Mechanisms of coexistence and competition between ants and land hermit crabs in a Bahamian archipelago

Ants and land crabs are common inhabitants of many coastal and insular communities across the tropics and subtropics, and yet direct evidence of interspecific competition between ants and land crabs has only recently been documented. I conducted a series of observational and manipulative experiments to further elucidate the mechanisms of competition, as well as coexistence, in these two groups in an archipelago of small Bahamian islands. Diel baiting trials demonstrated a significant temporal difference in foraging activity between the land hermit crab, Coenobita clypeatus (Herbst), and ant Brachymyrmex obscurior Forel, suggesting this is one mechanism underlying their coexistence on small oceanic islands. Reciprocal manipulative baiting experiments, in which one of a pair of species was removed from baits, documented that aggressive interspecific interactions underlie patterns of complementary distribution and temporal turnover at rich food resources. This was true for competition between hermit crabs and B. obscurior, and between B. obscurior and a second ant species, Dorymyrmex pyramicus Roger. Negative species associations at baits were found to be common throughout an archipelago of 69 small islands. A trade-off in exploitative and interference abilities may be a second mechanism allowing species coexistence on these small islands. Interspecific interactions such as competition and predation may occur commonly between ants and land crabs and have important consequences for the structure and function of tropical and subtropical insular ecosystems.     

Invasive ants compete with and modify the trophic ecology of hermit crabs on tropical islands

Invasive species can dramatically alter trophic interactions. Predation is the predominant trophic interaction generally considered to be responsible for ecological change after invasion. In contrast, how frequently competition from invasive species contributes to the decline of native species remains controversial. Here, we demonstrate how the trophic ecology of the remote atoll nation of Tokelau is changing due to competition between invasive ants (Anoplolepis gracilipes) and native terrestrial hermit crabs (Coenobita spp.) for carrion. A significant negative correlation was observed between A. gracilipes and hermit crab abundance. On islands with A. gracilipes, crabs were generally restricted to the periphery of invaded islands. Very few hermit crabs were found in central areas of these islands where A. gracilipes abundances were highest. Ant exclusion experiments demonstrated that changes in the abundance and distribution of hermit crabs on Tokelau are a result of competition. The ants did not kill the hermit crabs. Rather, when highly abundant, A. gracilipes attacked crabs by spraying acid and drove crabs away from carrion resources. Analysis of naturally occurring N and C isotopes suggests that the ants are effectively lowering the trophic level of crabs. According to δ¹⁵ N values, hermit crabs have a relatively high trophic level on islands where A. gracilipes have not invaded. In contrast, where these ants have invaded we observed a significant decrease in δ¹⁵ N for all crab species. This result concurs with our experiment in suggesting long-term exclusion from carrion resources, driving co-occurring crabs towards a more herbivorous diet. Changes in hermit crab abundance or distribution may have major ramifications for the stability of plant communities. Because A. gracilipes have invaded many tropical islands where the predominant scavengers are hermit crabs, we consider that their competitive effects are likely to be more prominent in structuring communities than predation. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Land hermit crab (Coenobita clypeatus) densities and patterns of gastropod shell use on small Bahamian islands

Aim To examine patterns of abundance, density, size and shell use in land hermit crabs, Coenobita clypeatus (Herbst), occurring on three groups of small islands, and to determine how these variables change among islands. Location Small islands in the Central Exuma Cays and near Great Exuma, Bahamas. Methods Land hermit crabs were captured in baited pitfall traps and were separately attracted to baits. A mark–recapture technique was used in conjunction with some pitfall traps monitored for three consecutive days. The size of each crab and the type of adopted gastropod shell were recorded, along with physical island variables such as total island area, vegetated area, island perimeter, elevation and distance to the nearest mainland island. Results Relative abundances, densities and sizes of crabs differed significantly among the three island groups. Densities of land hermit crabs were as high as 46 m⁻² of vegetated island area. In simple and multiple linear regressions, the only variable that was a significant predictor of the abundance of hermit crabs was the perimeter to area ratio of the island. Patterns of gastropod shell use varied significantly among the island groups, and the vast majority of adopted shells originated from gastropod species that inhabit the high intertidal and supratidal shorelines of the islands. Main conclusions Although densities of land hermit crabs varied, they were relatively high on many islands, and land hermit crabs may play an important role in these insular food webs. Patterns of shell use may be strongly restricted by island geomorphology: irregular shorelines provide relatively more habitat for the gastropod species that account for the majority of adopted shells and the steep sides of the islands prevent the accumulation of marine gastropod shells. The size of adult hermit crabs appears to be limited by the relatively small gastropod shells available, while the abundance of hermit crabs may be limited by the number of shells available.     

