The Relationship Between Pagurus Anachoretus and Cerithium
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The relationship between Pagurus anachoretus and Cerithium vulgatum as affected by shell size and availability at Station de Recherches Sous-Marines et Oceanographiques in Corsica, France. Wade Dugdale, Ryan Baker, Eleni Christoforou University of California at Santa Cruz - BIO159 - Corsica 2014 Abstract The paper expands on the hermit crab species Pagurus anachoretus and the gastropod shell Cerithium vulgatum that it inhabits. The association between claw size of P. anachoretus and C. vulgatum shell and aperture size were tested on specimens collected in situ and measured in a lab. Contrary to current literature, we found no correlation between shell and crab size with crabs from the wild. We also ran experiments in a confined tank space to test whether P. anachoretus will exchange shells for a more optimum fit, when given the opportunity. Hermit crabs, left in a tank overnight, moved into more preferable shells after empty shells were added. They arranged themselves into even more preferred shells, suggesting that they do re-sort based on shell size preference. Our results suggest that this movement occurs to improve the hermit crabs’ general fitness. Introduction 1976; Bertness 1980), growth rate (Bertness 1980), survival and fecundity (Angel 2000). Hermit crabs, have interested biologists and As hermit crabs grow, they must find larger naturalists for centuries, dating back to shells, therefore they are frequently in the Aristotle, in 350 B. C, (translated in English search for a new shell. Hermit crabs most by D’Arcy Wentworth Thompson in 1910) often obtain shells in one of two ways. who was the first to study their nature Either directly after the snail’s death (Reese 1962). Hermit crabs (Paruroidea) following its decomposition, after the shell belong to the class called Malacostraca is deserted on the bottom’s substrate (Laidre (Aristotle 350 B. C. E), which is derived 2011) or by competition with other hermit from the Greek words ‘µαλακό’ and crabs, while most of the times both invader ‘όστρακο’ meaning soft shell. All and defender should have a benefit for the organisms’ fitness depends on resources exchange to occur (Hazlett 1981; Briffa and provided by other taxa. Hermit crabs are not Elwood 2000). A factor influening hermit an exception (Laidre 2011), relying on crabs’ fitness is that empty gastropod shells empty gastropod shells or in limited cases, are not easily found in most habitats other types of cavities (Hazlett 1981). (Childress 1972). This makes the study of Gastropod shells are of a great importance to hermit crabs and their relation to shells hermit crabs fitness, providing protection essential to ecology (Arce 2012) in order to from predation (Vance 1972). The main understand the severity of shell limitation in aspects of shells that are of interest to hermit nature (Kellogg 1976). The importance of crabs are their shape and size which hermit crabs to the environment is also influence the crab’s reproduction (Kellogg ! 1! suggested by the analysis of the 550 optimum shell in order to improve its invertebrates that live harmonically and are fitness. This prediction was supported by the dependent on hermit crabs (Williams and statement that a hermit crab that has a large McDermott 2004). Despite the limitation in choice of shells will decide on a shell closer gastropod shells, not all shells are to the optimum size, compared to a hermit appropriate (Childress 1972) for the hermit crab that has a limited amount of shells crabs, who choose shells based on their size, available (Hazlett 1992). With that said, the species (Bertness 1980; Reese 1962) i.e. following methods and results expand on shape and abundance (Reese 1962; Reese whether there is a relationship between the 1969; Bertness 1980). size of P. anachoretus and C. vulgatum and if P. anachoretus change into more optimum After observing hermit crabs at the Station shells when provided with the opportunity to de Recherches Sous-Marines et resort with other individuals in a confined Oceanographiques (STARESO), in Corsica, tank space. France, we became interested in their shell choice patterns. Therefore, our main goal was to observe, study, test and understand the relationship between the hermit crabs Materials and Methods and the shell they inhabit, in the specific Location area. We first observed that P. anachoretus has a strong preference for Cerithium The study was contacted South of vulgatum shells, which was the most STARESO field station harbor, which is abundant gastropod shell species. Hazlett located on the northwest coast of Corsica, (1981; 1992) found a strong correlation France. Fieldwork was conducted over large between the size of the shell and the size of (1-5m) sub-tidal boulders at a depth of 4- hermit crabs, and therefore we decided to 10m. The surfaces of these boulders were test it ourselves. Following the collection of primarily covered with algae and detritus. C. vulgatum shells inhabited