- f., r:, Fscarch Center -;-:iirt: Service ~~4cvard -. r”jxl Biological Report 82(11.40) TR EL-82-4 June 1985 Library IQtioxM Wetlands Research Centw U. S. Fish and Wildlife Service 700 Cajundome Boulevard Lafayette, La. 70506 SpeciesProfiles: Life Histories and EnvironmentalRequirements of CoastalFishes and Invertebrates(Mid-Atlantic)
MUMMICHOGAND STRIPED KILLIFISH
QL 155 .S63 no. 82 11.40
Coastal Ecology Group Fish and Wildlife Service Waterways Experiment Station U.S. Department of the Interior U.S. Army Corps of Engineers This is one of the first reports to be published in the new "Biological Report" series. This technical report series, published by the Research and Development branch of the U.S. Fish and Wildlife Service, replaces the "FWS/OBS" series published from 1976 to September 1984. The Biolog- ical Report series is designed for the rapid publication of reports with an application orientation, and it continues the focus of the FWS/OBS series on resource management issues and fish and wildlife needs. Biological Report 82(11.40) TR EL-82-4 June 1985
Species Profiles: Life Histories and Environmental Requirements of Coastal Fishes and Invertebrates (Mid-Atlantic)
MUMMICHOG AND STRIPED KILLIFISH
Barbara J. Abraham Department of Biological Sciences P.O. Box 6565 Hampton University Hampton, VA 23668
Project Manager Carroll L. Cordes Project Officer Pepsi Nunes National Coastal Ecosystems Team U.S. Fish and Wildlife Service 1010 Gause Boulevard Slidell, LA 70458
Performed for Coastal Ecology Group Waterways Experiment Station U.S. Army Corps of Engineers Vicksburg, MS 39180
and
National Coastal Ecosystems Team Division of Biological Services Research and Development Fish and Wildlife Service U.S. Department of the Interior Washington, DC 20240 This series should be referenced as follows:
U.S. Fish and Wildlife Service. 1983-19 . Species profiles: life histories and environmental requirements of coasts fishes and invertebrates. U.S. Fish Wildl. Serv. Biol. Rep. 82(11). U.S. Army Corps of Engineers, TR EL-82-4.
This profile should be cited as follows:
Abraham, B.J. 1985. Species profiles: life histories and environmental requirements of coastal fishes and invertebrates (Mid-Atlantic)--mummichog and striped killifish. U.S. Fish Wildl. Serv. Biol. Rep. 82(11.40). U.S. Army Corps of Engineers, TR EL-82-4. 23 PP. PREFACE
This species profile is one of a series on coastal aquatic organisms, principally fish, of sport, commercial, or ecological importance. The profiles are designed to provide coastal managers, engineers, and biologists with a brief comprehensive sketch of the biological characteristics and environmental requirements of the species and to describe how populations of the species may be expected to react to environmental changes caused by coastal development. Each profile has sections on taxonomy, life history! ecological role, environmental requirements, and economic importance, if applicable. A three-ring binder is used for this series so that new profiles can be added as they are prepared. This project is jointly planned and financed by the U.S. Army Corps of Engineers and the U.S. Fish and Wildlife Service.
Suggestions or questions regarding this report should be directed to one of the following addresses.