Interference and exploitation components in interespecific competition between sympatric intertidal hermit crabs

This study was designed to evaluate the effect of interference and exploitation competition in shell partitioning between two hermit crab species (Pagurus criniticornis and Clibanarius antillensis). Field samples revealed that shells of the gastropod Cerithium atratum were the main resource used by both hermit crab species and that Pagurus used eroded or damaged shells in higher frequency than Clibanarius. The exploitative ability of each species was compared between species in the laboratory using dead gastropod (Cerithium) baits to simulate predation events and signalize newly available shells to hermit crabs. Pagurus reached the baits more rapidly than Clibanarius, but this higher exploitative ability did not explain shell utilization patterns in nature. Another experiment evaluated the dominance hierarchy between these two hermit crab species and revealed that Clibanarius was able to outcompete Pagurus for higher quality shells in agonistic encounters. This higher interference competitive ability of Clibanarius in relation to Pagurus may explain field observations. Nevertheless, Pagurus may be responsible to enhance shell availability to other hermit crab species that have lower ability to find and use newly available shells. Differently, the poorer condition of shells used by Pagurus, the higher ability of this species to attend gastropod predation events and its higher consumption rate by shell-breaking crabs (Menippe nodifrons) may increase its predation risks, thus revealing the disadvantages of such an exploitative competitive strategy for hermit crabs.     

Plant species persistence and turnover on small Bahamian islands

I conducted surveys of the plant species occupying 136 small islands in the Exuma Cays and 58 small islands near Andros, Bahamas. Most species occurred on relatively few islands, and most islands contained relatively few species. Identities of the most common species differed between the two archipelagos. Comparisons with earlier surveys revealed species extinctions and immigrations. Turnover was relatively low on both a per island and a per species basis on both archipelagos, although significant spatial variation in turnover rates between archipelagos was found. Most islands experienced no turnover; islands on which turnover did occur were larger and had higher species richness. Likewise, most species did not turnover, although much variation existed in turnover rates among those that did. Experimental introductions of two species to very small islands naturally devoid of vegetation revealed that these islands could support plant life. One species survived on eight of ten islands for >9 years, including the effects of a moderate (class 2) hurricane. This hurricane caused substantial damage and loss of plant biomass, but resulted in no species extinctions on 30 small islands. Data for the small islands in this region, now spanning almost a decade, reveal that most populations are persistent over periods of years to decades, rarely going extinct or immigrating. Even moderate hurricanes seem to have little impact on species compositions.     

Why small hermit crabs have large shells

Frequent shell exchanges among hermit crabs imply the enigmatic circumstance that large crabs frequently obtain large shells from smaller crabs. This seeming anomaly is explored as a key to the shell resource system. It is hypothesized to reflect how, where, and how often shells become available to the crabs. Shells become available infrequently, as snails die, and are available to the crabs for only a brief time before they become inaccessible. The standing crop of empty shells is almost always low and is irrelevant to rates of shell turnover in the crab population. Crabs are most likely to encounter shells of the wrong size, and the chance of encountering a shell of the desired size decreases as a crab grows. Snails and crabs are usually found on different portions of the shore; thus, crabs must make “foraging trips” for shells. Under this regime of shell supply, a crab will get a suitable shell the fastest when it accepts any fresh shell that is larger than its initial shell. It can then trade with other crabs to improve its shell fit. This behavior will make small crabs into a regular source of large shells for large crabs, and a shell exchange ritual will be strongly favored because both participants will benefit. Shells are an unusual resource because they are the object of both competitive and mutualistic interactions. This ambiguous quality is revealed in the intraspecific and interspecific responses of crabs to each other and to shells.     

