United States Department of Agriculture Sand Lance: A Review Forest Service Pacific Northwest of Biology and Predator Research Station Research Paper Relations and Annotated PNW-RP 521 September 1999 Bibliography Authors MARTIN D. ROBARDS is a graduate student, Memorial University of Newfoundland, and is supported by the U.S. Geological Survey, Biological Resources Division, 1011 E. Tudor Road, Anchorage, AK 99503; MARY F. WILLSON was a research ecologist, U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station, 2770 Sherwood Lane, Juneau, AK 99801, and currently is with The Nature Conser- vancy, Chicago, IL; ROBERT H. ARMSTRONG was a biologist (now retired), Alaska Department of Fish and Game, Juneau, AK; and JOHN F. PIATT is a senior research scientist, U.S. Geological Survey, Biological Resources Division, 1011 E. Tudor Road, Anchorage, AK 99503. The Exxon Valdez Oil Spill Trustee Council conducts all programs and activities free from discrimination, consistent with the Americans With Disabilities Act. This publica- tion is available in alternative communication formats upon request. Please contact the Restoration Office to make any necessary arrangements. Any person who believes she or he has been discriminated against should write to either the EVOS Trustee Council, 645 G Street, Suite 401, Anchorage, AK 99501, or O.E.O., U.S. Department of the Interior, Washington, DC 20240. Cover photo by Robert H. Armstrong; drawing by Richard Carstensen. Sand Lance: A Review of Biology and Predator Relations and Annotated Bibliography Martin D. Robards Mary F. Willson Robert H. Armstrong John F. Piatt Editors Exxon Valdez Oil Spill Restoration Project 99346 Final Report U.S. Department of Agriculture Forest Service Pacific Northwest Research Station Portland, Oregon Research Paper PNW-RP-521 September 1999 This page has been left blank intentionally. Document continues on next page. Abstract Robards, Martin D.; Willson, Mary F.; Armstrong, Robert H.; Piatt, John F., eds. 1999. Sand lance: a review of biology and predator relations and annotated bibli- ography. Res. Pap. PNW-RP-521. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station. 327 p. Six species of sand lance (Ammodytes) in temperate and boreal regions are currently recognized. Sand lance can occupy a wide range of environmental conditions, but all appear to be dormant predominantly in winter, and one species is in summer also. They lack a swim bladder and spend much of their time buried in specific substrates. Copepods are the primary food. Spawning usually occurs in fall or winter (although some species also spawn in spring), eggs are demersal, and larvae may hatch at times of low food abundance. Sand lance usually occur in schools and are regarded as a relatively high-quality forage fish. Sand lance constitute a major prey for at least some populations of over 100 species of consumer, including 40 species of birds, 12 species of marine mammals, 45 species of fishes, and some invertebrates. Variation in the availability of sand lance (and other forage fishes) can have major effects on the breeding success and survival of their predators. Commercial fishing and other pressures on sand lance populations potentially have ramifying effects on many species of wildlife. The bibliography contains over 1,700 references on the family Ammodytidae, with an emphasis on the genus Ammodytes. Keywords are provided for each reference and have been further organized into taxonomic, geographic, subject, and predator indexes. Keywords: Sand lance, sand eel, Ammodytes, personatus, hexapterus, americanus, dubius, tobianus, marinus, predator/prey relations, seabirds, marine mammals, forage fish, predatory fish, Alaska, adaptations, spawning, phenology, morphology, habitat, recruitment, ecology, bibliography. Summary Although much taxonomic confusion exists within the genus Ammodytes, six species are currently recognized: personatus, hexapterus, americanus, dubius, tobianus, and marinus. Sand lance are both euryhaline and eurythermal, as well as tolerant of reduced oxygen concentrations. The absence of a swim bladder allows this narrow, elongate fish to spend much time buried dormant in intertidal and shallow subtidal substrates, venturing out only to feed or spawn. All Ammodytes species appear to be relatively dormant in winter and one (A. personatus) also estivates during sum- mer. Copepods are the primary food source, allowing for rapid energy accumulation during secondary production blooms. Life spans range from 3 to 12 years within the genus. Spawning usually occurs in fall or winter (although some species spawn in spring) with the production of demersal, slightly adhesive eggs. Early development is oxygen and temperature dependent. Larval