Pagurus bernhardus

Geographic Range

Common hermit crabs are found in the near-shore North Atlantic waters of northwestern Europe, from the White Sea to the British Isles, including the North, Baltic, and Barents Seas. They are found as far south as Portugal, including in the Mediterranean Sea, as well as along the coasts of the Azorean Islands. ("ARKive", 2012; Anger, 1989; Reese, 1969; Reiss, et al., 2005; Sokolov, 2006; Tϋrkay, 2013)

Habitat

These crabs live along coasts in most types of seabeds, including rocky and shell bottoms, in sea grass beds, and sandy or silty sediments, but excluding muddy bottoms. Larvae live mainly in pools and may be found under objects such as rocks and seaweed. Common hermit crabs are most often found at depths up to 80 m, although they may be found as deep as 200 m. Smaller individuals live in shallower waters than larger individuals. (Burton and Burton, 2002; Lancaster, 1990; Tϋrkay, 2013)

  • Aquatic Biomes
  • temporary pools
  • coastal
  • Range depth
    50 to 200 m
    164.04 to 656.17 ft
  • Average depth
    80 m
    262.47 ft

Physical Description

Common hermit crabs are relatively large (maximum body length 8 cm), with bodies that are divided into two segments: cephalothorax and abdomen. The cephalothorax is encased by a carapace consisting of three thick cuticle layers: epicuticle, exocuticle, and endocuticle (maximum carapace length is 4.5 cm). The abdomen is soft and coiled to the right and body color is typically reddish or brown. These crabs inhabit the abandoned shells of animals, such as edible periwinkles (Littorina littorea), flat periwinkles (Littorina obtusata), dog whelks (Nucella lapillus) and other whelks (Buccinum sp.), using them as protection for their soft bodies. Their last two pairs of walking legs are greatly reduced, and are used to hold the shell in place. Compared to most hermit crabs, Common hermit crabs prefer a lighter shell. The size of the shell is important because it affects the fitness of the hermit crab-a shell that is too large does not offer the best protection and a shell that is too small restricts its growth. Individuals may attack each other in attempts to claim ownership of a shell. These crabs have 5 pairs of walking legs; the first pair are enlarged claws (known as chelae); those of males are larger than those of females. Chelae are used for gathering food and for protection. Of the two claws, one is larger and is covered by an operculum; this claw is used in fighting and defense. Chelae and walking leg surfaces are rough. Common hermit crabs have compound eyes and four short antennae. ("ARKive", 2012; "World Association of Zoos and Aquariums", 2012; Babu and Anger, 1987; Briffa, et al., 2008a; Briffa, et al., 2008b; Doake, et al., 2010; Elwood and Briffa, 2001; Reiss, et al., 2005; Tϋrkay, 2013)

  • Sexual Dimorphism
  • male larger
  • sexes shaped differently
  • Range mass
    0.13 to 54 g
    0.00 to 1.90 oz
  • Average mass
    7.3 g
    0.26 oz
  • Range length
    8 (high) cm
    3.15 (high) in

Development

Eggs of this species are black in color. Females carry eggs until they hatch (2 months, on average). Common hermit crabs pass through four zoeal stages and a glaucothoe/megalopa stage before reaching adulthood. Zoeal stages take 39-47 days to complete, with the megalopa stage requiring an additional 13 days on average; total development time averages 60 days. Zoeal larvae are thin, long, and yellowish-red in color. More chromatophores are added in each stage of zoeal development: megalops have areas of red and pale-yellow coloration and adults are reddish-brown. Megalops do not eat and must seek out a suitable shell in which to complete metamorphosis. Shell type and size effectively determine survival odds, as well as body size, time to sexual maturity, and longevity. Sexual differentiation occurs during the zoeal stages; female development is complete upon reaching adulthood, while males continue to develop morphological characteristics after adulthood. Sexual maturity is typically reached within a year. ("ARKive", 2012; Anger, 1989; Bookhout, 1964; Carvacho, 1988; Darwirs, 1979; Lancaster, 1990; Macdonald, 1957)