Resource partitioning and competition for shells between intertidal hermit crabs on the outer coast of Washington

Summary Resource partitioning was quantified for 6 species of intertidal hermit crabs in the genus Pagurus, that occur on the outer coast of Washington. This, together with field evidence of shell shortage and with laboratory experiments to quantify the mechanism of interactions for shells, allowed estimation of the relative intensities of inter-and intraspecific competition between these species. The findings were that: (1) the magnitude of intraspecific competition was greater than any single interspecific competitive effect for all of the species; and (2) the relative proportion of intraspecific competition was greater for the middle and upper intertidal species than for the lower intertidal species. Studies at several outer coast sites supported these generalizations. Both of these findings are consistent with the hypothesis that competitive divergence has occurred in the past. The structure of the outer coast hermit crab assemblage is compared with that of the San Juan Archipelago hermit crab assemblage. Differences between the two do not seem to be the result of adaptive responses to the presence of more competing species in the former group.     

Alternative methods of measuring competition applied to two Australian hermit crabs

Summary This article describes the intertidal hermit crab species assemblage at One Tree Island, Great Barrier Reef, Australia. Competition for shells between the two most abundant species, Clibanarius virescens and Calcinus latens, is studied in more detail. Competition appears to be primarily exploitative. The relative intensities of inter- and intra-specific competition between this pair of species are estimated using two different methods. The first is based on habitat overlap data in conjunction with a mathematical model of shell population dynamics described in Abrams (1980). The second method is more direct, and is based on following the fate of marked empty shells. Results of the two methods are very similar. This supports the validity of assumptions made in applying the first method. The relative amount of interspecific competition between these two species appears to be greater than that for other hermit crab species pairs studied previously.     

Frequencies of interspecific shell exchanges between hermit crabs

Frequencies of interspecific shell exchange due to shell fighting were determined for a number of species pairs of hermit crabs from several different locations. Frequencies were determined in the laboratory using a standardized experimental design. Results suggest that most individuals of most species are able to retain adequate or good quality shells in the presence of members of another species occupying poor quality shells. High frequencies of shell exchange always seem to be associated with very asymmetric relationships in which one member of the species pair is clearly dominant over the other. Dominant species usually attain larger sizes than subordinates, are found lower in the intertidal habitat, and are less abundant.     

Evolutionary palaeoecology of symbioses between bryozoans and hermit crabs

Modern hermit crabs form associations with many organisms which encrust, bore into, or cohabit the living chambers of gastropod shells occupied by the crabs. Among these hermit crab symbionts are bryozoan species which develop massive, commonly multilayered, colonies encrusting hermit crab shells. These colonies extend the living chamber of the crab through a characteristic process of helicospiral tubular growth originating from the shell aperture. The scant information available on the ecology of Recent bryozoan‐hermit crab symbioses is reviewed. Symbioses have been recorded from intertidal to upper slope environments, and from tropical to cold temperate zones. None of the hermit crab species are obligatory symbionts of bryozoans, and the majority of the modern bryozoan species involved are also not obligatory symbionts. Fossil examples always lack the hermit crabs, which have a poor fossilization potential; however, the distinctive tubular growth pattern and other features of the bryozoans enable recognition of ancient examples of the symbiosis. The earliest inferred associations between bryozoans and hermit crabs date from the Mid Jurassic, but associations remained uncommon until the Neogene. A remarkably wide taxonomic diversity of Recent and fossil bryozoans are known or inferred symbionts of hermit crabs. The broad evolutionary pattern of the association demonstrates multiple originations of the symbiosis by bryozoans belonging to at least 5 cyclostome and 12 cheilostome families. Only the Miocene‐Recent cheilostome family Hippoporidridae has an evolutionary history closely tied to symbiosis with hermit crabs. There is no evidence for coevolution.     

