Scientific Literature - Classic
GRUNION SCIENTIFIC LITERATURE REVIEW
Journal Articles, Books, Chapters, Theses, Dissertations
Classic Grunion References (before 1990s)
The original description of the species
- Ayres, W. O. 1860. [Descriptions of fishes.] Proceedings of the California Academy of Sciences (Ser. 1): 73-77.
Early descriptions of spawning behavior and biology
- Barnhart, P. S. 1918. The spawning of the little-smelt Leuresthes tenuis (Ayres). California Fish & Game 4: 181-182.
- Clark, F. N. 1925. The life history of Leuresthes tenuis, an atherine fish with tide controlled spawning habits. Calif. Fish & Game 10: 1-51 plus plate.
- Clark, F. N. 1926. Conservation of the grunion. Calif. Fish & Game 12(4): 163-166.
- Clark, F. N. 1928a. Grunion at La Jolla. Calif. Fish & Game 14(1): 89-90.
- Clark, F. N. 1928b. Grunion on Cabrillo Beach. Calif. Fish & Game 14(4): 273-274.
- Clark, F. N. 1938. Grunion in southern California. Calif. Fish & Game 24(1): 49-54.
- Hubbs, Carl L., 1916. Notes on the marine fishes of California. University of California Publications in Zoology 16(13): 153-169.
- Jordan, D. S. 1926. The habits of the grunion. Science 63: 454.
- Jordan, D. S., and Carl L. Hubbs. 1919. A monographic review of the family of Atherinidae or silversides. Leland Stanford Junior Univ. Publ., Univ. Ser. (No.40):1–87, pls. I-XII.
- Thompson, W. F. 1919. The spawning of the grunion (Leuresthes tenuis). Calif. Fish and Game 5:1-27.
- Thompson, W. F. and J. B. Thompson. 1919. The spawning of the grunion (Leuresthes tenuis). Fish Bulletin 3: 1-29.

Boyd Walker, PhD was a professor of ichthyology at UCLA. His 1949 dissertation research on Scripps Beach in San Diego led to better understanding of the timing of California Grunion runs and improved information on the natural history of this charismatic animal.
He also organized the first example of grunion greeting, with volunteers in many locations along the coast observing grunion runs on the same two nights.
The Walker Scale, our metric for evaluating grunion runs, is named in honor of Dr. Boyd Walker. It was developed in the early 1990s by Karen Martin, Michael Schaadt, and Suzanne Lawrenz-Miller, published here in detail.
He also organized the first example of grunion greeting, with volunteers in many locations along the coast observing grunion runs on the same two nights.
The Walker Scale, our metric for evaluating grunion runs, is named in honor of Dr. Boyd Walker. It was developed in the early 1990s by Karen Martin, Michael Schaadt, and Suzanne Lawrenz-Miller, published here in detail.
Boyd Walker’s work on grunion
- Walker, B. W. 1947. The beach-spawning grunion. Aquarium J. Nov. 1947: 8-12.
- Walker, B. W. 1949. The periodicity of spawning by the grunion, Leuresthes tenuis, an atherine fish. PhD. Thesis, Univ. of Calif., Los Angeles, 1-166. https://escholarship.org/uc/item/1j33928x
- Walker, Boyd W. 1952. A guide to the grunion. Calif. Fish & Game 38(3): 409-420.
- Walker, Boyd W. 1959. The timely grunion. Natural History 68(6): 301-307.
- Walker, B. W. 1964. Fish, Moon and Tides: The Grunion Story. Academy Films, Los Angeles, California.
Grunion science by others 1930s-1950s
- David, L.R. 1939. Embryonic and early larval stages of the grunion, Leuresthes tenuis, and the sculpin, Scorpaena guttata. Copeia 1939(2): 75-81.
- Gallup, F. N. 1949. Banding recoveries of Tytlo alba. Bird-Banding 20: 150.
- McHugh, J.L. 1954. The influence of light on the number of vertebrae in the grunion Leuresthes tenuis. Copeia 1954(1): 23-25.
- Korringa, P. 1947. Relations between the moon and periodicity in the breeding of marine animals. Ecological Monographs 17: 349-381.
- Olson, A.C. 1950. Ground squirrels and horned larks as predators upon grunion eggs. Calif. Fish and Game 36(3): 323-327.
- Olson, A.C., Jr. 1955. Parasites of the grunion, Leuresthes tenuis (Ayres). PhD Dissertation, Oregon State College, 36 pp.
