Marine Biology

, Volume 157, Issue 12, pp 2783–2789 | Cite as

Effects of crowding and wave exposure on penis morphology of the acorn barnacle, Semibalanus balanoides

  • J. Matthew HochEmail author
Original Paper


Wave action and low population density can strongly reduce the ability of sessile acorn barnacles to find mates and copulate. For Semibalanus balanoides, penis morphology varies with wave exposure and with characteristics of the mating neighborhood. Field experiments were conducted at five intertidal sites on Long Island, New York, USA from July to December 2005 to determine how wave exposure and aggregation structure influence the length, diameter, mass, and number of annulated folds of the penis. Sparsely crowded barnacles had more annulations in the penis and are inferred to have greater ability to stretch. At higher wave exposure, the diameter of the penis was greater, but the mass was not. This study identifies density of crowding as the most important cue that barnacles respond to when perceiving their mating group and details how penis morphology varies in response to wave exposure.


Jetty Mating Group Wave Exposure Test Volume Penis Length 
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Valuable ideas were incorporated into this manuscript deriving from conversations with J. Levinton, D. Padilla, D. Rand, R. Strathmann, J. True, and B. Yuen. This manuscript was meaningfully improved thanks to comments from the associate editor and anonymous reviewers. Help in the field was provided by B. Allen, P. Bourdeau, L. Brown, A. Ehmer, M. Fung, R. Junkins, C. McGlynn, B. Rodgers and E. Woo. Laboratory assistance was provided by E. O’Donnell, W. Wang and B. Yuen. Laboratory work was carried out in the Levinton Laboratory at Stony Brook University and the Functional Ecology Research and Training Lab (FERTL) in the department of Ecology and Evolution at Stony Brook University. Logistical support was provided by M. Doall. Assistance with statistical analysis was provided by Paulette Johnson at the FIU Statistical Consulting Center. Funding was provided by a Grant-in-aid-of-research from the Society for Integrative and Comparative Biology, a Student Research Fellowship from the Crustacean Society and a Doctoral Dissertation Improvement Grant from the National Science Foundation. An earlier version of this manuscript appears as a chapter in the author’s PhD dissertation.

Supplementary material

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Supplementary material 4 (DOC 73 kb)


