋骨段位中4-10段骨折发生率最高为69.83%,骨折肋骨水平阶段中腋肋骨折发生率最高为59.78%。相较于DR平片,16层螺旋CT三维重建诊断外伤性肋骨骨折的准确率、灵敏度、特异性、阳性预测值、阴性预测值更高,漏诊率更低,差异有统计学意义(P0.05)。结论:相较于DR平片检查,应用16层螺旋CT三维重建检查外伤性肋骨骨折能明显提高临床诊断的准确率、灵敏度和特异性,减少漏诊,可为临床诊治提供更可靠的信息,值得临床推广。     

REMOTE BIOMINERALIZATION IN DIVARICATE RIBS OF STRIGILLA AND SOLECURTUS (TELLINOIDEA: BIVALVIA)

Deposits composed of aragonite prisms, which were formed afterthe outer shell layer, have been found at the posterior steepslopes of divaricate ribs in two species of Strigilla and anothertwo of Solecurtus. These prisms have their axes oriented perpendicularto the outer shell surface and differ in morphology from fibresof the surface-parallel composite prisms forming the outer shell.They display crystalline features indicating that, unlike crystalsforming the outer shell surface, their growth front was free,unconstrained by the mantle or periostracum. These particulardeposits are called free-growing prisms (FGPs). In these generathe periostracum is clearly not the substrate for biomineralizationand, upon formation, does not adhere to the steep slope of ribs,but detaches at the rib peak and reattaches towards the posterior,just beyond the foot of the posterior scarps of ribs. In thisway, a sinus or open space developed between the internal surfaceof the periostracum and the outer shell surface along each steeprib slope. These spaces could remain filled with extrapallialfluid after the mantle advances beyond that point during shellsecretion. FGPs grow within this microenvironment, out of contactwith the mantle. Other species with divaricate ribs do not developFGPs simply because the periostracum adheres tightly to both ribslopes (which are never so steep as in Solecurtus and Strigilla).FGPs constitute one of the rare cases of remote biomineralizationin which aragonite is produced and direct contact with the mantlenever takes place. (Received 22 November 1999; accepted 20 February 2000)     

Rib fabrication in Ostreoidea and Plicatuloidea (Bivalvia,Pteriomorphia) and its evolutionary significance

Ribs of Ostreoidea and Plicatuloidea are defined as antimarginal, that is, perpendicular to the margin throughout growth. Morphogenetically, these ribs are unique, since, unlike radial ribs, they are secreted by a homogeneous mantle margin. Based also on the reconstructed shell secretion cycle in Bivalvia, we propose that ribs of Ostreoidea and Plicatuloidea are formed by a mantle margin which, upon extension from the shell margin, stretches and folds by taking the preformed ribs as templates. In extending perpendicular to the margin (as in all Bivalvia growing isometrically), such a mantle extends the rib pattern antimarginally. Ribs of this kind are purely mechanical structures, as their arrangement depends on the mechanical properties of the mantle and on the environmental conditions. This explains the high irregularity of such ribbing patterns. The presence of antimarginal ribs in both the Ostreoidea and Plicatuloidea sheds light on their origin. The first known oyster, Actinostreon cristadifformis, probably derived from an antimarginally ribbed Prospondylidae gen. indet. in the Late Permian or Early Triassic. Antimarginally ribbed Triassic species formerly included in Placunopsis originated both the Dimyidae Atreta in the Late Triassic and Enantiostreon in the Mid Triassic, which was transitional to Plicatulidae. Therefore, Dimyidae and Plicatulidae are closely connected and grouped under Plicatuloidea, to which Ostreoidea is phylogenetically unrelated.     

Morphological evolution of silica scales in the freshwater genus Synura (Stramenopiles)

A high degree of morphological variability is expressed between the ornately sculptured siliceous scales formed by species in the chrysophycean genus, Synura. In this study, we aimed to uncover the general principles and trends underlying the evolution of scale morphology in this genus. We assessed the relationships among thirty extant Synura species using a robust molecular analysis that included six genes, coupled with morphological characterization of the species‐specific scales. The analysis was further enriched with addition of morphological information from fossil specimens and by including the unique modern species, Synura punctulosa. We inferred the phylogenetic position of the morphologically unique S. punctulosa, to be an ancient Synura lineage related to S. splendida in the section Curtispinae. Some morphological traits, including development of a keel or a labyrinth ribbing pattern on the scale, appeared once in evolution, whereas other structures, such as a hexagonal meshwork pattern, originated independently several times over geologic time. We further uncovered numerous construction principles governing scale morphology and evolution, as follows: (i) scale roundness and pore diameter decreased during evolution; (ii) elongated scales became strengthened by a higher number of struts or ribs; (iii) as a consequence of scale biogenesis, scales with spines possessed smaller basal holes than scales with a keel and; and (iv) the keel area was proportional to scale area, indicating its potential value in strengthening the scale against breakage.     