- Pearse, A. S. 1953. A new copepod parasite from the grunion. J. Parasitol. 39: 332-333.
- Phillips, J. B. 1943. Grunion in Monterey Bay. California Fish & Game 29: 82.
- Rechnitzer, A. 1952. A wave of fish. Natural History 61(7): 321-325.
- Stock, J. H. 1955. A new Bomolochus (Copepoda parasitica) from the California grunion. Beaufortia 5(49): 55-59.
Grunion science in the 1960s-1970s
- Breder, C. M. and D. E. Rosen. 1966. Modes of Reproduction in Fishes. Natural History Press, Garden City, NY.
- Brothers, E. B., C. P. Mathews, R. Lasker. 1976. Daily growth increments in otoliths from larval and adult fishes. Fishery Bulletin 74: 1-8.
- Castro, T., and M. Yoshida. 1974. Los grunones curiosas criaturas del mar. Ciencias Marinas, 1(2):92-95.
- Crane, J.M. 1969. The response of male grunion to a wiggling stick. Bull. So. Calif. Acad. Sci., 68(3): 191-193.
- Constant, C. L., 1976. The effects of hypersalinity upon the eggs and prolarvae of the Gulf of California grunion, Leuresthes sardina (Jenkins and Evermann, 1888). Univ. Ariz. M.S. thesis, 53 pp.
- Daugherty, A. E. 1962. California grunion, Leuresthes tenuis. Pp. 29-30 in California Ocean Fisheries Resources to the Year 1960. Calif. Dept. of Fish and Game, Sacramento, CA.
- Ehrlich, Karl Forrest. 1969. The effect of some environmental factors upon the incubation and rearing of Grunion, Leuresthes tenuis (Ayres). MS thesis. San Diego State College.
- Ehrlich, K. F. 1977. Inhibited hatching success of marine fish eggs by power plant effluent. Marine Pollution Bulletin 8: 228-229.
- Ehrlich, K. F. and D. A. Farris. 1970. Effect of temperature on the number of myomeres in newly hatched grunion (Leuresthes tenuis). Copeia 1970(4): 774-775.
- Ehrlich, K. F. and D. A. Farris. 1971. Some influences of the temperature on the development of the grunion, Leuresthes tenuis (Ayers). Calif. Fish & Game. 57(1): 56-68.
- Ehrlich, K. F. and D. A. Farris. 1972. Some influences of temperature on the development of the grunion Leuresthes tenuis, an atherine fish. Marine Biology 12: 267-271.
- Garey, W. F. 1962. Cardiac responses of fishes in asphyxic environments. Biological Bulletin 122: 362-368.
- Hubbs, Clark. 1965. Developmental temperature tolerance and rates of four southern California fishes, Fundulus parvipinnis, Atherinops affinis, Leuresthes tenuis, and Hypsoblennius sp. California Fish & Game 51(2): 113-122.
- Idyll, C. P. 1969. Grunion, the fish that spawns on land. National Geographic 135(5): 714-723.
- MacGregor, J. S. 1976. Ovarian development and fecundity of five species of California Current fishes. Calif. Coop. Oceanic Fish. Invest. Rep., 1976: 181-188.
- May, R. C. 1971. Effects of delayed initial feeding on larvae of the grunion, Leuresthes tenius (Ayers). U.S. Fish. Bull. 69(2): 411-425.
- May, R. C. 1974. Larval mortality in marine fishes and the critical period concept. Pp. 3-19 in The Early Life History of Fish. Springer Berlin Heidelberg.
- Mercieca, A., and R. C. Miller. 1969. The spawning of the grunion. Pacific Discovery 22(1): 26-27.
- Moffatt, N. M. 1974. A morphometric and meristic comparison of the Gulf grunion, Lenresthes sardina (Jenkins and Evermann) and the California grunion, Leuresthes tenuis (Ayres). Master's Thesis, University of Arizona, Tucson.
- Moffatt, N. M. 1977. Thermal effects on the survival and development of embryonic grunions, Leuresthes sardina and L. tenuis. Ph.D. dissertation, University of Arizona, Tucson, 88 pp.
- Moffatt, N. M. & D.A. Thomson. 1975. Taxonomic status of the Gulf grunion (Leuresthes sardina) and its relationship to the California grunion (L. tenuis). Trans. San Diego Soc. Nat. Hist. 18(4): 75-84.
