Evolving possibilities: Post-embryonic axial elongation in salamanders with biphasic (Eurcyea cirrigera, E. longicauda, E. quadridigitata) and paedomorphic life cycles (Eurycea nana and Ambystoma mexicanum). Submitted Acta Zoologica.
ABSTRACT: Typically the number of vertebrae an organism will have post-embryonically is determined during embryogenesis via the development of paired somites. Our research investigates the phenomenon of post-embryonic vertebral addition in salamander tails. We describe body and tail growth, and patterns of postsacral vertebral addition and elongation in context with caudal morphology for four plethodontids (Eurycea), and one ambystomatid. Eurycea nana and A. mexicanum have paedomorphic life cycles; E. cirrigera, E. longicauda and E. quadridigitata are biphasic. Specimens were collected, borrowed and/or purchased, and cleared and stained for bone and cartilage. Data collected include snout-vent length (SVL), tail length (TL), vertebral counts and centrum lengths. Eurycea species with biphasic life cycles had TLs that surpassed SVL following metamorphosis. Tails in paedomorphic species elongated but rarely exceeded body length. Larger TLs were associated with more vertebrae and longer vertebrae in all species. We observed that rates of postsacral vertebral addition varied little amongst species. Regional variation along the tail becomes prominent following metamorphosis in biphasic developers. In all species vertebrae in the posterior one-half of the tail taper towards the tip. We suggest a developmental link might exist between the ability to continually add vertebrae and regeneration in salamanders.
Project description:Macaques (genus Macaca) are known to have wide variation in tail length. Within each species group tail length varies, which could be associated with a phylogenetic trend seen in caudal vertebral morphology. We compared numbers and lengths of caudal vertebrae in species of the fascicularis group, M. assamensis (sinica group), M. nemestrina (silenus group), and those obtained from reports for an additional 11 species. Our results suggest different trends in number and lengths. The caudal vertebral length profiles revealed upward convex patterns for macaques with relative tail lengths of ?15%, and flat to decreasing for those with relative tail lengths of ?12%. They varied between species groups in terms of the lengths of proximal vertebrae, position and length of the longest vertebra, numbers and lengths of distal vertebrae, and total number of vertebrae. In silenus and sinica group, the vertebral length is the major skeletal determinant of tail length. On the other hand, the vertebral number is the skeletal determinant of tail length in the fascicularis group. Tail length variation among species groups are caused by different mechanisms which reflect the evolutionary history of macaques.
Project description:BACKGROUND:Body shape can be both a consequence and cause of a species' evolution and ecology. There are many examples of phenotypes associated with specific ecological niches, likely as a result of specific selective regimes. A classic example of this is the phenotypic change associated with colonization of caves, including body and limb elongation. However, studies explicitly testing for differences in body shape between cave-dwelling and non-cave-dwelling lineages have been limited and so the role of the cave environment in determining morphological characteristics is still not completely understood. Here we examine variation in body shape among 405 individuals representing 20 species in the salamander genus Eurycea (Plethodontidae) and select outgroups exhibiting great diversity in morphology, ecological niche, and life history. RESULTS:After analyzing morphometric data in a phylogenetic context using phylogenetic MANOVA and examination of the phylomorphospace, we found significant differences in body shape among cave-dwelling and non-cave-dwelling species and between aquatic and terrestrial species. Notably, limb elongation and reduced body and tail size characterized cave-dwelling species. Terrestrial species also exhibited elongation of the limbs and digits. We also observed differences in shape variance among paedomorphic and biphasic species. Our results suggest that the functional limitations imposed by habitat and life history played a key role in the evolution of body shape in this group in the context of their phylogenetic history.
Project description:The transformation of ancestral phenotypes into novel traits is poorly understood for many examples of evolutionary novelty. Ancestrally, salamanders have a biphasic life cycle with an aquatic larval stage, a brief and pronounced metamorphosis, followed by a terrestrial adult stage. Repeatedly during evolution, metamorphic timing has been delayed to exploit growth-permissive environments, resulting in paedomorphic salamanders that retain larval traits as adults. We used thyroid hormone (TH) to rescue metamorphic phenotypes in paedomorphic salamanders and then identified quantitative trait loci (QTL) for life history traits that are associated with amphibian life cycle evolution: metamorphic timing and adult body size. We demonstrate that paedomorphic tiger salamanders (Ambystoma tigrinum complex) carry alleles at three moderate effect QTL (met1-3) that vary in responsiveness to TH and additively affect metamorphic timing. Salamanders that delay metamorphosis attain significantly larger body sizes as adults and met2 explains a significant portion of this variation. Thus, substitution of alleles at TH-responsive loci suggests an adaptive pleiotropic basis for two key life-history traits in amphibians: body size and metamorphic timing. Our study demonstrates a likely pathway for the evolution of novel paedomorphic species from metamorphic ancestors via selection of TH-response alleles that delay metamorphic timing and increase adult body size.