The ontogeny of the shell in side‐necked turtles,with emphasis on the homologies of costal and neural bones

Although we are starting to understand the molecular basis of shell development based on the study of cryptodires, basic comparative ontogenetic data for the other major clade of living turtle, the pleurodires, are largely missing. Herein, the developmental and phylogenetic relation between the bony shell and endoskeleton of Pleurodira are examined by studying histological serial sections of nine specimens of three different species, including an ontogenetic series of Emydura subglobosa. Emphasis is given to the portion of the carapace in which ribs and vertebral spinous processes become part of the carapace. Central questions are how neurals and costals are formed in pleurodiran turtles, whether costals and neurals are of endoskeletal or exoskeletal origin, and what ontogenetic factors relate to neural reduction of some Pleurodira. The neurals and costals do not develop as independent ossification centers, but they are initial outgrowths of the periosteal collar of endoskeletal ribs and neural arches. Slightly later in development, the ossification of both shell elements continues without a distinct periosteum but by metaplastically ossifying precondensed soft‐tissue integumentary structures. Through ontogeny, ribs of the turtles studied are closely associated with the hypaxial intercostalis musculature while epaxial interspinalis musculature connects the neural arches. We here propose an alternative structural hypothesis for the neural reduction and, ultimately, the complete loss of the neural series. The complete reduction of neurals in Emydura spp. may be linked to heterochrony, accompanied by a restricted influence of epaxial musculature and epidermal–dermal interaction in shell bone formation. J. Morphol., 2008. © 2008 Wiley‐Liss, Inc.     

ACCELERATED EVOLUTION OF LAND SNAILS MANDARINA IN THE OCEANIC BONIN ISLANDS: EVIDENCE FROM MITOCHONDRIAL DNA SEQUENCES

An endemic land snail genus Mandarina of the oceanic Bonin (Ogasawara) Islands shows exceptionally rapid evolution not only of morphological and ecological traits, but of DNA sequence. A phylogenetic relationship based on mitochondrial DNA (mtDNA) sequences suggests that morphological differences equivalent to the differences between families were produced between Mandarina and its ancestor during the Pleistocene. The inferred phylogeny shows that species with similar morphologies and life habitats appeared repeatedly and independently in different lineages and islands at different times. Sequential adaptive radiations occurred in different islands of the Bonin Islands and species occupying arboreal, semiarboreal, and terrestrial habitat arose independently in each island. Because of a close relationship between shell morphology and life habitat, independent evolution of the same life habitat in different islands created species possesing the same shell morphology in different islands and lineages. This rapid evolution produced some incongruences between phylogenetic relationship and species taxonomy. Levels of sequence divergence of mtDNA among the species of Mandarina is extremely high. The maximum level of sequence divergence at 16S and 12S ribosomal RNA sequence within Mandarina are 18.7% and 17.7%, respectively, and this suggests that evolution of mtDNA of Mandarina is extremely rapid, more than 20 times faster than the standard rate in other animals. The present examination reveals that evolution of morphological and ecological traits occurs at extremely high rates in the time of adaptive radiation, especially in fragmented environments.     

Mechanics of sculpture formation in Magadiceramus? rangatira rangatira (Inoceramidae,Bivalvia) from the Upper Cretaceous of New Zealand

The surface sculpture of the inoceramid bivalve Magadiceramus? rangatira rangatira consists of commarginal ribs and curious, transverse wrinkles. The wrinkles typically are at a high angle or orthogonal to the shell margin (‘antimarginal’) and thus differ from purely radial structures. They show features of distribution and morphology that reveal them to be products of margin‐parallel compression of the shell‐secreting mantle and its adjacent, flexible, uncalcified periostracum. The interaction of wrinkles with commarginal ribs indicates that the ribs also formed as folds of the mantle margin. During growth, commarginal folding caused withdrawal of the entire mantle margin towards the umbo, with a consequent reduction in perimeter length. Measurement of specimens indicates that fabrication of the commarginal ribs resulted in the magnitude of commarginal shortening that is required for the formation of transverse wrinkles. We infer that early in ontogeny, at the first development of these sculptures, the wrinkles resulted entirely from mantle contraction and resultant commarginal shortening. With subsequent growth, total wrinkling included a component of ‘pre‐wrinkling’ inherited from the preceding growth stage; the contribution of pre‐wrinkling to total wrinkling increased with shell size. The proposed mechanical model is two‐phase. First, the transversely corrugated (pre‐wrinkled) mantle and periostracum advanced and secreted a slightly concave growth increment. Secondly, the mantle subsequently contracted to create a commarginal rib and increase the number and amplitude of transverse wrinkles. This model is consistent with a homogeneous mantle lacking any differentiated and specialized rib‐constructing segments.     

