Inter-sexual contests in the hermit crab <Emphasis Type="Italic">Pagurus bernhardus</Emphasis>: females fight harder but males win more encounters

Contests often occur between members of the same sex when they compete for access to mates, but inter-sexual contests may occur over access to other essential resources such as food or shelter. Despite the possibility that such contests are common, most studies focus on male fighting, and very few have analysed fights between males and females. Because males and females differ physically and physiologically, fighting ability or resource-holding potential (RHP) may also be subject to inter-sexual variation. In this study, we investigate size-controlled inter-sexual contests over the ownership of empty gastropod shells in the hermit crab Pagurus bernhardus. During these fights, there are two roles, attacker and defender, and we use a fully orthogonal experimental design to compare the performance of males and females in each role, when fighting either a male or female opponent. Although females fight more intensely, male attackers have an advantage when compared to females playing the attacker role, as they are more likely to evict the defender from its shell and thus win the resource. Further, in the defender role, male defenders are subject to shorter attacks than female defenders. The differences in agonistic performance could not be attributed to differences in perceived resource value between the sexes or to differences in body or weapon size. There are clear differences in the agonistic behaviour of males and females, and this possibility should be incorporated into models of contest behaviour. In particular, evolutionarily stable strategies may be expected to vary with sex ratios.     

Binary individual recognition in hermit crabs

One mechanism that permits the maintenance of dominance hierarchies is individual recognition, defined as the ability of an animal to recognize a conspecific on the basis of one or more identifying cues, and to associate it with experiences of victories or defeats that the animal has gained from preceding encounters with that particular individual. We examined whether the long-clawed hermit crab, Pagurus longicarpus, could differentiate between unfamiliar and familiar opponents. The experimental protocol was designed to control in pairs of interacting individuals several factors together, such as status and relative size of the opponent, as well as species, quality, and fit of the inhabited shell. The hermit crabs were more reactive and their agonistic level was higher in unfamiliar than in familiar pairs; in addition, betas were more prone to initiate an interaction with unfamiliar than with familiar alphas. The alternative explanation—that the ability to discriminate between familiar and novel shells can explain our results per se— was tested following, in part, Jackson and Elwoods (1989) protocol for Pagurus bernhardus and was, at least for this species, rejected. This study did not determine whether a true individual recognition occurs, but demonstrated that P. longicarpus categorizes the individuals into two heterogeneous subgroups, thus being capable of a binary discrimination among opponents.Communicated by T. Czeschlik     

Shell acquisition by hermit crabs: which tactic is more efficient?

Gastropod shells are limited resources for most hermit crab species, acting as primary factors in various aspects of their biology. To investigate the efficacy of different behavioral tactics adopted for their acquisition (locomotion, attendance at shell-supplying sites, interactions with conspecifics in aggregation) we conducted observations and experiments at a salt marsh in New England (USA). Locomotion, fast and meandering, significantly increased the chances of encountering empty shells and conspecifics. However, shell exchanges were rare. Simulated gastropod predation sites quickly attracted a larger number of hermit crabs than the other shell-supplying sites tested (live and dead conspecifics, live snails) and induced the rapid occupancy of all the empty shells offered, usually by the first crabs arriving at the site. Aggregations seemed not to function as shell exchange markets, as previously suggested for several other species. In the short run, exploitation seems to be more efficient for the acquisition of new shells by Pagurus longicarpus. In the long run, it is the density of nondestructive gastropod predators that regulates the availability of new shells of good quality in the pool available to this hermit crab species.     

Fighting behavior in hermit crabs: the combined effect of resource-holding potential and resource value in Pagurus longicarpus

A study was conducted to determine whether asymmetries in both resource-holding potential (RHP) and resource value (RV) influence dominance and fighting behavior in the hermit crab Pagurus longicarpus. A total of 120 groups of three crabs were observed for 10 min in four experiments that tested all diverse combinations of equal/different RHP (i.e. ‘body size’) and equal/different RV (i.e. ‘shell size’ and ‘shell quality’). In a fifth experiment, dominant and subordinate individuals of the same size category (26 groups) were forced to enter shells of opposite quality than those previously occupied, and then the behavior of the reconstituted original groups was observed for additional 10 min. As expected, crabs in lower quality shells were more willing to initiate and to escalate fights. However, their attacks were directed to any crab of the group, independently of the defender's shell quality, and the fight duration did not vary with the different value of the resources at stake. This may indicate that P. longicarpus is unable to assess the quality of the shells available in its social environment but bases its tactical decisions during fights solely on its own resource. This suggestion was confirmed by the change in the fighting behavior of crabs whose shell quality was experimentally altered. This manipulation induced an overall increase in the intensity of aggression, drastic modification of crab behavior, and inversion of the hierarchy even though these crabs have had previous experiences of wins/losses and were familiar to the other members of the group. In this species, large crab size and/or the occupancy of adequate (and oversized) shells appeared to be the most likely determinant of contest resolution. Individuals seemed to retain a memory of the previously held resource and behaved accordingly.     