Information Transfer Specialist National Coastal Ecosystems Team U.S. Fish and Wildlife Service NASA-Slide11 Computer Complex 1010 Gause Boulevard Slidell, LA 70458
or
U.S. Army Engineer Waterways Experiment Station Attention: WESER-C Post Office Box 631 Vicksburg, MS 39180
iii CONVERSION TABLES
Metric to U.S. Customary
Multiply !!Y To Obtain millimeters (mn) 0.03937 inches centimeters (cm) 0.3937 inches meters (m) 3.281 feet kilometers (km) 0.6214 miles square meters (m2) 10.76 square feet square kilometers (km2) 0.3861 square miles hectares (ha) 2.471 acres liters (1) 0.2642 gallons cubic meters (m3) 35.31 cubic feet cubic meters 0.0008110 acre-feet milligrams (mg) 0.00003527 ounces grams (9) 0.03527 ounces kilograms (k ) 2.205 pounds metric tons 9 t) 2205.0 pounds metric tons 1.102 short tons kilocalories (kcal) 3.968 British thermal units
Celsius degrees 1.8("C) + 32 Fahrenheit degrees
U.S. Customary to Metric inches 25.40 millimeters inches 2.54 centimeters feet (ft) 0.3048 meters fathoms 1.829 meters miles (mi) 1.609 kilometers nautical miles (mni) 1.852 kilometers square feet (ft2) 0.0929 square meters acres 0.4047 hectares square miles (mi2) 2.590 square kilometers gallons (gal) 3.785 liters cubic feet (ft3) 0.02831 cubic meters acre-feet 1233.0 cubic meters ounces (02) 23.35 grams pounds (lb), . 0.4536 kilograms short tons (ton) 0.9072 metric tons British thermal units (Btu) 0.2520 kilocalories
Fahrenheit degrees 0.5556("F - 32) Celsius degrees
iv CONTENTS
Page
PREFACE ...... iii CONVERSION TABLE ...... iv ACKNOWLEDGMENTS ...... vi
NOMENCLATURE/TAXONOMY/RANGE ...... 1 MORPHOLOGY/IDENTIFICATION AIDS ...... REASON FOR INCLUSION IN SERIES ...... : LIFE HISTORY ...... 5 Reproductive Physiology ...... Spawning Season and Periodicity ...... z Spawning Behavior ...... 6 Spawning Site ...... 6 Eggs ...... 6 Yolk-Sac Larvae ...... 7 Larvae ...... 7 Juveniles ...... 7 Adults ...... 7 Movement Patterns and Seasonal Use of Habitats ...... GROWTH CHARACTERISTICS ...... : Length-Weight Relationships ...... 8 Seasonal and Annual Growth and Production ...... 8 POPULATION DYNAMICS ...... 9 Population Size and Structure ...... 9 Mortality ...... 9 ECOLOGICAL RqLE ...... 10 Feeding Behavior/Food Habits ...... 10 Predators ...... 11 Competitors ...... 11 Parasites ...... 12 ENVIRONMENTAL REqUIREMENTS/TOLERANCES ...... 12 Temperature ...... Salinity ...... :: Temperature-Salinity Interactions ...... 14 Contaminants ...... 14 Other Environmental Factors ...... 14
LITERATURE CITED ......
V ACKNOWLEDGMENTS
I am grateful to M.H. Taylor, University of Delaware, and two anonymous reviewers who commented on an early draft of the manuscript.
vi Adult male killifish
Adult female killifish
Figure 1. Mummichog and striped killifish.
MUMMICHOG AND STRIPED KILLIFISH
NOMENCLATURE/TAXONOMY/RANGE Class ...... Osteichthyes Order ...... Atheriniformes Scientific name . . . Fundulus hetero- Family ...... Cyprinodontidae cl;.l;s (L innaeus) -(Robins et al. Geographic range ...... Gulf of Preferred common name ...... mummichog St. Lawrence south to northeastern (Figure 1, top) Florida (Figure 2). (F. grandis, Other common names ...... mud minnow, previously E. h_ grandi>, includes pike minnow, mud dabbler, gudgeon populations which range from peninsu- (Hildebrand and Schroeder 1928); lar Florida to Tampico, Mexico.) saltwater minnow, killie, mummie Also found on Sable Island, Nova (Ursin 1977); marsh minnow (Teal and Scotia (Garside 1969) and Bermuda Burns 1979); common killifish, (Rosen 1973). Salt, brackish, and brackish water chub (Northcutt and freshwater (Rosen 1973); character- Davis 1983) istically close to shore (Daiber 1982). 1 PE NSYLVANIA
PHILADELPHIA
h ATLANTIC OCEAN MARYLAND ~ -7 BALTIMOREjj
IASHINGTON, D.C., 4
VIRGINIA _ a
0 50 100 KILOMETERS
B Coastal distribution
NORTH CAROLINAcn h CAPE HATTERAS
Figure 2. Mid-Atlantic coast distribution of the Mid-Atlantic region close to shore (within 110 for mummichogs: Armstrong and Child 1965) and in bays, estuaries and tidal creeks (Hardy 1978). Striped killifish penetrate tidal rivers only as far as the mean boundary between fresh and brackish water; mummichogs penetrate as 75
2 Recent systematic studies . . . . . On the In the Cyprinodontidae there is no basis of differences in egg morphol- lateral line and the upper surface of ogy, gene frequencies, and spawning the head is conspicuously flattened sites between populations north and (Hardy 1978). The anal fin in the male south of northern New Jersey, Morin is not modified as an intromittent and Able (1983) proposed retaining organ, and the third anal ray is the subspecies names F. h. hetero- branched. In the Fundulinae the jaw clitus for the southern population teeth are conical without cusps. Among and F. h. macrolepidotus for the members of the genus Fundulus, the body northern.- Populations in the upper is not sharply angular or trapezoidal, Delaware and Chesapeake Bays seem to and there is more than one series of be relicts of the northern sub- jaw teeth (Rosen 1973). The mouth is species. (These distributions do not terminal or the lower jaw projects match those of the subspecies as slightly (Hildebrand and Schroeder originally named.) 1928).