- Moffatt, N. M. & D.A. Thomson. 1978. Reciprocal hybridization between the California and Gulf of California grunions, Leuresthes tenuis and Leuresthes sardina (Atherinidae). Fish. Bull. 76(2): 476-480.
- Moffatt, N. M. & D. A. Thomson. 1978. Tidal influence on the evolution of egg size in the grunions (Leuresthes, Atherinidae). Environmental Biol. Fish 3: 267-273.
- Muench, K. A. 1977. Behavioral ecology and spawning periodicity of the Gulf of California grunion, Leuresthes sardina. PhD dissertation, Univ. Arizona, 92 pp.
- Olson, A.C., Jr. 1972. Argulus melanostictus and other parasitic crustaceans on the California grunion, Leuresthes tenuis (Osteichthyes: Atherinidae). Journal of Parasitology 58: 1201-1204.
- Olson, A. C., Jr. 1975. Metacercaria of Bucephalopsis labiatus (Trematoda: Bucephalidae) in the California grunion, Leuresthes tenuis. Journal of Parasitology 61: 50.
- Olson, A. C., Jr. 1976. Asymphylodora atherinopsidis (Trematoda: Monorchiidae) from the California grunion, Leuresthes tenuis, including a redescription. Journal of Parasitology 63: 295-298.
- Olson, A. C., Jr. 1978. Lepocreadium manteri sp. N. (Trematoda: Lepocreadiidae) from the California grunion, Leuresthes tenuis, and its hyperparasitic microsporidan. Proceedings Helminthological Society Washington 45: 155-157.
- Olson, L. J. 1979. Host-parasite associations of the grunions, Leuresthes sardina and Leuresthes tenuis from the Gulf of California. PhD Dissertation, University of Arizona, 97 pp.
- Reynolds, W. W. 1973. Orientation responses of laboratory-reared larval and juvenile Gulf grunion (Leuresthes sardina) to artificial gradients. PhD Dissertation, Univ. Arizona, 93 pp.
- Reynolds, W. W. & D. A. Thomson. 1974. Ontogenetic change in the response of the Gulf of California grunion, Leuresthes sardina (Jenkins & Evermann), to a salinity gradient. J. Exp. Mar. Biol. Ecol. 14: 211-216.
- Reynolds, W. W. & D. A. Thomson. 1974. Responses of young Gulf grunion, Leuresthes sardina, to gradients of temperature, light, turbulence and oxygen. Copeia 1974: 747-758.
- Reynolds, W. W., D. A. Thomson & M. E. Casterlin. 1976. Temperature and salinity tolerances of larval California grunion, Leuresthes tenuis (Ayres): A comparison with gulf grunion, L. sardina (Jenkins & Evermann). Journal of Experimental Marine Biology and Ecology 24: 73-82.
- Reynolds, W. W., D. A. Thomson & M. E. Casterlin. 1977. Responses of young California grunion Leuresthes tenuis to gradients of temperature and light. Copeia 1977(1): 144-149.
- Scholander, P. F., E. Bradstreet & W. F. Garey. 1962. Lactic acid response in the grunion. Comp. Biochem. Physiol. 6: 201-203.
- Southern California Ocean Studies Consortium of the California State University and Colleges. 1978. Report on grunion spawning at McGrath State Beach, Ventura County, California. Prepared for the United States Army Corps of Engineers, Los Angeles District. Contract # DACW09-78-C0008.
- Thomson, D. A. & K. A. Muench. 1976. Influence of tides and waves on the spawning behavior of the Gulf of California grunion, Leuresthes sardina (Jenkins and Evermann). Bull. Southern California Acad. Sci. 75: 198-203.
- Valentine, D. W. & M. Soulé. 1973. Effect of p'-DDT on developmental stability of pectoral fin rays in the grunion, Leuresthes tenuis. Fishery Bulletin 71, 921–920.
Grunion science in the 1980s
- Allen, L. G., M. H. Horn, F. A. Edmonds II & C. A. Usui. 1983. Structure and seasonal dynamics of the fish assemblage in the Cabrillo Beach area of Los Angeles Harbor, California. Bull. Southern California Acad. Sci. 82: 47-70.
- Ambrose, M., N. S. Walker & J. Hawk. 1981. Captain Mike’s Complete Guide to Grunion Hunting. Oak Tree Press, La Jolla, CA.