Vulnerability and reliable signaling in conflicts between hermit crabs

When interacting organisms have opposing genetic interests,the integrity of communication systems may be undermined. Forsignaling in such conflict circumstances to remain evolutionarilystable, cheaters must be handicapped. Agonistic threat signals,however, are not always constrained or costly to produce, andyet these signals occur in the severest of conflicts where strongincentives exist for dishonesty. A leading hypothesis for howreliability is stabilized under these conditions is that signalingentails a risk, making signalers vulnerable to injury. HereI experimentally alter vulnerability to show how risk can modifyorganisms' willingness to escalate disputes, affecting the useof threat signals. The vulnerability to injury of hermit crabs(Pagurus bernhardus) was manipulated by varying the exposureof their soft uncalcified abdomens. When faced with potentiallydamaging conspecific attacks, more vulnerable crabs were conflictaverse, showing reluctance to claim ownership over contestablefood, frequently retreating from threats, and refraining fromthreatening others. The risk an organism can bear in escalatedconflict can thus mediate its agonistic behavior and usage ofthreats. Postural nuances can consequently provide reliableinformation about aggressive intentions despite minimal productioncosts and opposing interests between communication parties.     

Ecological relations between hermit crabs and their shell-supplying gastropods: Constrained consumers

Hermit crabs are critically dependent upon gastropod shells for their survival and reproductive fitness. While anecdotal reports have suggested that hermit crabs may be capable of removing live gastropods from their shells to access the essential shell resource, no systematic experiments have been conducted to investigate this possibility. This paper reports experiments on both marine (Pagurus bernhardus) and terrestrial (Coenobita compressus) hermit crabs in which crabs were paired in the laboratory with the gastropods whose shells they inhabit in the field. Pairings included both shelled and naked crabs and spanned the full range of the gastropod life cycle. Neither marine nor terrestrial hermit crabs were successful at removing live gastropods from their shells. Furthermore, only a small fraction of the crabs (5.7%) were capable of accessing shells in which the gastropod had been killed in advance, with its body left intact inside the shell. Finally, although hermit crabs readily entered empty shells positioned on the surface, few crabs (14.3%) were able to access empty shells that were buried just centimeters beneath them. These results suggest that hermit crabs are constrained consumers, with the shells they seek only being accessible during a narrow time window, which begins following natural gastropod death and bodily decomposition and which typically ends when the gastropod's remnant shell has been buried by tidal forces. Further experiments are needed on more species of hermit crabs as well as fine-grained measurements of (i) the mechanical force required to pull a gastropod body from its shell and (ii) the maximum corresponding force that can be generated by different hermit crab species' chelipeds.     

红火蚁与热带火蚁间的干扰竞争

【目的】为了探讨入侵火蚁在我国成功定殖及其之间的竞争机制。【方法】运用行为学方法研究红火蚁Solenopsis invicta(Buren)和热带火蚁Solenopsis geminata(Fabricius)在个体水平和群体水平上的攻击性、攻击手段及合作能力。【结果】一对一攻击试验中,红火蚁和热带火蚁之间攻击级别多集中在3级,两种入侵蚂蚁间以相互威胁为主;红火蚁大型工蚁与热带火蚁兵、工蚁间最为好斗,其攻击级别达到4级的比例最高,分别为33.04%、37.92%。热带火蚁兵蚁与各型红火蚁间攻击强度差异不显著;热带火蚁工蚁与红火蚁小型工蚁之间的攻击性最强,其攻击性(3.49)显著高于热带火蚁工蚁与红火蚁大、中型工蚁的攻击性(3.32和2.97)。在攻击手段上,3级打斗时各型红火蚁更倾向以物理攻击主动威胁热带火蚁,而热带火蚁兵、工蚁会采取多种方式主动攻击红火蚁,双方皆以躲避应对为主;4级打斗时两种火蚁主要以混合攻击为主动或应对手段。群体攻击试验显示,红火蚁群体间攻击强度和合作性会随着群体数量的增加而显著增加,热带火蚁合作性较差,其群体对抗红火蚁的优势仅仅是由于个体数量的增加。【结论】红火蚁比热带火蚁具有更强的竞争优势。研究结果为入侵蚂蚁间不对称竞争机制和长期群落替代的内在原因提供理论基础。     