Scientific name . . . . . Fundulus majalis In both mummichogs and striped (Walbaum) (Robins et al. 1980) killifish, males are smaller than Preferred common name ...... striped females after the first year. The killifish (Figure 1, bottom) origin of the dorsal fin in the male is Other common names ...... bull minnow, directly above or in advance of the gudgeon (Hildebrand and Schroeder origin of the anal fin (Rosen 1973). 1928); striped mummichog, banded The caudal fin is rounded in both killifish, killie, mummie (Ursin species (Hardy 1978). 1977) Class ...... Osteichthyes Fundulus heteroclitus: The snout is Order ...... Atheriniformes short, rounded, a little longer than family ...... Cyprinodontidae diameter of eye in side view, with 8 mandibular pores. A well-developed Geographic range ...... New Hampshire fleshy pouch is found at the anterior south to northeastern Florida (Figure base of the anal fin. Scales along 2). Mainly in saltwater, less often the lateral line number 31-35 (Rosen in brackish (Rosen 1973). Most 1973, but Hardy 1978 gives 31-39). authors do not record it from There are 11-12 dorsal fin rays freshwater (but see Hardy 1978). (Hildebrand and Schroeder 1928). (See also "Salinity.") There is no conspicuous silvery sheen on the sides; adult females have no dark spot on dorsal fin (Rosen 1973), and are brownish-green above, paler MORPHOLOGY/IDENTIFICATION AIDS below. Small females have 13-15 dark vertical bars (Hildebrand and The name "killifish" may be Schroeder 1928); adult females may be applied to members of three families: confused with those of F. con- Cyprinodontidae, Anablepidae, and fluentis (Rosen 1973). Adult males Poecilidae (Rosen 1973). Killifishes are dark green or olive above, yellow are soft-rayed, with no dorsal spine or beneath; have sides with about 15 spine preceding the anal fin. Most narrow silvery vertical bars and species are sexually dimorphic. Sexes numerous white or yellowish spots; are polymorphic with respect to pigment and may have a dark spot on posterior patterns and body and fin shape and 4-5 rays of dorsal fin. Sex-specific size. Rosen (1973) suggested fin-ray color patterns appear when fish are and scale counts to identify correctly 38-44 mm long (Hildebrand and males and females within species. His Schroeder 1928); until then, young paper contains a key to species of may be confused with adult female F. these three families. luciae (Rosen 1973). Adults are
3 commonly 51-102 mm long (Armstrong al. 1980). (See also "Movement and Child 1965). The largest Patterns" and "Food Habits.") specimen reported from Chesapeake Bay was 125 mm (Hildebrand and Schroeder Mummichogs are sold as bait in 1928). (Throughout this paper, sport fisheries for summer founder length measurements are as noted by or fluke (Paralichthys dentatus) and the original authors. Total length young bluefish or snappers (Pomatomus may be assumed where the authors did saltatrix) in New York and New Jersey not specify standard length.) (Perlmutter 1961), and for flounder in Delaware (M.H. Taylor, University of Fundulus majalis: The snout is long, Delaware, Newark, pers. comm.). pointed, and a little less than two eye diameters in profile (Rosen Mummichogs are the primary 1973). Overall body shape is slimmer predators in the Open Marsh Water than that of F. heteroclitus (Ursin Management mosquito control program 1977). There-are 13-15 dorsal fin currently being tested in New Jersey, rays (Hildebrand and Schroeder 1928); Delaware, and Maryland (Winner et al. and 32-37 scales in lateral series Although Gambusia (Hardy 1978). A conspicuous silvery ;~osquipt%ss)h Family Poeciliidae) is sheen appears on the sides of young more often u&d as a biological control and adults of both sexes (Rosen agent, the portion of the diet con- 1973); adults of both sexes have a sisting of mosquito larvae is higher black spot on the last rays of the studied Fundulus dorsal fin. Adult females have one &andis and c~r%~~~tis~' According to several dark longitudinal lines to Harrington and Harrington (1961), and one or more disrupted vertical the role of Gambusia affinis against bars near the base of the tail; adult Aedes mosquitoes has been overvalued females are olive above, white below. because of its reputation