Project description:Life cycle strategies have evolved extensively throughout the history of metazoans. The expression of disparate life stages within a single ontogeny can present conflicts to trait evolution, and therefore may have played a major role in shaping metazoan forms. However, few studies have examined the consequences of adding or subtracting life stages on patterns of trait evolution. By analysing trait evolution in a clade of closely related salamander lineages we show that shifts in the number of life cycle stages are associated with rapid phenotypic evolution. Specifically, salamanders with an aquatic-only (paedomorphic) life cycle have frequently added vertebrae to their trunk skeleton compared with closely related lineages with a complex aquatic-to-terrestrial (biphasic) life cycle. The rate of vertebral column evolution is also substantially lower in biphasic lineages, which may reflect the functional compromise of a complex cycle. This study demonstrates that the consequences of life cycle evolution can be detected at very fine scales of divergence. Rapid evolutionary responses can result from shifts in selective regimes following changes in life cycle complexity.
Project description:BACKGROUND: Habitat variation strongly influences the evolution of developmentally flexible traits, and may drive speciation and diversification. The plethodontid salamander Eurycea tynerensis is endemic to the geologically diverse Ozark Plateau of south-central North America, and comprises both strictly aquatic paedomorphic populations (achieving reproductive maturity while remaining in the larval form) and more terrestrial metamorphic populations. The switch between developmental modes has occurred many times, but populations typically exhibit a single life history mode. This unique system offers an opportunity to study the specific ecological circumstances under which alternate developmental and life history modes evolve. We use phylogenetic independent contrasts to test for relationships between a key microhabitat feature (streambed sediment) and this major life history polymorphism. RESULTS: We find streambed microstructure (sediment particle size, type and degree of sorting) to be highly correlated with life-history mode. Eurycea tynerensis is paedomorphic in streams containing large chert gravel, but metamorphoses in nearby streams containing poorly sorted, clastic material such as sandstone or siltstone. CONCLUSION: Deposits of large chert gravel create loosely associated streambeds, which provide access to subsurface water during dry summer months. Conversely, streambeds composed of more densely packed sandstone and siltstone sediments leave no subterranean refuge when surface water dries, presumably necessitating metamorphosis and use of terrestrial habitats. This represents a clear example of the relationship between microhabitat structure and evolution of a major developmental and life history trait, and has broad implications for the role of localized ecological conditions on larger-scale evolutionary processes.
Project description:The avian tail played a critical role in the evolutionary transition from long- to short-tailed birds, yet its ontogeny in extant birds has largely been ignored. This deficit has hampered efforts to effectively identify intermediate species during the Mesozoic transition to short tails. Here we show that fusion of distal vertebrae into the pygostyle structure does not occur in extant birds until near skeletal maturity, and mineralization of vertebral processes also occurs long after hatching. Evidence for post-hatching pygostyle formation is also demonstrated in two Cretaceous specimens, a juvenile enantiornithine and a subadult basal ornithuromorph. These findings call for reinterpretations of Zhongornis haoae, a Cretaceous bird hypothesized to be an intermediate in the long- to short-tailed bird transition, and of the recently discovered coelurosaur tail embedded in amber. Zhongornis, as a juvenile, may not yet have formed a pygostyle, and the amber-embedded tail specimen is reinterpreted as possibly avian. Analyses of relative pygostyle lengths in extant and Cretaceous birds suggests the number of vertebrae incorporated into the pygostyle has varied considerably, further complicating the interpretation of potential transitional species. In addition, this analysis of avian tail development reveals the generation and loss of intervertebral discs in the pygostyle, vertebral bodies derived from different kinds of cartilage, and alternative modes of caudal vertebral process morphogenesis in birds. These findings demonstrate that avian tail ontogeny is a crucial parameter specifically for the interpretation of Mesozoic specimens, and generally for insights into vertebrae formation.
Project description:Based on morphological and genetic evidence we evaluated the taxonomic status of a newly discovered forest-dwelling population of skink (genus Scincella) from the Keo Seima Wildlife Sanctuary, Mondulkiri Province, Cambodia. From phylogenetic analysis of a 668-bp fragment of the mtDNA COI and diagnostic morphological characters we allocate the newly discovered population to the Scincella reevesii-S. rufocaudata species complex and describe it as Scincella nigrofasciata sp. nov. The new skink species can be distinguished from all other Southeast Asian congeners by the following combination of morphological characters: snout-vent length (SVL) 40.0-52.6 mm; relative tail length (TaL/SVL ratio) 1.25-1.94; prefrontals in broad contact; infralabials 6; primary temporals 2; relative forelimb length (FIL/SVL ratio) 0.20-0.22; relative hindlimb length (HIL/SVL ratio) 0.30-0.33; relative forearm length (FoL/SVL ratio) 0.14-0.16; adpressed forelimbs and hind limbs either overlapping (0.4-2.2 mm) or separated (1.9-2.3 mm); midbody scale rows 32-33, paravertebral scales 69-74, vertebral scales 65-69; dorsal scales between dorsolateral stripes 8; comparatively slender fingers and toes, subdigital lamellae under fourth toe 15-17; dark discontinuous regular dorsal stripes 5-7; distinct black dorsolateral stripes, narrowing to lateral sides and extending to 52%-86% of total tail length. We provide additional information on the holotype of Scincella rufocaudata (Darevsky & Nguyen, 1983), and provide evidence for the species status of Scincella rupicola. Our discovery brings the number of Scincella species in Cambodia to five and emphasizes the incompleteness of knowledge on the herpetofaunal diversity of this country.