Reproduction

Unlike some species of crabs, copulation does not necessarily follow a female's molting. Before copulation, a male carries or pulls a female around by her shell opening for hours, even days in some cases, using his minor cheliped. Just prior to copulation, he holds her by one of her right walking legs, pulling her toward him and tapping her appendages with his major cheliped. This is followed by a period where both male and female tap each other with their chelipeds (the female also palpates the male's mouthparts) for 15-20 minutes. When a female is ready to mate, she gives either a tactile or chemical signal, causing a male to turn her around to face him. Both individuals leave their shells and copulate for 4-6 minutes. They remain together, outside of their shells, for up to 10 minutes after copulation. (Hazlett, 1970; Lancaster, 1988)

Unlike other Pagurus hermit crabs, Common hermit crabs are able to mate outside of the period after a female molts; there have been records of breeding occurring during all phases of a female molt cycle. There are many environmental factors that affect whether or not breeding occurs, such as photoperiod, water temperature (itself and in relation to air temperature), salinity, and availability of resources. Common hermit crabs may mate continuously throughout the year, although populations living in shallower water tend to have reproductive peaks in January and Feburary. After one encounter, a female produces 200-300 eggs, up to 30% of which may by lost or destroyed. ("ARKive", 2012; "World Association of Zoos and Aquariums", 2012; Burton and Burton, 2002; Hazlett, 1970; Lancaster, 1988)

  • Breeding interval
    Common hermit crabs breed multiple times throughout the year.
  • Breeding season
    Populations living in deeper waters breed year round, while those living in shallower waters tend to breed in January and February.
  • Range number of offspring
    200 to 300
  • Average gestation period
    43 days
  • Average age at sexual or reproductive maturity (female)
    1 years
  • Average age at sexual or reproductive maturity (male)
    1 years

After fertilization, a female carries eggs until they hatch (43 days, on average). During this time, they exhibit greater care in shell choice, switching shells more often than when they are not brooding. (Lancaster, 1988; Neil and Elwood, 1985)

Lifespan/Longevity

Common hermit crabs are known to live for up to four years in the wild. Shell selection, as well as molting, affects longevity. Molting, in particular, has short-term benefits and long-term costs; short term benefits include limb regeneration, while long-term costs include potential negative shifts in dominance hierarchy, reproductive success, feeding, communication, and locomotion. (Burton and Burton, 2002; Lancaster, 1988)

  • Range lifespan
    Status: wild
    4 (high) hours

Behavior

Common hermit crabs are nocturnal. They aggregate into loose communities, living around each other but not necessarily interacting. In these populations, breeding occurs at almost any time during the year and there is usually intense competition over resources, often resulting in physical damage to individuals. Each community has a dominant male; this individual wins the most fights and may assert its dominance by taking resources from others. (Briffa and Bibost, 2009; Briffa and Dallaway, 2007; Hazlett, 1968; Hazlett, 1970; Jackson and Elwood, 1989; Lancaster, 1988; Lancaster, 1990)

Before any physical encounter, Common hermit crabs engage in visual displays. During aggressive encounters, they either raise their bodies high off the ground, displaying their chelipeds and walking legs, or lower their bodies in submission. If neither individual submits, the crabs may try to dislodge each other from their shells, an interaction known as shell fighting. One hermit crab will crawl onto its opponent's shell, hitting its opponents entire body, including its shell, and using its chelipeds to rap its opponent's shell (over 500 raps have been observed in a single encounter). Around half of such encounters result in the aggressor exchanging shells with its opponent. Larger crabs, and those with larger chelipeds, are usually more successful than small- and medium-sized crabs in these encounters. (Briffa and Bibost, 2009; Briffa and Dallaway, 2007; Hazlett, 1968; Hazlett, 1970; Jackson and Elwood, 1989; Lancaster, 1988; Lancaster, 1990)

Another behavior that has been extensively studied is shell selection. Common hermit crabs are more likely to exchange and fight over shells when a new shell confers an advantage, such as more space. They are also able to remember shells that they have rejected before. If a crab is naked or in a shell that does not offer enough protection or space, then it will make the decision to move into a shell very quickly, regardless of advantages it may confer. If a new shell is of a much higher quality than the currently inhabited shell, the decision to move will also be made very quickly. (Briffa and Bibost, 2009; Briffa and Dallaway, 2007; Hazlett, 1968; Hazlett, 1970; Jackson and Elwood, 1989; Lancaster, 1988; Lancaster, 1990)

