Of the Biology, Ecology, and Fisheries of the Clupeoid Fishes Occurring in the Gulf of Maine
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Northeast Fisheries Science Center Reference Document 00-02 An Overview· of the Biology, Ecology, and Fisheries of the Clupeoid Fishes Occurring in the Gulf of Maine by Thomas A. Munroe National Marine Fisheries Serv., National Systematics Lab., National Mus. of Natural History, 10th & Constitution Ave., N W, Washington, DC 20560-0153 U.S. DEPARTMENT OF COMMERCE National Oceanic and Atmospheric Administration National Marine Fisheries Service Northeast Region Northeast Fisheries Science Center Woods Hole, Massachusetts March 2000 Northeast Fisheries Science Center Reference Documents This series is an informal report series designed to assUre the long-term documentation and to enable the timely transmission of research results emanating from various Center laboratories. The reports in this series receive internal scientific review but no technical or copy editing. The National Marine Fisheries Service does not endorse any proprietary material, process, or product mentioned in these reports. To obtain additional copies of reports in this series, contact the senior Center author of the desired report. Refer to the title page of the desired report for the senior Center author's name and mailing address. This report's publication history is as follows: manuscript submitted for review--February 7, 2000; manuscript accepted through technical review--February 17,2000; manuscript accepted through policy review--February 24 2000; and camera-ready copy submitted for publication- March 30, 2000. This report may be cited as: Munroe, T. 2000. An overview of the biology, ecology, and fisheries of the clupeoid fishes occurring in ·the GulfofMaine. Northeast Fish. Sci. Cent. Ref Doc. 00-02; 226 p. Available from: National Marine Fisheries Service, 166 Water Street, Woods Hole, MA 02543-1026. Contents Abstract .................................... : .................................................................................................... iv Chapter 1. Introduction................................................................................................................ 1 Chapter 2. Order Clupeiforrnes .......................................................................................... :......... 2 Suborder C1upeoidei .......................................................................................................... 3 Key to Families of Clupeoid Fishes in the Gulf of Maine ................................................ 4 Chapter 3. Family C1upeidae, the herrings .................................................................................. 5 Key to genera and species of Gulf of Maine Herrings ....................................................... 6 Chapter 4. Subfamily Clupeinae .................................................................................................. 7 Chapter 5. Genus Clupea Description and distinctions ................................................................ 7 Chapter 6. Atlantic herring, Clupea Jwrengus. account ..... ... ... ...... ........ .... ..... ... ............ .... .......... 8 Chapter 7. Genus Opisthonema Description and distinctions ..................................................... 69 Chapter 8. Opisthonema oglinum, Atlantic thread-herring, account ........................................... 70 Chapter 9. Sub-family Alosinae .................................................................................................. 72 Chapter 10. Genus A/osa Description and distinctions ................................................................ 72 Chapter 11. Alewife, A/osa pseudoharengus, account ................................................................ 73 Chapter 12. Blueback herring, Alosa aestivalis, account ............................................................ 93 Chapter 13. American Shad, A/osa sapidissima, account ......................................................... 105 Chapter 14. Hickory shad, A/osa mediocris, account ................................................................ 139 Chapter 15. Genus Brevoortia Description and distinctions (with 1. Smith) ............................ 142 Chapter 16. Atlantic menhaden, Brevoortia tyrannus, account (with J. Smith) ........................ 143 Chapter 17. Subfamily Dussumieriinae .................... " ............................................................... 160 Chapter 18. Genus Etrumeus Description and distinctions ....................................................... 161 Chapter 19. Atlantic Round Herring, Etrumeus teres, account ................................................. 161 Chapter 20. Suborder Engrauloidea. Family Engraulidae ........................................................ 164 Chapter 21. Genus Anchoa Description and distinctions .......................................................... 165 Key to species of Anchovies occurring in the Gulf of Maine ............................................ 166 Chapter 22. Bay Anchovy, Anchoa mitchilli, account ............................................................... 166 Chapter 23. Striped Anchovy, Anchoa hepsetus, account ......................................................... 172 Acknowledgments ...................................................................................................................... 174 Personal Communications .......................................................................................................... 175 Literature Cited ............................................................................................................................ 176 iii ABSTRACT Clupeoid species are important ecological components within marine and riverine ecosystems of the Gulf of Maine. This report summarizes information (1953-1995) for eight species of c1upeid fishes (Clupeidae) and two species of anchovy (Engraulidae) occurring in the Gulf of Maine. Accounts are presented for Atlantic herring (Clupea harengus), Atlantic thread herring (Opisrhonema oglinum), Atlantic round herring (Etrumeus teres), Atlantic menhaden (Brevoortia tyrannus), Hickory shad (Alosa mediocris), American shad (A. sapidissima), Alewife (A. pseudoharengus), Blueback herring (A. aestivalis), Bay anchovy (Anchoa mitchilli) and Striped anchovy (Anchoa hepsetus). Identification keys to families, genera and species are. provided. Systematic, biological, ecological, distributional and fisheries data are summarized for each species, with emphasis on data derived from populations in the Gulf of Maine. Whenever possible, natural and human-induced environmental factors affecting populations of c1upeoid species in the Gulf are discussed. An extensive bibliography of the literature on these species is also provided, with special concentration on populations occurring in the western Atlantic. Recent declines in abundance, especially of anadromous species and the Atlantic herring, are discussed relative to environmental and fishery practices, as well as the conservation measures being afforded these species. Although considerable information is available for most c1upeoid species occurring in the Gulf of Maine, significant gaps in our information remain for some species. Information needs should be addressed to improve understanding of the population biology and ecological interactions of these species within Gulf of Maine communities. iv INTRODUCTION Because of the relative size of their populations and their position within trophic food webs, clupeoid fishes play significant roles in many ecosystems in which they occur. The Gulf of Maine is no exception. Here, as elsewhere, clupeoid fishes serve important roles both as predator and prey species. Eight species of the Family Clupeidae, including the Atlantic herring (Clupea harengus), Atlantic thread herring {Opisthonema oglinum), Atlantic round herring (Etrumeus teres), Atlantic menhaden (Brevoortia tyrannus), Hickory shad (Alosa mediocris), American shad (A. sapidissima), Alewife (A. pseudoharengus) and Blueback herring (A. aestivalis), and two species of anchovies (Engraulidae), the Bay anchovy (Anchoa mitchilli) and Striped anchovy (Anchoa hepsetus), occur in the Gulf of Maine. Large popUlation sizes (historically, at least, for some species) and general availability to fishing gears have subjected many of these fishes to heavy exploitation in commercial, industrial and recreational fisheries in the Gulf and elsewhere. Populations of most commercially important Gulf of Maine clupeoid fishes have undergone considerable fluctuations, with most species, especially the anadromous alosines, presently represented by much smaller populations than those recorded historically. Smaller population sizes have resulted from a number of factors, including such anthropogenic activities as overfishing and habitat loss and destruction. Because of their ecological and commercial value, clupeoid fishes in the Gulf of Maine such as the Atlantic herring, Atlantic menhaden, American shad, Alewife and Bltleback herring, have been the subject material for much detailed research. This report compiles and summarizes available information on the systematics, identification, biology, ecology and fisheries of species of the Family Clupeidae, as well as, two species of anchovies (Engraulidae), occurring in the Gulf of Maine. This report was conducted while preparing chapters summarizing available information for species accounts for a forthcoming revised edition of Bigelow and Schroeder's Fishes of the Gulf of Maine, edited by B.B. Collette and G. Klein-McPhee. However, after preliminary