Marine Biology

, Volume 153, Issue 4, pp 523–528 | Cite as

Claw allometry in green crabs, Carcinus maenas: heterochely, handedness, and sex

  • F. JuanesEmail author
  • K. T. Lee
  • A. McKnight
  • K. Kellogg
Research Article


Claw loss and reversal of handedness during regeneration are common phenomena in heterochelous decapod crustaceans, which typically have one large ‘crusher’ claw on the right side and a smaller ‘cutter’ claw on the left. Little is known about the relative importance of claw growth vs. body growth during claw regeneration. Here the relationship between claw size and body size of green crabs, Carcinus maenas, was examined to test for differences in claw allometry as a function of handedness and sex, as there are differences in how males and females use their claws. A total of 730 crabs (range = 15.7–83.6 mm CW) were collected from Maine to New Jersey, USA from May to October 1997, 2000, and 2004–2005. Claw growth, particularly crushers, was accelerated in left-handed crabs and in males compared to right-handed crabs and females respectively. These differing growth strategies highlight the role of sexual dimorphism in claw usage and the importance of achieving heterochely after claw injury. These results imply that handedness should be an important factor to consider in future studies of crab morphology, behavior, and morphometrics.


Carapace Width Body Growth Green Crab Male Crab Stone Crab 
These keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.



This project was funded in part by the Richard Cronin Fisheries Research Fund and the Woods Hole Scholarship Fund from the University of Massachusetts, by a Cooperative State Research, Extension, and Education Service, U.S. Department of Agriculture, MA Agricultural Experiment Station Hatch grant, and by a University of Pittsburgh at Johnstown Research Council Small Grant. We thank Connecticut College for use of space and equipment, Normandeau Associates for help with crab collections, and Rutgers University Marine Field Station, Suffolk University Friedman Field Station, and Wells National Estuarine Research Reserve for access to lab space and field sites. Writing was completed while the senior author was a Center Fellow at the National Center for Ecological Analysis and Synthesis, a Center funded by NSF (Grant #DEB-0553768), the University of California, Santa Barbara, and the State of California. We thank LD Smith for a thorough review of the manuscript and his insightful comments, as well as three anonymous reviewers, all of whom helped in producing a better manuscript.

Supplementary material

227_2007_826_MOESM1_ESM.doc (29 kb)
Claw allometry in green crabs, Carcinus maenas: heterochely, handedness, and sex