Home Range

Studies have shown that individuals may move within a range of 16-312 meters per month. (Bell, 2009)

Communication and Perception

Males and females communicate with each other through tactile and chemical signaling, and visual cues are used during aggressive encounters. (Briffa and Bibost, 2009; Briffa and Dallaway, 2007; Hazlett, 1968; Hazlett, 1970; Lancaster, 1988)

Food Habits

Common hermit crabs are scavengers. Their diet includes a wide variety of food sources, including microscopic bivalves, scraps of dead animals, and plants; they are known to be cannibalistic when resources are low. This diet is good for survival in environments that are unpredictable and can seasonally change. (Lancaster, 1990)

  • Animal Foods
  • mollusks
  • aquatic or marine worms
  • aquatic crustaceans
  • other marine invertebrates
  • Other Foods
  • microbes

Predation

Predators of common hermit crabs include fishes, starfish, octopi, other crabs, and some birds. Seagulls are known to pick up common hermit crabs in their beaks and drop them on rocks to break their shells. These hermit crabs may also exhibit cannibalism. (Haug, et al., 2002; Lancaster, 1988)

Ecosystem Roles

Common hermit crabs are valuable members of their ecosystems as scavengers and detritovores. As detritovores, these hermit crabs help decompose dead materials and contribute to nutrient cycles. Common hermit crabs may host a variety of epibionts on their shells, including protozoans, hydrozoans, entroprocts, barnacles, and polychaete worms. These hermit crabs prefer shells with epibonts over clean shells; some epibionts, such as anenomes, can protect the hermit crab from predators. They may also host parasites (most often isopods or barnacles), which bore into their shells, residing most often in the abdomen or brachial cavities, sometimes even causing castration in males. Infestation levels are as low as 1.5%. Eggs of this species have antibiotic properties. ("World Association of Zoos and Aquariums", 2012; Bell, 2009; Fernandez-Leborans and Gabilondo, 2006; Haug, et al., 2002; Lancaster, 1988)

Commensal/Parasitic Species

Economic Importance for Humans: Positive

Common hermit crabs play an important role in their environments as decomposers. They are also kept by some people in aquaria and may be used as fishing bait, once removed from their shells. (Lancaster, 1988)

Economic Importance for Humans: Negative

There are no known adverse effects of common hermit crabs on humans.

Conservation Status

This species has not yet been assessed for conservation by the IUCN or any other agency. There are no current conservation efforts specific to this species. (IUCN, 2013; Lancaster, 1988)

Contributors

Kathleen Wang (author), University of Michigan-Ann Arbor, Jeremy Wright (editor), University of Michigan-Ann Arbor.

Glossary

Atlantic Ocean

the body of water between Africa, Europe, the southern ocean (above 60 degrees south latitude), and the western hemisphere. It is the second largest ocean in the world after the Pacific Ocean.

World Map

Palearctic

living in the northern part of the Old World. In otherwords, Europe and Asia and northern Africa.

World Map

bilateral symmetry

having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.

biodegradation

helps break down and decompose dead plants and/or animals

carnivore

an animal that mainly eats meat

chemical

uses smells or other chemicals to communicate

coastal

the nearshore aquatic habitats near a coast, or shoreline.

detritivore

an animal that mainly eats decomposed plants and/or animals

dominance hierarchies

ranking system or pecking order among members of a long-term social group, where dominance status affects access to resources or mates

ectothermic

animals which must use heat acquired from the environment and behavioral adaptations to regulate body temperature

female parental care

parental care is carried out by females

fertilization

union of egg and spermatozoan

herbivore

An animal that eats mainly plants or parts of plants.

heterothermic

having a body temperature that fluctuates with that of the immediate environment; having no mechanism or a poorly developed mechanism for regulating internal body temperature.

indeterminate growth

Animals with indeterminate growth continue to grow throughout their lives.