Rib fractures under anterior–posterior dynamic loads: Experimental and finite-element study

The purpose of this study was to investigate whether using a finite-element (FE) mesh composed entirely of hexahedral elements to model cortical and trabecular bone (all-hex model) would provide more accurate simulations than those with variable thickness shell elements for cortical bone and hexahedral elements for trabecular bone (hex–shell model) in the modeling human ribs. First, quasi-static non-injurious and dynamic injurious experiments were performed using the second, fourth, and tenth human thoracic ribs to record the structural behavior and fracture tolerance of individual ribs under anterior–posterior bending loads. Then, all-hex and hex–shell FE models for the three ribs were developed using an octree-based and multi-block hex meshing approach, respectively. Material properties of cortical bone were optimized using dynamic experimental data and the hex–shell model of the fourth rib and trabecular bone properties were taken from the literature. Overall, the reaction force–displacement relationship predicted by both all-hex and hex–shell models with nodes in the offset middle-cortical surfaces compared well with those measured experimentally for all the three ribs. With the exception of fracture locations, the predictions from all-hex and offset hex–shell models of the second and fourth ribs agreed better with experimental data than those from the tenth rib models in terms of reaction force at fracture (difference <15.4%), ultimate failure displacement and time (difference <7.3%), and cortical bone strains. The hex–shell models with shell nodes in outer cortical surfaces increased static reaction forces up to 16.6%, compared to offset hex–shell models. These results indicated that both all-hex and hex–shell modeling strategies were applicable for simulating rib responses and bone fractures for the loading conditions considered, but coarse hex–shell models with constant or variable shell thickness were more computationally efficient and therefore preferred.     

Molecular phylogeny and evolution of phenotype in silica‐scaled chrysophyte genus Mallomonas

The evolution of phenotypes is highly understudied in protists, due to the dearth of morphological characters, missing fossil record, and/or unresolved phylogeny in the majority of taxa. The chrysophyte genus Mallomonas (Stramenopiles) forms species‐specific silica scales with extraordinary diversity of their ornamentation. In this paper, we molecularly characterized three additional species to provide an updated phylogeny of 43 species, and combined this with evaluations of 24 morphological traits. Using phylogenetic comparative methods, we evaluated phylogenetic signal in traits, reconstructed the trait evolution, and compared the overall phylogenetic and morphological diversity. The majority of traits showed strong phylogenetic signal and mostly dynamic evolution. Phylogenetic relatedness was often reflected by the phenotypic similarity. Both V‐rib and dome are very conservative structures that are presumably involved in precise scale overlap and bristle attachment, respectively. Based on modern species, it seems the dome firstly appeared on apical and/or caudal scales, and only later emerged on body scales. Bristle was presumably present in the common ancestor and gradually elongated ever since. However, most other morphological traits readily changed during the evolution of Mallomonas.     

Broken ribs: Paleopathological analysis of costal fractures in the human identified skeletal collection from the Museu Bocage,Lisbon, Portugal (late 19th to middle 20th centuries)

Although rarely reported in the anthropological literature, rib fractures are commonly found during the analysis of human skeletal remains of past and modern populations. This lack of published data precludes comparison between studies and restricts an accurate understanding either of the mechanisms involved in thoracic injuries or their impact on past societies. The present study aimed: 1) to report rib fracture prevalence in 197 individuals, 109 males, and 88 females, with ages at death ranging from 13 to 88 years old, from the Human Identified Skeletal Collection, Museu Bocage, Portugal (late 19th‐middle 20th centuries); 2) to test the hypothesis that a higher prevalence of rib stress fractures existed in the 133 individuals who died from respiratory diseases, in a period before antibiotics. The macroscopic analysis revealed 23.9% (n = 47) of individuals with broken ribs. 2.6% (n = 124) out of 4,726 ribs observed were affected. Males presented more rib fractures, and a significantly higher prevalence was noted for older individuals. Fractures were more frequently unilateral (n = 34), left sided (n = 19) and mainly located on the shaft of ribs from the middle thoracic wall. Nineteen individuals presented adjacent fractured ribs. Individuals who died from pulmonary diseases were not preferentially affected. However, a higher mean rate of fractures was found in those who died from pneumonia, a scenario still common nowadays. Since rib involvement in chest wall injury and its related outcomes are important issues both for paleopathology and forensic anthropology, further investigations are warranted. Am J Phys Anthropol 2009. © 2009 Wiley‐Liss, Inc.     