- Borthwick, P. W., J. M. Patrick, Jr. & D. P. Middaugh. 1985. Comparative acute sensitivities of early life stages of Atherinid fishes to chlorpyrifos and thiobencarb. Arch. Environ. Contam. Toxicol. 14: 465-473.
- Canning, E. U. & A. C. Olson, Jr. 1980. Nosema lepocreadii sp. n., a parasite of Lepocreadium manteri (Digenea: Lepocreadiidae) from the gut of the California grunion, Leuresthes tenuis. J. Parasitology 66: 154-159.
- Canning, E. U. & A. C. Olson, Jr. 1983. The ultrastructure of Nosema lepocreadii Canning and Olson, 1979 (Microspora, Nosematidae) and its relevance to the generic diagnosis of Nosema Nageli, 1857. J. Parasitology 69: 143-151.
- Clark, J. R., J. M. Patrick, Jr., D. P. Middaugh & J. C. Moore. 1985. Relative sensitivity of six estuarine fishes to carbophenothion, chlorpyrifos, and fenvalerate. Ecotoxicology and Environmental Safety 10: 382-390.
- Crabtree, C. B. 1983. Assessment of genetic differentiation within the atherinid genus Leuresthes. Isozyme Bull. 16: 77.
- Crabtree, C. B. 1987. Allozyme evidence for the phylogenetic relationships within the silverside subfamily Atherinopsinae. Copeia 1987: 860-867.
- Ehrlich, K.F. & Muszynski, G. 1981. The relationship between temperature-specific yolk utilization and temperature selection of larval grunion. Rapp. P.-V. Reun. Cons. Int. Explor. Mer 178: 312-313.
- Ehrlich, K. F. & G. Muszynski. 1982. Effects of temperature on interactions of physiological and behavioral capacities of larval California grunion: adaptations to the planktonic environment. J. Exp. Mar. Biol. Ecol. 60: 223-224.
- Fritzsche, R.A., R.H. Chamberlin & R.A. Fisher. 1985. Species profiles: life histories and environmental requirements of coastal fishes and invertebrates (Pacific Southwest) – California grunion. U.S. Fish and Wildlife Service Biological Report 82(11.28). U.S. Army Corps of Engineers, TR EL-82-4, 12 pp.
- Goodman, L. R., D. J. Hansen, G. M. Cripe, D. P. Middaugh & J. C. Moore. 1985. New early life-stage toxicity test using the California grunion (Leuresthes tenuis) and results with chlorpyrifos. Ecotoxicology and Environmental Safety 10:12-21.
- Gruber, D., E. H. Ahlstrom & M. M. Mullin. 1982. Distribution of ichthyoplankton in the Southern California Bight. Calif. Coop. Ocean. Fish. Invest. Rep. 23: 172-179.
- Hose, J.E. & H. W. Puffer. 1984. Oxygen consumption rates of grunion (Leuresthes tenuis) embryos exposed to the petroleum hydrocarbon, benzo[a]pyrene. Environmental Research 35: 413-420.
- Middaugh, D. P., H. W. Kohl & L. E. Burnett. 1983. Concurrent measurement of intertidal environmental variables and embryo survival for the California grunion, Leuresthes tenuis, and Atlantic silverside, Menidia menidia (Pisces: Atherinidae). Calif. Fish and Game 69: 89-96.
- Newton, F., A. Thum, B. Davidson, A. Valkirs & P. Seligman. 1985. Effects on the growth and survival of eggs and embryos of the California grunion (Leuresthes tenuis) exposed to trace levels of tributylin. Naval Oceans Systems Center, Technical Rep. 1040, San Diego, CA 35 pp.
- Spratt, J. D. 1980. California grunion spawn in Monterey Bay. Calif. Dept of Fish and Game 67: 134.
- Spratt, J. D. 1986. The amazing grunion. Marine Resource Leaflet No. 3, Calif. Dept. of Fish and Game.
- Taylor, M. 1984. Lunar synchronization of fish reproduction. Transactions of the American Fisheries Society 113: 484-493.
- Watson, W. & R. L. Davis, Jr. 1989. Larval fish diets in shallow coastal waters off San Onofre, California. Fishery Bulletin, U.S. 87: 569-591.
- Winkler, D. L., K. L. Duncan, J. E. Hose & H. W. Puffer. 1983. Effects of benzo(a)pyrene on the early development of the California grunion, Leuresthes tenuis (Pisces, Atherinidae). U.S. National Marine Fisheries Services, Fishery Bulletin 41(3): 473-481.