internal fertilization

fertilization takes place within the female's body

iteroparous

offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).

macroalgae

seaweed. Algae that are large and photosynthetic.

metamorphosis

A large change in the shape or structure of an animal that happens as the animal grows. In insects, "incomplete metamorphosis" is when young animals are similar to adults and change gradually into the adult form, and "complete metamorphosis" is when there is a profound change between larval and adult forms. Butterflies have complete metamorphosis, grasshoppers have incomplete metamorphosis.

molluscivore

eats mollusks, members of Phylum Mollusca

motile

having the capacity to move from one place to another.

native range

the area in which the animal is naturally found, the region in which it is endemic.

nocturnal

active during the night

oceanic islands

islands that are not part of continental shelf areas, they are not, and have never been, connected to a continental land mass, most typically these are volcanic islands.

omnivore

an animal that mainly eats all kinds of things, including plants and animals

oviparous

reproduction in which eggs are released by the female; development of offspring occurs outside the mother's body.

pet trade

the business of buying and selling animals for people to keep in their homes as pets.

planktivore

an animal that mainly eats plankton

polygynandrous

the kind of polygamy in which a female pairs with several males, each of which also pairs with several different females.

saltwater or marine

mainly lives in oceans, seas, or other bodies of salt water.

seasonal breeding

breeding is confined to a particular season

sedentary

remains in the same area

sexual

reproduction that includes combining the genetic contribution of two individuals, a male and a female

social

associates with others of its species; forms social groups.

solitary

lives alone

tactile

uses touch to communicate

temperate

that region of the Earth between 23.5 degrees North and 60 degrees North (between the Tropic of Cancer and the Arctic Circle) and between 23.5 degrees South and 60 degrees South (between the Tropic of Capricorn and the Antarctic Circle).

territorial

defends an area within the home range, occupied by a single animals or group of animals of the same species and held through overt defense, display, or advertisement

visual

uses sight to communicate

year-round breeding

breeding takes place throughout the year

References

2012. "ARKive" (On-line). Common hermit crab (Pagurus bernhardus). Accessed January 21, 2012 at http://www.arkive.org/common-hermit-crab/pagurus-bernhardus/#text=All.

2012. "World Association of Zoos and Aquariums" (On-line). Common Hermit Crab (Pagurus bernhardus). Accessed July 31, 2013 at http://www.waza.org/en/zoo/choose-a-species/invertebrates/other-aquatic-invertebrates/pagurus-bernhardus.

Adema, H. 1980. The hermit crab (Pagurus bernhardus Linne 1785) and its parasites. Zeepaard, 40/2: 32-.

Anger, K. 1989. Growth and exuvial loss during larval and early juvenile development of the hermit crab Pagurus bernhardus reared in the laboratory. Marine Biology, 103/4: 503-511. Accessed March 25, 2012 at http://www.springerlink.com/content/n7331w1122606j2q/fulltext.pdf.

Ates, R. 2010. The unknown about the hermit crab, Pagurus bernhardus (Linnaeus, 1785). Strandvlo, 30/4: 102-.

Babu, D. 1988. 'Glandular pockets' of the integument and feeding mechanism in Pagurus bernhardus (Crustacea: Anomura). Marine Biology, 99/3: 315-323. Accessed July 30, 2013 at http://link.springer.com/article/10.1007/BF02112123#page-1.

Babu, D., K. Anger. 1987. The structure and modification of integumental tissues in Pagurus bernhardus (L.) (Decapoda: Anomura). Journal of Experimental Marine Biology and Ecology, 112/3: 267-281. Accessed February 03, 2012 at http://www.sciencedirect.com/science/article/pii/0022098187900736.

Bell, J. 2009. Hitching a ride on a hermit crabs home: Movement of gastropod shells inhabited by hermit crabs. Estaurine, Coastal, and Shelf Science, 85/2: 173-178. Accessed February 24, 2012 at http://www.sciencedirect.com/science/article/pii/S027277140900359X.