Morphological study of the anthropoid thoracic cage: scaling of thoracic width and an analysis of rib curvature

While a relatively broad thorax and strongly curved ribs are widely regarded as common features of living hominoids, few studies have quantitatively examined these traits by methods other than calculating the chest index. The present study aims to quantify variations in thoracic cage morphology for living anthropoids. The odd-numbered ribs (first to eleventh) were articulated with the corresponding vertebrae and the cranial and lateral views subsequently photographed. Rib profiles were digitized in both views and line-fitted by a Bézier curve to create a three-dimensional morphological data set. When thoracic cage width was scaled against body mass, Hylobates (and possibly Pongo) plotted above non-hominoid anthropoids at almost all rib levels, while Pan did not differ from non-hominoid anthropoids. The overall pattern of the normalized thoracic width differed between Hylobates and other hominoids. In Hylobates, an upward convex curve was seen between the first and seventh ribs while a more linear pattern was observed in Pan and Pongo. This result quantitatively confirmed that the barrel-shaped thoracic cage in Hylobates can be distinguished from the funnel-shaped form in other hominoids. Conversely, all hominoids shared two distinct features in the upper half-thorax: (1) a pronounced dorsal protrusion of the proximal part of the rib in accordance with ventral displacement of the thoracic spine and (2) a relatively medially projecting sternal end. Although these features are likely to provide some mechanical advantage in orthograde and/or suspensory positional behaviors, they were barely present in the suspensory Ateles. An erratum to this article can be found at     

Disentangling the effects of intraspecies variability,phylogeny, space,and climate on the evolution of shell morphology in endemic Greek land snails of the genus Codringtonia

Extensive variation in land snail shell morphology has been widely documented, although few studies have attempted to investigate the ecological and evolutionary drivers of this variation. Within a comparative phylogenetic framework, we investigated the temporal and spatial evolution of the shell morphology of the Greek endemic land snail genus Codringtonia. The contribution of both inter‐ and intraspecies shell differentiation in the overall shell variability is assessed. The effect of climate, space, and evolutionary history on the shell variability was inferred using a variance partitioning framework. For Codringtonia species, intraspecies divergence of shell traits contributes substantially to the overall shell variability. By decomposing this variability, it was shown that the overall shell size of Codringtonia clades is phylogenetically constrained, related to early speciation events, and strongly affected by large‐scale spatial variability (latitudinal gradient). The effect of climate on shell size cannot be disentangled from phylogeny and space. Shell and, to a larger extent, aperture shape are not phylogenetically constrained, and appear to be mostly related to conspecific populations divergence events. Shell shape is substantially explained by both climate and space that greatly overlap. Aperture shape is mainly interpreted by medium to small‐scale spatial variables. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 796–813.     

Internal embryonic brooding and development in the southern Australian micro-snail Tricolia rosea (Angas, 1867) (Vetigastropoda: Phasianellidae: Tricoliinae)*

ABSTRACT

Internal embryonic brooding has been suggested as an adaptation to enhance reproductive success in minute gastropods. It is rare in vetigastropods, previously known in only two species of Spectamen Iredale, 1924 (Solariellidae) from South Africa. Herein it is confirmed in the temperate Australian micro-snail Tricolia rosea (Angas, 1867), with up to 46 embryos observed within a gravid female. Embryos are brooded to an advanced stage, possessing a translucent, calcified shell and a calcified operculum. The initial protoconch is colourless, spirally sculptured and delineated by a consistent axial demarcation at the 0.75 whorl mark, when it measures 320?µm across. Beyond this, the second part of the protoconch is tinted pink, the strong spiral sculpture continues but the ribs and interstices are broader, with smoother surface microsculpture. At the 1.1 to 1.125 whorl mark the protoconch measures 400?µm across when transition to smooth teleoconch sculpture occurs. No brooded embryos possessed teleoconch sculpture. The potential relationship of protoconch morphology to embryonic development, hatching, feeding and release are considered. The mechanism of fertilisation is unknown, but embryos in a brood are at the same developmental stage. Unanswered questions in embryonic development and problems with protoconch terminologies in vetigastropods are also discussed.