Reduced Density and Nest Survival of Ground‐Nesting Songbirds Relative to Earthworm Invasions in Northern Hardwood Forests

Abstract: European earthworms (Lumbricus spp.) are spreading into previously earthworm‐free forests in the United States and Canada and causing substantial changes, including homogenization of soil structure, removal of the litter layer, and reduction in arthropod abundance and species richness of understory plants. Whether these changes affect songbirds that nest and forage on the forest floor is unknown. In stands with and without earthworms in the Chequamegon‐Nicolet National Forest, Wisconsin (U.S.A.), we surveyed for, monitored nests of, and measured attributes of habitat of Ovenbirds (Seiurus aurocapillus) and Hermit Thrushes (Catharus guttatus), both ground‐dwelling songbirds, and we sampled earthworms at survey points and nests. Bird surveys indicated significantly lower densities of Ovenbirds and Hermit Thrushes in relation to Lumbricus invasions at survey point and stand extents (3.1 and 15–20 ha, respectively). Modeling of Ovenbird nest survival (i.e., the probability that nestlings successfully fledge) indicated that lower survival probabilities were associated with increased sedge cover and decreased litter depth, factors that are related to Lumbricus invasions, possibly due to reduced nest concealment or arthropod abundance. Our findings provide compelling evidence that earthworm invasions may be associated with local declines of forest songbird populations.     

Using Distance from Putative Source Woodlots to Predict Occurrence of Forest Birds in Putative Sinks

Abstract:  For some species of forest-breeding birds, productivity is much higher in large forest fragments than in small forest fragments and is higher than the level needed to replace individuals within the fragment (sources). Thus large forest fragments potentially provide excess individuals to neighboring small fragments where productivity is much lower and not adequate for replacement (sinks). We used occurrence data and distances between putative sources and sinks for four species of forest birds with this pattern of demography in large and small forest fragments to predict occupancy in small fragments. For the Ovenbird (  Seiurus aurocapilla ), Wood Thrush (  Hylocichla mustelina ), Veery ( Catharus fuscescens ), and Rose-breasted Grosbeak ( Pheucticus ludovicianus ), distance from large woodlots was a significant predictor of occurrence in small woodlots. Distance from large woodlots did not predict occupancy of Red-eyed Vireos (  Vireo olivaceus ), a species with no apparent difference in productivity in relation to fragment size. Neither vegetation features nor area of the small woodlots adequately explained patterns of occupancy for any species. These results suggest that maintenance of large, productive forest fragments benefit regional occurrence of forest-breeding birds by providing individuals to neighboring small woodlots. Limited reproduction in these small woodlots means that they may act as a reservoir for excess individuals in the case of catastrophic events that result in vacancies in source woodlots. Conservation and sound forest management of small forests should be directed particularly toward those closer to large regional forests.     

The past ownership of a resource affects the agonistic behavior of hermit crabs

Resource value is often considered the most important nonstrategic variable in the fighting behavior of invertebrates. In our study, we tested whether the experience of shells of a given quality, occupied in the recent past, might affect the agonistic behavior of the hermit crab Pagurus longicarpus. We analyzed the fights battled by 84 test crabs against size-matched unknown rivals before (premanipulation phase) and after (postmanipulation phase) having modified the quality of the domicile shell of the contestants. Specifically, we compared the behavior of crabs that had been subject to either a worsening or an improvement in the quality of their shell with crabs that, although being similarly subject to a shell manipulation, occupied a shell of the same quality as in the premanipulation phase. We found that the crabs subject to a worsening in the quality of their shell were more aggressive than those subject to its improvement and that the former were even more aggressive than those that occupied a bad quality shell also in the premanipulation phase. Crabs seemed not to gather information about the opponent’s shell during fights or not to use this information, most often behaving in accordance with the quality of the domicile shell. These results are clear in showing the role played by the experience of a previously occupied shell, also confirming that the agonistic behavior of P. longicarpus is mainly based on decision rules of the type “own resource value dependency.”