Bookhout, C. 1964. Salinity effects on the larval development of Pagurus bernhardus (L.) reared in the laboratory. Ophelia, 1/2: 275-294. Accessed March 25, 2012 at http://www.tandfonline.com/doi/pdf/10.1080/00785326.1964.10416283.

Briffa, M., M. Austin. 2009. Effects of predation threat on the structure and benefits from vacancy chains in the hermit crab Pagurus bernhardus. Ethology, 115/11: 1029-1035. Accessed January 21, 2012 at http://onlinelibrary.wiley.com/doi/10.1111/j.1439-0310.2009.01687.x/abstract.

Briffa, M., A. Bibost. 2009. Effects of shell size on behavioural consistency and flexibility in hermit crabs. Canadian Journal of Zoology-Revue Canadienne de Zoologie, 87/7: 597-603. Accessed January 21, 2012 at http://www.nrcresearchpress.com/doi/abs/10.1139/Z09-047?journalCode=cjz#.UffsKa6veRk.

Briffa, M., D. Dallaway. 2007. Inter-sexual contests in the hermit crab Pagurus bernhardus: females fight harder but males win more encounters. Behavioral Ecology and Sociobiology, 61/11: 1781-1787. Accessed February 03, 2012 at http://www.jstor.org/stable/27823563.

Briffa, M., P. Haskell, C. Wilding. 2008. Behavioural colour change in the hermit crab Pagurus bernhardus: reduced crypticity when the threat of predation is high. Behaviour, 145/7: 915-929. Accessed February 03, 2012 at http://www.ingentaconnect.com/content/brill/beh/2008/00000145/00000007/art00003.

Briffa, M., S. Rundle, A. Fryer. 2008. Comparing the strength of behavioural plasticity and consistency across situations: animal personalities in the hermit crab Pagurus bernhardus. Proceedings of the Royal Society Biological Sciences Series B, 275/1640: 1305-1311. Accessed July 30, 2013 at http://rspb.royalsocietypublishing.org/content/275/1640/1305.full.pdf+html.

Briffa, M., C. Twyman. 2006. Do I stand out of blend in? Conspicuousness awareness and consistent behavioural differences in hermit crabs. Biology Letters, 7/3: 330-332. Accessed January 21, 2012 at http://rsbl.royalsocietypublishing.org.proxy.lib.umich.edu/content/7/3/330.

Briffa, M., R. Williams. 2006. Use of chemical cues during shell fights in the hermit crab Pagurus bernhardus. Behaviour, 143/10: 1281-1290. Accessed February 03, 2012 at http://www.jstor.org/stable/4536406.

Burton, M., R. Burton. 2002. The International Wildlife Encyclopedia, Volume 1. New York: Marshall Cavendish Corporation.

Carvacho, A. 1988. Early development of Pagurus bernhardus L. (Crustacea, Decapoda). Cahiers de Biologie Marine, 29/1: 109-133. Accessed March 25, 2012 at http://www.sb-roscoff.fr/cbm/article.htm.

Darwirs, R. 1979. Effects of Temperature and Salinity on Larval Development of Pagurus bernhardus (Decapoda, Paguridae). Marine Ecology Progress Series, 1: 323-329. Accessed March 25, 2012 at http://www.int-res.com/articles/meps/1/m001p323.pdf.

Doake, S., R. Elwood. 2011. How resource quality differentially affects motivation and ability to fight in hermit crabs. Proceedings of the Royal Society B-Biological Sciences, 278/1705: 567-573. Accessed January 21, 2012 at http://rspb.royalsocietypublishing.org.proxy.lib.umich.edu/content/278/1705/567.

Doake, S., M. Scantlebury, R. Elwood. 2010. The costs of bearing arms and armour in the hermit crab Pagurus bernhardus. Animal Behavior, 80/4: 637-642. Accessed January 21, 2012 at http://www.sciencedirect.com/science/article/pii/S0003347210002617.

Elwood, R., M. Briffa. 2001. Information gathering and communication during agonistic encounters: A case study of hermit crabs. Advances in the Study of Behavior, 30: 53-97. Accessed February 24, 2012 at http://www.sciencedirect.com/science/article/pii/S006534540180005X.

