Sand Lance - Alaska Science Center - the USGS

340 Pages·2007·1.01 MB·English

United States Sand Lance: A Review
Department of
Agriculture
of Biology and Predator
Forest Service
Pacific Northwest
Research Station Relations and Annotated
Research Paper
PNW-RP521
Bibliography
September 1999
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; MARYF. 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; ROBERTH. ARMSTRONG was a biologist (now retired), Alaska
Department of Fish and Game, Juneau, AK; and JOHN F. PIATTis 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
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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
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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.
Copepodsare 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 ValdezOil 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 in intertidal and shallow subtidal sub-
strates, venturing out only to feed or spawn. All Ammodytes species appear to be rela-
tively dormant in the winter and one (A. personatus) also estivates during summer.
Copepods are their 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 plankton blooms and
have several mechanisms to resist starvation. Density-dependent conditions are com-
mon for both adults and juveniles. Sand lance are host to a wide variety of parasites,
although prevalence is unclear. Because of their abundance, schooling behavior, ener-
getic content, and size, sand lance are an important forage species throughout their
range for a wide variety of marine predators.
Keywords: Sand lance, sand eel, Ammodytes, personatus, hexapterus, americanus,
dubius, tobianus, marinus,forage fish.
1
Taxonomy and Phylum: Chordata
Distribution Subphylum: Vertebrata (Craniata)
Superclass: Gnathostomata
Grade: Pisces
Class: Osteichthyes
Subclass: Actinopterygii
Infraclass: Neopterygii
Division: Halecostomi
Subdivision: Teleostei
Infradivision: Euteleostei
Superorder: Acanthopterygii
Order: Perciformes
Suborder: Ammodytoidei
Family: Ammodytidae
Subfamily: Ammodytinae
Genus: Ammodytes
Species: personatus, hexapterus, americanus, dubius, marinus, tobianus
Ammodytidae is the sole family within the suborder Ammodytoidei. Numerous papers
have been written on the designations below this level of classification, for which
there is considerable confusion, continuing the thought of Gill (1904) who stated:
There are few fishes respecting whose affinities there has been so much
diversity of opinion...as the sand launces or Ammodytids.
Current literature suggests that the Ammodytidae includes 7 genera including a total
of 20 species (Ida and others 1994, Stevens and others 1984): Gymnammodytes,
Hyperoplus,and Ammodytesusually live in temperate and boreal conditions, unlike
Protammodytes, Lepidammodytes, Bleekeria,andAmmodytoides, which live in more
tropical regions (Ida and others 1994, Stevens and others 1984).
Artedi (1738) derived the name for the genus Ammodytes from the Latin ammos
meaning “sand” and dytes meaning “diver.” Species designations and names have
changed often during the past 30 years. Confusion results largely from the overlapping
meristic characteristics among species, which differ with environmental conditions
and are therefore suggestive, rather than conclusive, for making species designations
(Inoue and others 1967, Okamoto 1989). The wide latitudinal range of sand lance
2