Tracks and Trails

  • Carl N. ShusterJr.


In summary, in sleuthing the trails of horseshoe crabs, whether observing spawning or in interpreting the trails they have left behind, a camera is a prerequisite for a complete record. No spawning group or any trail appear to be exactly alike another although there may be several similarities. That is what makes the deciphering so interesting; even when watching spawning or a trail being made. Afterwards, with only the trail remaining, much has to be imagined. Since the trails are so transitory, a good photograph is needed to refresh ones memory.

At least two main considerations must be made when looking at a trail. The characteristics and condition of the substrate and the various evidence of activity left by the crab. It is the attempt to unravel of all the possible combinations of conditions and activity that make the study of these prints so fascinating.


Sandy Beach Horseshoe Crab Migratory Shorebird Male Mating Tactic Ebbing Tide 
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.


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  1. Allee, Warder C. (1922). Studies in marine ecology II: An annotated catalog of the distribution of common Invertebrates of the Woods Hole littoral. Manuscript in the Marine Biological Library, Woods Hole, Massachusetts.Google Scholar
  2. Barthel, K. Werner. (1974). Limulus: a living fossil. Die Naturwissenschaften 61: 428–433.CrossRefGoogle Scholar
  3. Botton, Mark L. (1981). The gill books of the horseshoe crab Limulus polyphemus) as a substrate for the blue mussel (Mytilus edulis). Bulletin New Jersey Academy of Science 26: 26–28.Google Scholar
  4. Botton, Mark L. and Robert E. Loveland. (1987). Orientation of the horseshoe crab, Limu!us po!yphemus, on a sandy beach. Biological Bulletin 173: 289–298. 1989. Reproductive risk: high mortally associated with spawning by horseshoe crabs (Limulus poyphernus) in Delaware Bay, USA. Marine Biology 101: 143–151.Google Scholar
  5. Botton, Mark L., Robert E. Loveland, and Timothy R. Jacobsen. (1988). Beach erosion and geochemical factors: influence on spawning success of horseshoe crabs (Limulus polyphemus) in Delaware Bay. Marine Biology 99: 325–432. 1994. Site selection by migratory shorebirds in Delaware Bay, and its relationship to beach characteristics and abundance of horseshoe crab (Limulus po!yphemus) eggs. The Auk 111: 605–616.CrossRefGoogle Scholar
  6. Botton, Mark L. and John W. Ropes. (1988). An indirect method for estimating longevity of the horseshoe crab (Limulus polyphemus) based in epifaunal slipper snails (Crepidu!a fornicata). Journal Shellfish Research 7: 407–412.Google Scholar
  7. Brockmann, H. Jane. (1990). Mating behavior of horseshoe crabs, Limulus polyphemus. Behavior 114: 1992. The effects of age on male behavior in Limulus po!yphemus. Galaxea 11:61–62.Google Scholar
  8. Brockman, H. Jane. 1996. Satellite male groups in horseshoe crabs, Limulus po!yphemus, Ecology 102: 1–21.Google Scholar
  9. Brockmann, H. Jane and Dustin Penn. (1992). Male mating tactics in the horseshoe crab, Limulus polyphemus. Animal Behavior 44: 653–665.Google Scholar
  10. Caster, Kenneth E. (1938). A restudy of the tracks of Paramphibius. Journal Paleontology 12: 1–60.Google Scholar
  11. Clark, Kathleen E., Lawrence J. Niles and Joan Burger. (1993). Abundance and distribution of migrant shorebirds in Delaware Bay. Condor 95: 694–705.Google Scholar
  12. Eldredge, Niles. (1970). Observations on the burrowing behavior in Limulus polyphemus (Chelierata, Merostomata), with implications on the functional morphology of trilobites. American Museum Novitates 2436:17pp.Google Scholar
  13. Lockwood, Samuel. (1870). The horseshoe foot crab. American Naturalist 4: 257–274.Google Scholar
  14. Malz, Heinz. (1976). Solnhofener Plattenkalk: Eine Welt in Stein. Museum beim Solenhofer Aktien Verein, Maxberg, Germany: 109 pages.Google Scholar
  15. Packard, Alpheus Springer. (1900). On supposed merostomatous and other Paleozoic trails, with notes on those of limu!us. American Academy of Arts, Proceedings 36: 63–71.Google Scholar
  16. Penn, Dustin and H. Jane Brockmann. (1994). Nest-site selection in the horseshoe crab, Limulus polyphemus. Biological Bulletin 187: 373–384. (1995). Age-biased stranding and righting in horseshoe crabs, Limulus po!yphemus. Animal Behavior 49: 1531–1539.Google Scholar
  17. Rudloe, Anne and Jack Rudloe (photographs by Robert Sisson). (1981). The changeless horseshoe crab. National Geographic 159: 562–572.Google Scholar
  18. Shuster, Carl N. Jr. (1982). A pictorial review of the natural history and ecology of the horseshoe crab Limulus polyphemus, with reference to oilier Limulidae. In: Bonaventura, Joseph, Celia Bonaventura, and Shirley Tesh (eds.), Physiology and Biology of Horseshoe Crabs: Studies on Normal and Environmentally Stressed Animals. Alan R. Liss, Inc.: 1–52.Google Scholar
  19. Shuster, Carl N. Jr. and Mark L. Botlon. (1985). A contribution to the population biology of horseshoe crabs, Limulus polyphemus (L.), in Delaware Bay. Estuaries 8:363–372.Google Scholar
  20. Swan, Benjie Lynn. (1997). Personal communication on a tag return from an adult female tagged at Reeds Beach, on Delaware Bay, New Jersey on 23 May 1988 and recovered, alive, at Highs Beach, NJ on 31 May 1996. (also in: Swan, Benjie Lynn, William R. Hall Jr. and Carl N. Shuster, Jr. 1998. An Overview of Horseshoe Crab Spawning Activity Along the Shores of Delaware Bay, 1990–1997: 35 pages processed.Google Scholar

Copyright information

© Kluwer Academic Publishers 2002

Authors and Affiliations

  • Carl N. ShusterJr.
    • 1
  1. 1.The College of William and MaryVirginia Institute and School of Marine ScienceUSA

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