Interspecific competition between alien and native congeneric species

A good way to check hypotheses explaining the invasion of ecosystems by exotic plants is to compare alien and native congeneric species. To test the hypothesis that invasive alien plants are more competitive than natives, we designed a replacement series experiment to evaluate interspecific competition between three Senecio species representing the same bushy life form: two alien species (S. inaequidens and S. pterophorus, both from South Africa) and a native species from the south-east of the Iberian Peninsula and Maghreb (S. malacitanus). While S. inaequidens is widespread throughout western Europe and is expanding towards the south of Spanish–French border, the geographical distribution of the recently introduced S. pterophorus is still limited to north-eastern Spain. Plants from each species were grown in pure and in mixed cultures with one of their congeners, and water availability was manipulated to evaluate the effects of water stress on competitive abilities. Our results show that the alien S. inaequidens is the most competitive species for all water conditions. The native S. malacitanus is more competitive that the alien S. pterophorus in water stress conditions, but this situation is reversed when water availability is not limiting.     

Shell fighting and competition between two hermit crab species in Panama

Summary This article is a study of shell fighting between two intertidal hermit crab species in Panama. Laboratory results showed some cases of high exchange frequencies between Calcinus obscurus and Clibanarius albidigitus when the former occupied poor-quality shells. Exchange frequencies varied considerably between collecting sites, and were always low when the defending Clibanarius came from Venado Beach. Shell exchange frequencies estimated from field experiments were similar to those obtained in the laboratory. Observations on relative shell sizes occupied by both species in areas of sympatry and allopatry failed to provide clear evidence that Calcinus reduced the shell size of Clibanarius or that Clibanarius increased the shell size occupied by Calcinus. Results obtained here differ from those obtained in previous studies (Abrams 1980; Bertness 1981a, b), and these differences are discussed. Although shell fighting may be an important component of the interaction of these species, it is likely that roughly 90% of the competition experienced by each species is intraspecific.     

Interspecific competition and arbuscular mycorrhiza: Importance for the coexistence of two calcareous grassland species

Two naturally coexisting grassland species—Centaurea jacea (often predominating) andFragaria vesca (subordinate) were grown together in a pot experiment of factorial design, where competition and arbuscular mycorrhiza (AM) inoculation were used as treatments. The effects of competition were one-sided, i.e. the mass ofF. vesca decreased relatively more than that ofC. jacea as a result of competition. The root and total mass ofC. jacea increased with AM inoculation whileF. vesca did not respond. The mass difference betweenC. jacea andF. vesca in a particular pot increased when plants were mycorrhizal, mostly due to the mass increase ofC. jaceae. This can be explained by the differential response of these species to AM. We did not find any indirect indication of the interplant transfer of resources. The results of this experiment show the more unbalanced competition (larger differences in biomass) resulting from AM infection of plants. *** DIRECT SUPPORT *** A02DO006 00010     

Effect of gastropod shell characteristics and hermit crabs on shell Epifauna

The epifauna on gastropod shells occupied by the hermit crabs Pagurus pollicaris (Say) and P. longicarpus (Say) was examined, as was the utilization of shells by these two hermit crabs. In the study area in Tampa Bay, Florida, shells were not a limiting factor to the hermit crab population, and there apparently was little competition for shells. Interspecific competition for shells was limited because the two hermit crab species differed in size and hence occupied shells of different sizes. The total number and density of most epifaunal species were higher on shells occupied by hermit crabs than on unoccupied shells, possibly because hermit crabs prevent their shells from being buried and hence lengthen the time the epifaunal community can grow and develop. The hermit crab species also appeared to affect the epifaunal community, for the total number and density of most epifaunal species were larger on shells occupied by P. pollicaris than P. longicarpus. With increasing shell size, the populations of most epifaunal species, also were larger but not their density. Least influential in affecting the epifaunal community was the species of shells.