Elwood, R., N. Marks, J. Dick. 1995. Consequences of shell-species preference for female reproductive success in the hermit crab Pagurus bernhardus. Marine Biology, 123/3: 431-434. Accessed July 30, 2013 at http://link.springer.com/article/10.1007/BF00349221#page-1.

Elwood, R., A. Stewart. 1987. Reproduction in the littoral hermit crab Pagurus bernhardus. Irish Naturalists' Journal, 22/6: 252-255. Accessed July 30, 2013 at http://www.jstor.org/stable/25539155.

Fernandez-Leborans, G., R. Gabilondo. 2006. Taxonomy and distribution of the hydrozoan and protozoan epibionts on Pagurus bernhardus (Linaeus, 1785) (Crustacea, Decapoda) from Scotland. Acta Zoologica, 87/1: 33-48. Accessed February 03, 2012 at http://onlinelibrary.wiley.com/doi/10.1111/j.1463-6395.2006.00218.x/abstract.

Gherardi, F. 2010. Chemical Communication in Crustaceans. New York, NY: Springer.

Haug, T., A. Kjuul, K. Stensvag, E. Sandsdalen, O. Styrvold. 2002. Antibacterial activity in four marine crustacean decapods. Fish and Shellfish Immunology, 12/5: 371-385. Accessed March 26, 2012 at http://www.sciencedirect.com/science/article/pii/S105046480190378X.

Hazlett, B. 1968. Communicatory effect of body position in Pagurus bernhardus (L.) (Decapoda, Anomura). Crustaceana, 14/2: 210-214. Accessed March 26, 2012 at http://www.jstor.org/stable/pdfplus/20102937.pdf?acceptTC=true.

Hazlett, B. 1982. Resource value and communication strategy in the hermit crab Pagurus bernhardus (L.). Animal Behavior, 30/1: 135-139. Accessed February 03, 2012 at http://www.sciencedirect.com/science/article/pii/S0003347282802480.

Hazlett, B. 1970. Tactile Stimuli in the Social Behavior of Pagurus bernhardus (Decapoda, Paguridae). Behaviour, 36/1-2: 20-48. Accessed March 30, 2012 at http://www.jstor.org/stable/pdfplus/4533318.pdf?acceptTC=true&.

IUCN, 2013. "IUCN Red List of Threatened Species. Version 2013.1" (On-line). Accessed March 26, 2012 at iucnredlist.org.

Jackson, N., R. Elwood. 1989. How animals make assessments: information gathering by the hermit crab Pagurus bernhardus. Animal Behaviour, 38/6: 951-957. Accessed March 30, 2012 at http://www.sciencedirect.com/science/article/pii/S0003347289801368.

Kaiser, M., K. Ramsay, R. Hughes. 1998. Can fisheries influence interspecific competition in sympatric populations of hermit crabs?. Journal of Natural History, 32/4: 521-531. Accessed February 03, 2012 at http://www.tandfonline.com/doi/abs/10.1080/00222939800770281#.Ufkv_K6veRk.

Knickmeyer, R., H. Steinhart. 1988. Seasonal differences of cyclic organochlorines in eggs of the hermit crab Pagurus bernhardus L. from the North Sea. Sarsia, 73/4: 291-298. Accessed July 31, 2013 at http://www.tandfonline.com/doi/abs/10.1080/00364827.1988.10413416#.UfkwOK6veRk.

Laidre, M. 2011. Ecological relations between hermit crabs and their shell-supplying gastropods: Constrained consumers. Journal of Experimental Marine Biology and Ecology, 397/1: 65-70. Accessed January 21, 2012 at http://www.sciencedirect.com/science/article/pii/S0022098110004296.

Lancaster, I. 1988. Pagurus Bernhardus (L.)-An Introduction to the Natural History of Hermit Crabs. Field Studies, 7: 189-238. Accessed February 24, 2012 at http://www.eco-challenge-xtra.org/fieldstudies/documents/vol7.1_192.pdf.

Lancaster, I. 1990. Reproduction and life history strategy of the hermit crab Pagurus bernhardus. Journal of the Marine Biological Association of the United Kingdom, 70/1: 129-142. Accessed July 31, 2013 at http://journals.cambridge.org/action/displayAbstract?fromPage=online&aid=4366412.