sand lance hatch before spring plank- ton blooms and have several mechanisms to resist starvation. Density-dependent conditions are common for both adults and juveniles. Sand lance are host to a wide variety of parasites. Sand lance are a nutitrious and important prey for over 100 species of consumers, including 40 species of birds, 12 species of marine mammals, 45 species of fishes (including many of commercial importance, such as salmon and flatfishes), and some squid. Reproductive success of at least 10 avian species has been correlated with availability of sand lance prey: great skua, parasitic jaeger, shag, black-legged kitti- wake, arctic tern, common tern, common murre, Atlantic puffin, tufted puffin, and rhi- noceros auklet. Sand lance are among the more nutritious of forage fishes, with relatively high levels of certain vitamins and energy density; however, they also are capable of transmitting paralytic shellfish poisoning and other toxins to their preda- tors. Sand lance constitute an important element in many marine food webs. Cascading effects throughout the marine food web may occur when severe perturbations are imposed on critical parts of the web. In areas where sand lance are commercially harvested, future problems in resource exploitation may be avoided, not only by using more and better science but also by using conservative harvesting strategies. Over 1,700 published and unpublished references on sand lance are listed for the family Ammodytidae, with emphasis on the genus Ammodytes. Keywords accompany each reference and are organized further into taxonomic, geographic, subject, and predator indexes. The taxonomic listings use names as given by authors, with no attempt at making synonomies. Abstracts are provided when available. Study History: Exxon Valdez Oil Spill Trustee Council restoration project 98346 was initiated out of interest by M. Willson and R. Armstrong in the behavior of birds foraging for buried sand lance in intertidal areas of southeast Alaska. As the biblio- graphic work proceeded, these two authors learned of work begun earlier in APEX project 96306 on ecology and demographics of the sand lance in Lower Cook Inlet, Alaska, by M. Robards and J. Piatt. This latter work included a review of the known biology and literature pertaining to sand lance. APEX project 99346 was estab- lished to support publication of the review and a comprehensive bibliography of 1,700 citations in one volume. Cooperation with various libraries (particularly ARLIS and the Forestry Sciences Library in Juneau) was invaluable to this effort. Project Data: Description of data—The bibliography was compiled from a variety of sources, including computer searches and traditional literature searches. Format— The bibliographic data were entered into ClarisWork® and reformatted by the Pacific Northwest Research Station, Communications Group. Custodian—None. Availability— Hard copies will be available for 2 years from date of publication from the Pacific Northwest Research Station, PO Box 3890, Portland, OR 97208-3890. The entire document also is available online in Portable Document Format (pdf). To view publi- cations and see instructions about downloading the Adobe Acrobat Reader, navigate to http://www.fs.fed.us/pnw/pubs.htm. Contents 1 Biology of the Genus Ammodytes, the Sand Lances Martin D. Robards and John F. Piatt 2 Taxonomy and Distribution 5 Morphology 5 Habitat 6 Behavior 6 Daily 6 Seasonal 7 Physiological Adaptations 7 Temperature and Salinity 7 Exposure 7 Summer Estivation 7 Winter Hibernation 8 Schooling Behavior 8 Feeding 9 Seasonal Changes in Proximate Composition 10 Age, Growth, and Life Span 11 Maturation 11 Spawning 12 Eggs 12 Larvae 13 Recruitment 14 Mortality 14 Predation 15 Parasites 15 Acknowledgments 17 Sand Lance as Cornerstone Prey for Predator Populations Mary F. Willson, Robert H. Armstrong, Martin D. Robards, and John F. Piatt 18 Introduction 18 Sand Lance as Prey 21 Antipredator Behavior 22 Predators 24 Complex Interactions of Forage Fishes, Their Predators, and Commercial Fisheries 27 What Do We Need to Know? 28 Acknowledgments 39 Literature Cited 45 Annotated Bibliography Robert H. Armstrong, Mary F. Willson, Martin D. Robards, and John F. Piatt 45 Introduction 46 Bibliography 318 Keyword Index Biology of the Genus Ammodytes, The Sand Lances Martin D. Robards and John F. Piatt Abstract Although much taxonomic confusion exists within the genus Ammodytes, six species are currently recognized: personatus, hexapterus, americanus, dubius, tobianus, and marinus. Sand lance are both euryhaline and eurythermal, as well as tolerant of reduced oxygen concentrations. The absence of a swim bladder allows this narrow, elongate fish to spend much time buried dormant