Can spatial sorting associated with spawning migration explain evolution of body size and vertebral number in Anguilla eels?

Spatial sorting is a process that can contribute to microevolutionary change by assembling phenotypes through space, owing to nonrandom dispersal. Here we first build upon and develop the “neutral” version of the spatial sorting hypothesis by arguing that in systems that are not characterized by repeated range expansions, the evolutionary effects of variation in dispersal capacity and assortative mating might not be independent of but interact with natural selection. In addition to generating assortative mating, variation in dispersal capacity together with spatial and temporal variation in quality of spawning area is likely to influence both reproductive success and survival of spawning migrating individuals, and this will contribute to the evolution of dispersal‐enhancing traits. Next, we use a comparative approach to examine whether differences in spawning migration distance among 18 species of freshwater Anguilla eels have evolved in tandem with two dispersal‐favoring traits. In our analyses, we use information on spawning migration distance, body length, and vertebral number that was obtained from the literature, and a published whole mitochondrial DNA‐based phylogeny. Results from comparative analysis of independent contrasts showed that macroevolutionary shifts in body length throughout the phylogeny have been associated with concomitant shifts in spawning migration. Shifts in migration distance were not associated with shifts in number of vertebrae. These findings are consistent with the hypothesis that spatial sorting has contributed to the evolution of more elongated bodies in species with longer spawning migration distances, or resulted in evolution of longer migration distances in species with larger body size. This novel demonstration is important in that it expands the list of ecological settings and hierarchical levels of biological organization for which the spatial sorting hypothesis seems to have predictive power.     

Correlation between Hox code and vertebral morphology in archosaurs

The relationship between developmental genes and phenotypic variation is of central interest in evolutionary biology. An excellent example is the role of Hox genes in the anteroposterior regionalization of the vertebral column in vertebrates. Archosaurs (crocodiles, dinosaurs including birds) are highly variable both in vertebral morphology and number. Nevertheless, functionally equivalent Hox genes are active in the axial skeleton during embryonic development, indicating that the morphological variation across taxa is likely owing to modifications in the pattern of Hox gene expression. By using geometric morphometrics, we demonstrate a correlation between vertebral Hox code and quantifiable vertebral morphology in modern archosaurs, in which the boundaries between morphological subgroups of vertebrae can be linked to anterior Hox gene expression boundaries. Our findings reveal homologous units of cervical vertebrae in modern archosaurs, each with their specific Hox gene pattern, enabling us to trace these homologies in the extinct sauropodomorph dinosaurs, a group with highly variable vertebral counts. Based on the quantifiable vertebral morphology, this allows us to infer the underlying genetic mechanisms in vertebral evolution in fossils, which represents not only an important case study, but will lead to a better understanding of the origin of morphological disparity in recent archosaur vertebral columns.     

Vertebral number in adders, Vipera berus: direct and indirect effects on growth

Given the importance of body size, and thereby growth rate, for many reproductive parameters in snakes, morphological traits conferring an advantage in terms of growth may be important targets of selection. Studies have demonstrated effects of vertebral number of growth rate in garter snakes. In this study effects of total number of body vertebrae and of number of abnormal body vertebrae (obtained by counting number of ventral scutes and number of abnormal scutes) on growth rate in free-ranging male and female adders, Vipera berus (L.), are examined by calculating directional performance gradients (estimating linear effects) and stabilizing performance gradients (estimating curvilinear effects). After controlling for body size (SVL) female adders demonstrated a significant positive directional gradient for vertebral number, and a significant interaction between body size and vertebral number, showing that females with more vertebrae have higher size-specific growth rates, and that this effect is strongest among small, fast growing individuals. Females also showed a weak stabilizing effect of abnormal vertebrae. Males, on the other hand, showed a positive directional gradient for number of abnormal vertebrae, whereas no effect of vertebral number was observed. Indirect effects of the same variables were evaluated by use of path analysis. Generally, indirect effects were weak and did not substantially increase the amount of explained variance in growth rate. Field data showed that the correlation between vertebral number and growth rate in females was stronger in years with higher overall growth rate. To evaluate whether vertebral number and food availability show an interactive effect I used captive born juvenile adders in an experiment with two different food levels. The experiment confirmed the field data. No relationship between vertebral number and growth was observed in the low food level group, whereas in the high food level group a significant positive correlation was demonstrated. Finally, the heritability of vertebral number was examined using a mother-offspring regression and a full-sib analysis. The estimated heritabilities were 0.30 and 0.39, respectively. From these results it is concluded that both vertebral number and abnormal vertebral number may significantly affect growth in adders, but that this effect may differ between sexes and among years.     

Variability and constraint in the mammalian vertebral column

Patterns of vertebral variation across mammals have seldom been quantified, making it difficult to test hypotheses of covariation within the axial skeleton and mechanisms behind the high level of vertebral conservatism among mammals. We examined variation in vertebral counts within 42 species of mammals, representing monotremes, marsupials and major clades of placentals. These data show that xenarthrans and afrotherians have, on average, a high proportion of individuals with meristic deviations from species' median series counts. Monotremes, xenarthrans, afrotherians and primates show relatively high variation in thoracolumbar vertebral count. Among the clades sampled in our dataset, rodents are the least variable, with several species not showing any deviations from median vertebral counts, or vertebral anomalies such as asymmetric ribs or transitional vertebrae. Most mammals show significant correlations between sacral position and length of the rib cage; only a few show a correlation between sacral position and number of sternebrae. The former result is consistent with the hypothesis that adult axial skeletal structures patterned by distinct mesodermal tissues are modular and covary; the latter is not. Variable levels of correlation among these structures may indicate that the boundaries of prim/abaxial mesodermal precursors of the axial skeleton are not uniform across species. We do not find evidence for a higher frequency of vertebral anomalies in our sample of embryos or neonates than in post-natal individuals of any species, contrary to the hypothesis that stabilizing selection plays a major role in vertebral patterning.     

Ecomorphology of Locomotion in Labrid Fishes

The Labridae is an ecologically diverse group of mostly reef associated marine fishes that swim primarily by oscillating their pectoral fins. To generate locomotor thrust, labrids employ the paired pectoral fins in motions that range from a fore-aft rowing stroke to a dorso-ventral flapping stroke. Species that emphasize one or the other behavior are expected to benefit from alternative fin shapes that maximize performance of their primary swimming behavior. We document the diversity of pectoral fin shape in 143 species of labrids from the Great Barrier Reef and the Caribbean. Pectoral fin aspect ratio ranged among species from 1.12 to 4.48 and showed a distribution with two peaks at about 2.0 and 3.0. Higher aspect ratio fins typically had a relatively long leading edge and were narrower distally. Body mass only explained 3% of the variation in fin aspect ratio in spite of four orders of magnitude range and an expectation that the advantages of high aspect ratio fins and flapping motion are greatest at large body sizes. Aspect ratio was correlated with the angle of attachment of the fin on the body (r = 0.65), indicating that the orientation of the pectoral girdle is rotated in high aspect ratio species to enable them to move their fin in a flapping motion. Field measures of routine swimming speed were made in 43 species from the Great Barrier Reef. Multiple regression revealed that fin aspect ratio explained 52% of the variation in size-corrected swimming speed, but the angle of attachment of the pectoral fin only explained an additional 2%. Labrid locomotor diversity appears to be related to a trade-off between efficiency of fast swimming and maneuverability in slow swimming species. Slow swimmers typically swim closer to the reef while fast swimmers dominate the water column and shallow, high-flow habitats. Planktivory was the most common trophic associate with high aspect ratio fins and fast swimming, apparently evolving six times.