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.