by P. The area has very little daily tidal anachoretus, our first hypothesis was that disturbance but is affected by major storm there is a relationship between the shell size events annually. This study was conducted and the hermit crab size of these specific in October 2014 before the first major storm species in the local area. Additional event of the season. information was that, crabs in smaller than preferred shells experience a slower growth rate (Bertness 1981) and are more exposed to predation (Angel 2000; Hazlett 1981) because of their lack of space to withdraw and protect themselves. On the other hand, crabs in larger shells must carry more mass, leading to greater energy expenditure (Arce 2011). In a broader scale, a non-adequate shell negatively influences a crab’s fecundity (Vance 1972) and reduces its chance of survival (Childress 1972). Figure 1: Left: West Mediterranean with a Therefore our second hypothesis was that, in square around Corsica. Right: Corsica with a a high-density environment, P. anachoretus square around where STARESO is. (images would compete and change into a more produced by using Google maps) ! 2! Species from our study in order to prevent misrepresentation of the shell metrics. Pagurus anachoretus is a marine decapod from the family Paguridae. This sub-tidal species ranges from 1-40 meter depth and is found throughout the Mediterranean sea. Pagurus anachoretus is omnivorous, preying primarily on tiny marine animals and opportunistically scavenging on carrion. In the study area P. anachoretus was observed to be solitary while other species of hermit crabs are known to cluster. Additionally, P. anachoretus lay and carry their eggs within the shell they inhabit. Collections We collected P. anachoretus individuals inhabiting C. vulgatum shells by hand using SCUBA. The collections were conducted at night given the fact that P. anachoretus are nocturnal, when the animals are more active. We collected specimens primarily from the Figure 2: The illustrations represent the tops and sides of boulders and placed them standardized linear metrics that were into sealable bags filled with seawater until initially used to measure P. anachoretus our return to the laboratory. At that time we chelae and C. vulgatum shells: chelae length measured the chelae and associated shells of (CL) and width (CW), shell length (SL), each specimen, thus minimizing the time shell width (SW) as well as the aperture availability, to ensure that no shell length (AP) and aperture width (AW) of exchanges occurred before baseline each C. vulgatum shell. measurements. All of our measurements were taken by one Mass to Chelae Relationship technician in order to standardize the measurement data, using vernier calipers Twenty-three previously collected P. and rounding to the closest 0.5mm. We anachoretus specimens were removed from measured two linear metrics of crab size: their shells and we measured their chelae chelae length (CL) and chelae width (CW), length and width. Their mass was also to use as proxies for overall crab size measured to a thousandth of a gram. We without having to remove individuals from then ran a linear regression analysis of their shells. We measured four linear metrics bivariate fits of chelae length by crab mass of shell size: shell length (SL), shell width in order to determine whether chelae size (SW), aperture length (AL) and aperture was a viable representation of overall crab width (AW) of each shell (Figure 1). All size. This allowed us to measure the relative specimens with broken shells were excluded sizes of hermit crabs in subsequent analyses ! 3! and experiments without removing them All but 3 of the empty shells used in this from their shells. experiment were previously inhabited; we removed their inhabitants by anesthetizing them using clove oil. The reason it was Shell Crab to Shell Size Relationship necessary to use these shells was because of the lack of suitable empty shells found in the We performed a scatterplot matrix analysis local area. In the 36 hours in between the of all possible pairs between the two linear first and second trial the hermit crabs were chelae size metrics and four linear shell size kept in the same tank with constant water metrics to determine which pair of metrics flow and aeration. Small rocks and algae had the strongest correlation between crab like Padina pavonica were added to the tank and shell size. These two metrics would be for the hermit crabs to feed on. used as proxies for crab and shell size for all subsequent analyses and experiments. We ran a linear regression analysis of the best Results pair of size metrics using JMP (Statistical Discovery software for SAS) to determine Mass to Chelae relationship whether a significant relationship between chelae metrics and shell metrics existed in A regression analysis of the bivariate fit of wild hermit crabs. CL by crab mass (n=23) revealed a significant positive linear correlation between the chelae length and the mass of Shell Exchange – Trial 1 hermit crab individuals (Figure 3.