Project description:SummaryBackgroundThere is a close relationship between the vertebral trabecular morphology and the condition of the associated disc.ObjectiveThe relationship between disc degeneration and vertebral trabecular inhomogeneity is unclear. This study aimed to analyse the regional changes of vertebral trabecular morphology after disc degeneration.MethodsThirty male Sprague–Dawley rats were randomly assigned to five groups. Group 1 served as an experimental group for the assessment of disc degeneration induced by needle puncture. Group 2 served as a sham group for trabecular morphology analysis. In Group 3, rats had their tail bent between the eighth and tenth coccygeal vertebrae. In Group 4, the tail of rats was bent with a compression load of 4.5 N. In Group 5, rats first underwent disc degeneration induced by a needle puncture before their tail was bent with a compressive load of 4.5 N. Magnetic resonance imaging was performed on all groups, and histological examination was performed on rodents from Group 1. The ninth coccygeal vertebrae of rats from Groups 2–5 were scanned by Micro-computed tomography. Trabecular morphologic changes were assessed in the concave and convex regions by bone volume fraction, trabecular number, trabecular thickness and trabecular separation.ResultsVertebral trabecular morphology in the concave region improved significantly, whereas the convex region was of significantly lower trabecular morphologic parameters with disc degeneration. The difference in trabecular morphologic parameters between the convex and concave regions increased significantly after disc degeneration.ConclusionDisc degeneration promotes regional inhomogeneity in the vertebral trabecular morphology, with the convex region of the vertebrae having the worse trabecular bone morphology than the concave region.The translational potential of this articleOur study indicates that disc degeneration promotes regional inhomogeneity in the vertebral trabecular morphology. Regional variations in trabecular microarchitecture are helpful to predict vertebral fragility. This may help to elucidate the mechanisms by which disc degeneration contributes to vertebral fracture.
Project description:Very little is known about genetic factors that regulate life history transitions during ontogeny. Closely related tiger salamanders (Ambystoma species complex) show extreme variation in metamorphic timing, with some species foregoing metamorphosis altogether, an adaptive trait called paedomorphosis. Previous studies identified a major effect quantitative trait locus (met1) for metamorphic timing and expression of paedomorphosis in hybrid crosses between the biphasic Eastern tiger salamander (Ambystoma tigrinum tigrinum) and the paedomorphic Mexican axolotl (Ambystoma mexicanum). We used existing hybrid mapping panels and a newly created hybrid cross to map the met1 genomic region and determine the effect of met1 on larval growth, metamorphic timing, and gene expression in the brain. We show that met1 maps to the position of a urodele-specific chromosome rearrangement on linkage group 2 that uniquely brought functionally associated genes into linkage. Furthermore, we found that more than 200 genes were differentially expressed during larval development as a function of met1 genotype. This list of differentially expressed genes is enriched for proteins that function in the mitochondria, providing evidence of a link between met1, thyroid hormone signaling, and mitochondrial energetics associated with metamorphosis. Finally, we found that met1 significantly affected metamorphic timing in hybrids, but not early larval growth rate. Collectively, our results show that met1 regulates species and morph-specific patterns of brain transcription and life history variation.
Project description:Mice deficient in growth differentiation factor 11 (GDF11) signaling display anterior transformation of axial vertebrae and truncation of caudal vertebrae. However, the in vivo molecular mechanisms by which GDF11 signaling regulates the development of the vertebral column have yet to be determined. We found that Gdf11 and Acvr2b mutants are sensitive to exogenous RA treatment on vertebral specification and caudal vertebral development. We show that diminished expression of Cyp26a1, a retinoic acid inactivating enzyme, and concomitant elevation of retinoic acid activity in the caudal region of Gdf11(-/-) embryos may account for this phenomenon. Reduced expression or function of Cyp26a1 enhanced anterior transformation of axial vertebrae in wild-type and Acvr2b mutants. Furthermore, a pan retinoic acid receptor antagonist (AGN193109) could lessen the anterior transformation phenotype and rescue the tail truncation phenotype of Gdf11(-/-) mice. Taken together, these results suggest that GDF11 signaling regulates development of caudal vertebrae and is involved in specification of axial vertebrae in part by maintaining Cyp26a1 expression, which represses retinoic acid activity in the caudal region of embryos during the somitogenesis stage.