  1. Abby-Kalio NJ, Warner GF (1989) Heterochely and handedness in the shore crab Carcinus maenas (L.) (Crustacea: Brachyura). Zool J Linn Soc 96:19–26CrossRefGoogle Scholar
  2. Abelló P, Pertierra JR, Reid DG (1990) Sexual dimorphism, relative growth and handedness in Liocarcinus depurator and Macropipus tuberculatus (Brachyura: Portunidae). Sci Mar 54:195–202Google Scholar
  3. Abelló P, Warman CG, Reid DG, Naylor E (1994) Chela loss in the shore crab Carcinus maenas (Crustacea: Brachyura) and its effect on mating success. Mar Biol 121:247–252CrossRefGoogle Scholar
  4. Backwell PRY, Christy JH, Telford SR, Jennions MD, Passmore NI (2000) Dishonest signalling in a fiddler crab. Proc R Soc Lond B 267:719–724CrossRefGoogle Scholar
  5. Brock RA, Smith DL (1998) Recovery of claw size and function following autotomy in Cancer productus (Decapoda: Brachyura). Biol Bull 194:53–62CrossRefGoogle Scholar
  6. Cheung TS (1976) A biostatistical study of the functional consistency in the reversed claws of the adult male stone crab, Menippe mercenaria (Say). Crustaceana 31:137–144CrossRefGoogle Scholar
  7. Crothers JH (1967) The biology of the shore crab Carcinus maenas (L.). I. The background—anatomy, growth and life history. Field Stud 2:407–434Google Scholar
  8. Daniels SR (2001) Allometric growth, handedness, and morphological variation in Potamonautes warreni (Calman, 1918) (Decapoda, Brachyura, Potamonautidae) with a redescription of the species. Crustaceana 74:237–253CrossRefGoogle Scholar
  9. Elner RW (1980) The influence of temperature, sex and chela size in the foraging strategy of the shore crab, Carcinus maenas (L.). Mar Behav Physiol 7:15–24CrossRefGoogle Scholar
  10. Figiel CR, Miller GL (1995) The frequency of chelae autotomy and its influence on the growth and survival of crayfish Procambarus clarkii (Girard, 1852) (Decapoda, Cambaridae). Crustaceana 68:472–483CrossRefGoogle Scholar
  11. Govind CK, Blundon JA (1985) Form and function of the asymmetric chelae in blue crabs with normal and reversed handedness. Biol Bull 168:321–331CrossRefGoogle Scholar
  12. Hamilton PV, Nishimoto RT, Halusky JG (1976) Cheliped laterality in Callinectes sapidus (Crustacea: Portunidae). Biol Bull 150:393–401CrossRefGoogle Scholar
  13. Hartnoll RG (1982) Growth. In: Abele LG (ed) The biology of Crustacea. Embryology, morphology, and genetics, vol 2. Academic, NY, pp 111–119Google Scholar
  14. Jivoff P (1997) Sexual competition among male blue crab, Callinectes sapidus. Biol Bull 193:368–380CrossRefGoogle Scholar
  15. Juanes F, Hartwick EB (1990) Prey size selection in Dungeness crabs: the effect of claw damage. Ecology 71:744–758CrossRefGoogle Scholar
  16. Juanes F, Smith LD (1995) The ecological consequences of limb loss and damage in decapod crustaceans: a review and prospectus. J Exp Mar Biol Ecol 193:197–223CrossRefGoogle Scholar
  17. Ladle RJ, Todd PA (2006) A developmental model for predicting handedness frequencies in crabs. Acta Oecologica 30:283–287CrossRefGoogle Scholar
  18. Lee SY (1993) Chela height is an acceptable indicator of chela strength in Carcinus maenas (Linnaeus, 1758) (Decapoda: Brachyura). Crustaceana 65:115–116CrossRefGoogle Scholar
  19. Lee SY (1995) Cheliped size and structure: the evolution of a multi-functional decapod organ. J Exp Mar Biol Ecol 193:161–176CrossRefGoogle Scholar
  20. Lee SY, Seed R (1992) Ecological implications of cheliped size in crabs: some data from Carcinus maenas and Liocarcinus holsatus. Mar Ecol Prog Ser 84:151–160CrossRefGoogle Scholar
  21. Mariappan P, Balasundaram C, Schmitz B (2000) Decapod crustacean chelipeds: an overview. J Biosci 25:301–313CrossRefGoogle Scholar
  22. Mathews LM, McKnight AE, Avery R, Lee KT (1999) Incidence of autotomy in New England populations of green crabs, Carcinus maenas, and an examination of the effect of claw autotomy on diet. J Crust Biol 19:713–719CrossRefGoogle Scholar
  23. Mitchell SC, Kennedy SM, Williams PJ, DeMont ME (2003) Morphometrics and estimates of force generation by the chelae of a North American population of the invasive green crab, Carcinus maenas (L.). Can J Zool 81:203–215CrossRefGoogle Scholar
  24. Ng PKL, Tan LWH (1985) ‘Right handedness’ in heterochelous calappoid and xanthoid crabs—suggestion for a functional advantage. Crustaceana 49:98–100CrossRefGoogle Scholar
  25. Przibram H (1931) Connecting laws in animal morphology. University of London Press, LondonGoogle Scholar
  26. Reid DG, Abelló P, Warman CG, Naylor E (1994) Size-related mating success in the shore crab Carcinus maenas (Crustacea: Brachyura). J Zool 232:397–407CrossRefGoogle Scholar
  27. Rosenberg MS (2002) Fiddler crab claw shape variation: a geometric morphometric analysis across the genus Uca (Crustacea: Brachyura: Ocypodidae). Biol J Linn Soc 75:147–162Google Scholar
  28. Savage T, Sullivan JR (1978) Growth and regeneration of the stone crab, Menippe mercenaria. Fla Mar Res Publ 32:1–23Google Scholar
  29. Schenk SC, Wainwright PC (2001) Dimorphism and the functional basis of claw strength in six brachyuran crabs. J Zool 255:105–119CrossRefGoogle Scholar
  30. Shigemiya Y (2003) Does the handedness of the pebble crab Eriphia smithii influence its attack success on two dextral snail species? J Zool 260:259–265CrossRefGoogle Scholar
  31. Simonson JL (1985) Reversal of handedness, growth, and claw stridulatory patterns in the stone crab Menippe mercenaria (Say) (Crustacea: Xanthidae). J Crust Biol 5:281–293CrossRefGoogle Scholar
  32. Simonson JL, Steele P (1981) Cheliped asymmetry in the stone crab, Menippe mercenaria, with notes on claw reversal and regeneration. North Gulf Sci 5:21–30Google Scholar
  33. Skinner DM (1985) Molting and regeneration. In: Bliss DE, Mantel LH (eds) Biology of Crustacea, vol 9. Academic, New York, pp 43–146Google Scholar
  34. Smith LD (2004) Biogeographic differences in claw size and performance in an introduced crab predator Carcinus maenas. Mar Ecol Prog Ser 276:209–222CrossRefGoogle Scholar
  35. Smith LD, Hines AH (1991) The effect of cheliped loss on blue crab Callinectes sapidus Rathbun foraging rate on soft-shelled clams Mya arenaria L. J Exp Mar Biol Ecol 151:245–256CrossRefGoogle Scholar
  36. Sneddon LU, Huntingford FA, Taylor AC (1997) Weapon size versus body size as a predictor of winning in fights between shore crabs, Carcinus maenas (L.). Behav Ecol Sociobiol 41:237–242CrossRefGoogle Scholar
  37. Vermeij GJ (1977) Patterns in crab claw size: the geography of crushing. Syst Zool 26:138–151CrossRefGoogle Scholar

Copyright information

© Springer-Verlag 2007

Authors and Affiliations

  • F. Juanes
    • 1
    Email author
  • K. T. Lee
    • 2
  • A. McKnight
    • 1
  • K. Kellogg
    • 3
  1. 1.Department of Natural Resources ConservationUniversity of MassachusettsAmherstUSA
  2. 2.Department of BiologyUniversity of Pittsburgh at JohnstownJohnstownUSA
  3. 3.Environmental Studies Program, Skidmore CollegeSaratoga SpringsUSA

Personalised recommendations