  1. Arsenault D, Marchinko KB, Palmer AR (2001) Precise tuning of barnacle leg length to coastal wave action. P Roy Soc B-Biol Sci 268:2149–2154CrossRefGoogle Scholar
  2. Barnes M (1992) The reproductive periods and condition of the penis in several species of common cirripedes. Oceanogr Mar Biol 30:483–525Google Scholar
  3. Barnes H, Powell HT (1950) The development, general morphology, and subsequent elimination of barnacle populations after heavy initial settlement. J Anim Ecol 19:175–179CrossRefGoogle Scholar
  4. Bertness M, Gaines S, Yeh S (1998) Making mountains out of barnacles: the dynamics of acorn barnacle hummocking. Ecology 79:1382–1394CrossRefGoogle Scholar
  5. Chan BKK, Hung OS (2005) Cirral length of the acorn barnacle Tetraclita japonica (Cirripedia: Balanomorpha) in Hong Kong: effect of wave exposure and tidal height. J Crustacean Bio 25:329–332CrossRefGoogle Scholar
  6. Charnov EL (1980) Sex Allocation and local mate competition in barnacles. Mar Biol Lett 1:269–272Google Scholar
  7. Charnov EL (1982) The theory of sex allocation. Princeton University Press, PrincetonGoogle Scholar
  8. Crisp DJ, Patel BS (1958) Relation between brooding and ecdysis in cirripedes. Nature 181:1078–1079CrossRefGoogle Scholar
  9. Darwin C (1854) A monograph on the sub-class Cirripedia, with figures of all the species. The Balanidæ, (or Sessile Cirripedes); the Verrucidæ, etc. etc. etc. The Ray Society, LondonGoogle Scholar
  10. Gotelli NJ, Ellison AM (2004) A primer of ecological statistics. Sinauer, SunderlandGoogle Scholar
  11. Heath DJ (1977) Simultaneous hermaphroditism: cost and benefit. J Theor Biol 81:151–155CrossRefGoogle Scholar
  12. Hoch JM (2008) Variation in penis morphology and mating ability in the acorn barnacle, Semibalanus balanoides. J Exp Mar Biol Ecol 359:126–130CrossRefGoogle Scholar
  13. Hoch JM (2009) Adaptive plasticity of the penis in a simultaneous hermaphrodite. Evolution 63:1946–1953CrossRefPubMedGoogle Scholar
  14. Klepal W, Barnes H, Munn EA (1972) The morphology and histology of the cirripede penis. J Exp Mar Biol Ecol 10:243–265CrossRefGoogle Scholar
  15. Li N, Denny MW (2004) Limits to phenotypic plasticity: flow effects on barnacle feeding appendages. Biol Bull 206:121–124CrossRefPubMedGoogle Scholar
  16. Marchinko KB (2003) Dramatic phenotypic plasticity in barnacle legs (Balanus glandula Darwin): magnitude, age dependence and speed of response. Evolution 57:1281–1290PubMedGoogle Scholar
  17. Marchinko KB, Palmer AR (2003) Feeding in flow extremes: dependence of cirrus form on wave-exposure in four barnacle species. Zoology 106:127–141CrossRefPubMedGoogle Scholar
  18. Munn EA, Klepal W, Barnes H (1974) The fine structure and possible function of the sensory setae of the penis of Balanus balanoides (L.). J Exp Mar Biol Ecol 14:89–98CrossRefGoogle Scholar
  19. Neufeld C, Palmer AR (2008) Precisely proportioned: intertidal barnacles alter penis form to suit coastal wave action. P Roy Soc B-Biol Sci 275:1081–1087CrossRefGoogle Scholar
  20. Raimondi PT, Martin JE (1991) Evidence that mating group-size affects allocation of reproductive resources in a simultaneous hermaphrodite. Am Nat 138:1206–1217CrossRefGoogle Scholar
  21. Rasband WS (1997–2007) ImageJ. US National Institute of Health, Bethesda.
  22. Schärer L (2009) Tests of sex allocation theory in simultaneously hermaphroditic animals. Evolution 63:1377–1405CrossRefPubMedGoogle Scholar
  23. Sokal RR, Rohlf FJ (1995) Biometry, 3rd edn. W. H. Freeman and Company, New YorkGoogle Scholar
  24. Stubbings H (1975) Balanus balanoides. Liverpool University Press, LiverpoolGoogle Scholar
  25. Thompson TL, Glenn EP (1994) Plaster standards to measure water motion. Limnol Oceanogr 39:1768–1779CrossRefGoogle Scholar
  26. Vogel S (2003) Comparative biomechanics: life’s physical world. Princeton University Press, PrincetonGoogle Scholar
  27. Wethey DS (1984) Effects of crowding on fecundity in barnacles: Semibalanus (Balanus) balanoides, Balanus glandula and Chthamalus dalli. Can J Zool 62:1788–1795CrossRefGoogle Scholar
  28. Wu R, Levings C, Randall D (1977) Differences in energy partition between crowded and uncrowded barnacles (Balanus balanoides Darwin). Can J Zool 58:559–566CrossRefGoogle Scholar
  29. Yuen B, Hoch JM (2010) Factors influencing mating success in the acorn barnacle, Semibalanus balanoides. J Crustacean Biol 30:373–376CrossRefGoogle Scholar

Copyright information

© Springer-Verlag 2010

Authors and Affiliations

  1. 1.Department of Ecology and EvolutionStony Brook UniversityStony BrookUSA
  2. 2.Southeast Environmental Research CenterFlorida International UniversityMiamiUSA

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