Lancaster, I., G. Wigham. 1990. Influences on movement and migration in the hermit crab Pagurus bernhardus. Journal of the Marine Biological Association of the United Kingdom, 70/3: 483-491. Accessed July 31, 2013 at http://journals.cambridge.org/action/displayAbstract;jsessionid=0860964A8410A567470D2B9506D75B18.journals?fromPage=online&aid=4370580.

Macdonald, J. 1957. Larvae of the British Species of Diogenes, Pagurus, Anapagurus and Lithodes (Crustacea, Decapoda). Proceedings of the Zoological Society of London, 128/2: 209-258. Accessed March 25, 2012 at http://onlinelibrary.wiley.com/doi/10.1111/j.1096-3642.1957.tb00265.x/pdf.

Matzen da Silva, J., A. dos Santos, M. Cunha, F. Costa, S. Creer, G. Carvalho. 2011. Multigene Molecular Systematics Confirm Species Status of Morphologically Convergent Pagurus Hermit Crabs. PLOS ONE, 6/12. Accessed January 21, 2012 at http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0028233.

Mowles, S., P. Cotton, M. Briffa. 2010. Whole-organisms performance capacity predicts resource-holding potential in the hermit crab Pagurus bernhardus. Animal Behaviour, 80/2: 277-282. Accessed February 03, 2012 at http://bama.ua.edu/~rlearley/Mowles_2010.pdf.

Neil, S., R. Elwood. 1985. Behavioural modifications during egg-brooding in the hermit crab, Pagurus bernhardus L.. Journal of Experimental Marine Biology and Ecology, 94/1-3: 99-114. Accessed February 03, 2012 at http://www.sciencedirect.com/science/article/pii/0022098185900528.

Perkins, E. 1984. Some aspects of the biology of Pagurus bernhardus (L.). Transactions of the Dumfriesshire and Galloway Natural History and Antiquarian Society Third Series, 59: 21-26. Accessed July 31, 2013 at http://www.dgnhas.org.uk/transonline/SerIII-Vol59.pdf.

Ramsay, K., M. Kaiser, R. Hughes. 1996. Changes in hermit crab feeding patterns in response to trawling disturbance. Marine Ecology Progress Series, 144: 63-72. Accessed February 03, 2012 at http://www.int-res.com.proxy.lib.umich.edu/articles/meps/144/m144p063.pdf.

Reese, E. 1969. Behavioral adaptations of intertidal hermit crabs. American Zoologist, 9/2: 343-355. Accessed February 24, 2012 at http://www.jstor.org/stable/pdfplus/3881807.pdf?acceptTC=true.

Reiss, H., H. Neumann, I. Kröncke. 2005. Chela-height vs. body-weight relationships for North Sea hermit crabs (Paguridae). Ices Journal of Marine Science, 62/4: 723-726. Accessed February 24, 2012 at http://icesjms.oxfordjournals.org/content/62/4/723.full.

Sokolov, V. 2006. Distribution of two hermit crabs Pagurus pubescens and P. bernhardus (Anomura, Paguridae) in the Barents and White Seas. Zoologiceskij zurnal, 85/10: 1176-1186. Accessed February 24, 2012 at http://cat.inist.fr/?aModele=afficheN&cpsidt=18516115.

Tuerkay, M. 2006. The hermit crab Pagurus bernhardus. Natur und Museum (Frankfurt am Main), 136/3-4: 62.

Tϋrkay, M. 2013. "Pagurus bernhardus (Linnaeus, 1758)" (On-line). WoRMS: World Register of Marine Species. Accessed January 21, 2012 at http://www.marinespecies.org/aphia.php?p=taxdetails&id=107232.

de la Haye, K., J. Spicer, S. Widdicombe, M. Briffa. 2011. Reduced sea water pH disrupts resource assessment and decision making in the hermit crab Pagurus bernhardus. Animal Behavior, 82/3: 495-501. Accessed January 21, 2012 at http://www.sciencedirect.com/science/article/pii/S0003347211002405.