Behavioral patterns in the early-stage antipredator response change after tail autotomy in adult wall lizards.

Autotomy is a drastic antipredator defense consisting of the voluntary shedding of a body part to escape from the predators. The loss of a body part may impair locomotion, feeding or mating, so animals may face a higher predation risk shortly after autotomy. Thus, until regeneration is completed, prey may adjust their behavior to reduce predation risk, and this could involve secondary costs. We assessed the effect of tail loss on the antipredator behavior of wall lizards (Podarcis muralis), comparing the behavior of tailed and tailless individuals exposed to a predatory snake (Coronella austriaca) scent, under controlled experimental conditions. Tailless lizards spent significantly more time performing behaviors with antipredatory significance (e.g., moving slowly), whereas tailed individuals performed exploratory walking for significantly more time. Moreover, tailless lizards spent more time basking, which probably increases the effectiveness of their cryptic design and decreases detection by predators. Lizards intensified the tongue flick rates when exposed to a pungent control or snake scents, as compared to their response to a neutral control. Besides, both tailed and tailless lizards intensified some aspects of their antipredator behavior (walking slowly and avoiding refuge use) when exposed to snake scent, which indicates discrimination of the smell of predatory snakes. Lizards decreased refuge use when exposed to predator scents, probably because the refuges are evaluated as unsafe due to a high concentration of snake scents. To conclude, our experiments showed that, after losing their tails, wall lizards modify their behavior in a way that likely minimizes predation risk.

Allocation costs of regeneration: tail regeneration constrains body growth under low food availability in juvenile lizards.

The balance of energy allocated to development and growth of different body compartments may incur allocation conflicts and can thereby entail physiological and evolutionary consequences. Regeneration after autotomy restores the functionality lost after shedding a body part but requires a strong energy investment that may trade-off with other processes, like reproduction or growth. Caudal autotomy is a widespread antipredator strategy in lizards, but regeneration may provoke decreased growth rates in juveniles that could have subsequent consequences. Here, we assessed the growth of intact and regenerating hatchling wall lizards (Podarcis muralis) exposed to different food regimens. Regenerating juveniles presented slightly but significantly lower body growth rates than individuals with intact tails when facing low food availability, but there were no differences when food was supplied ad libitum. Regenerating individuals fed ad libitum increased their ingestion rates compared to intact ones during the period of greatest tail growth, which also reveals a cost of tail regeneration. When resources were scarce, hatchlings invested more in tail regeneration in relation to body growth, rather than delay regeneration to give priority to body growth. We propose that, in juvenile lizards, regeneration could be prioritized even at the expense of body growth to restore the functionality of the lost tail, likely increasing survivorship and the probability to reach reproductive maturity. Our study indicates that food availability is a key factor for the occurrence of trade-offs between regeneration and other growth processes, so that environmental conditions would be determinant for the severity of the costs of regeneration.

An integrative analysis of the short-term effects of tail autotomy on thermoregulation and dehydration rates in wall lizards.

Maintaining body temperature is essential for the optimal performance of physiological functions. Ectotherms depend on external heat sources to thermoregulate. However, thermoregulation may be constrained by body condition and hydration state. Autotomy (i.e., the voluntary shed of a body part) evolved in various animal lineages and allowed surviving certain events (such as predator attacks), but it may affect body condition and volume/surface ratios, increase dehydration and constrain thermoregulation. In the framework of a general analysis of the evolution of autotomy, here we assessed the effects of tail loss on the thermal preferences and evaporative water loss rates (EWL) in the lizard Podarcis bocagei, integrating the thermal and hydric factors. We did not observe shifts in the thermal preferences of experimentally autotomized lizards when compared to the controls, which contradicted the hypothesis that they would raise preferred temperature to increase metabolic rates and accelerate regeneration. Evaporative water loss rates were also similar for tailed and tailless individuals, suggesting negligible increase of water loss through the injury and no specific ecophysiological responses after autotomy. Therefore, the changes observed in autotomized lizards in the field are to be considered primarily behavioural, rather than physiological, and thermoregulation could be secondarily affected by behavioural compensations for an increased predation risk after autotomy. Functional studies are necessary to understand how lizards' interaction with the environment is altered after autotomy, and further studies including different dehydration levels would be useful to fully understand the effect of water shortage on lizards' performance after caudal autotomy.

The movement dynamics of autotomized lizards and their tails reveal functional costs of caudal autotomy.

Autotomy has evolved independently several times in different animal lineages. It frequently involves immediate functional costs, so regeneration evolved in many instances to restore the functionality of that body part. Caudal autotomy is a widespread antipredator strategy in lizards, although it may affect energy storage, locomotion dynamics, or survival in future encounters with predators. Here, we assessed the effect of tail loss on the locomotor performance of wall lizards (Podarcis muralis), as well as the recovery of locomotor functionality of lizards with regenerated tails, and the movement dynamics of shed tails that were either intact or having regenerated portions. Tail loss had no effect on locomotion over unhindered spaces, possibly due to compensation between a negative effect on the stride of front limbs, and a positive effect of losing mass and friction force. We found a clear negative impact of tail loss on locomotion in spaces with interspersed obstacles, in which tailed lizards jumped larger distances when leaving the obstacles. Besides, lizards that used the tail to push off the ground were able to approach the obstacles from further, so that the tail seemed to be useful when used during jumping. Regeneration fully restores lizard's locomotor capacities, but tail antipredator value, as indicated by the intensity of post-autotomic movements, is only partially retrieved. From these results, we propose that, together with the recovery of post-autotomy antipredator capacities, the restoration of the organismal locomotor performance may have been an important, yet frequently neglected factor in the evolution of lizard's regeneration ability.

Holothurian (Echinodermata) diversity and distribution in the central Cantabrian Sea and the Avilés Canyon System (Bay of Biscay).

Here we report on the holothurian fauna at the Avilés Canyon System (Natura 2000 Site of Community Importance), central Cantabrian Sea, northern Spain. The samples were collected during several cruises covering the shelf, the continental slope and the abyssal depth. We identified 174 specimens, belonging to 35 species of the seven orders of class Holothuroidea. Depth was the main structuring agent. Multivariate analysis allowed the differentiation of four main assemblages which corresponded to abyssal plain, lower continental slope, upper continental slope, and continental shelf. Depth had a significant effect on holothurian species richness (it increased with depth at an approximate rate of 1.7 species 1000 m-1), which is consistent with previously described global patterns.