as a predator Adult males are dark olive on back, on freshwater anophelines; in the and their sides and belly are salmon control of saltmarsh mosquitoes, yellow, with 15-20 vertical black primary consideration should be given stripes (Hildebrand and Schr;c$;; to the local endemic cyprinodontiform 1928; Hardy 1978 gives fishes rather than any single species. stripes). Sex-specific color patterns appear when fish reach 38-51 mm (Hildebrand and Schroeder 1928); As a group, killifishes are used the young have bars similar to those in research in experimental studies of of ma1 es (Rosen 1973). This species embryology, genetics, physiology, is the largest member of the genus in endocrinology, cytology, and behavior. the study area; it frequently reaches Some of this research has important 152-178 mm. The record is 203 mm medical implications, such as the long (Ursin 1977). action of steroids in the regulation and reversal of sexual development, inheritance and development of REASON FOR INCLUSION IN SERIES cancerous tissues, and the genetics of histocompatability in tissue and organ Although not valued as commercial transplants (Rosen 1973). or sport fishes, both killifishes are important in food chains because of Mummichogs are the nondomesticated their distribution and abundance. (See fish most frequently used in research also "Predators.") Because of their (Rosen 1973), including such disparate importance in marsh food chains, studies as bioassay for water pollution mummichogs may be instrumental in (Isai et al. 1979), effects of weight- movement of organic material within and lessness in outer space (Hoffman et al. out of salt marsh ecosystems (Kneib et 1978), ion transport in tissues (Evans
4 1980), and cycling and biological Food availability may be the magnification of radioisotopes (Huver ultimate control for egg production in 1973). mummichogs. If feeding ceases, vitellogenesis (yolk formation) ceases and maturational stages are flushed from the reproductive tract; renewed feeding causes reappearance of LIFE HISTORY maturational stages (Wallace and Selman 1980). Fritz and Garside (1975) Reproductive Physiology reported that in an estuarine population, spawning mummichog females During the breeding season, males averaged 65 mm long and 243 ova; in a of mummichogs and striped killifish population in less brackish water assume a brighter coloration (Ursin (0.6-15.5 ppt), averages were 60 mm and 1977) and grow contact organs (Hardy 161 ova. The difference was attributed 1978). to lower food density in the less brackish area. Weisberg (1981) also Fundulus spp. are oviparous. The attributed the reduced female GSI ovary is single, and the number of ova observed in crowded experimental produced depends upon the size of the populations to insufficient food fish. The largest number of ripe ova availability. Taylor (in press), in a mummichog from Chesapeake Bay however, cautioned that additional counted by Hildebrand and Schroeder sources of stress may contribute to (1928) was 460, in a female 98 mm long; reduced fecundity in fish populations the maximum for a striped killifish was in nature. 540 (length not given). Hardy (1978) gave a range of 200-800 ova for Spawning Season and Periodicity mummichogs and 460-800 for striped killifish in the Mid-Atlantic region. The spawning seasons of mummichogs According to Taylor (in press), and striped killifish vary with production of 100-300 eggs per day for latitude. In Chesapeake Bay striped 3-5 days is not unusual for Delaware killifish spawn from April through mummichogs early in the spawning September; from New Jersey northward, season. However, even early in the the season lasts from June through spawning season in North Carolina, up August (Hardy 1978). Spawning of to 50% of the ova may be reabsorbed mummichogs usually begins in spring (Kneib and Stiven 1978). (March to May) and ends in later summer or early autumn (July to September) In male and female mummichogs, (Hardy 1978; Kneib, in press). Although the gonadosomatic index (GSI = gonad timing and duration of spawning seasons wt./body wt x 100) rises steeply in the differ, the water temperature range spring and declines steeply in the over which spawning occurs is similar autumn (Kneib and Stiven 1978). GSI (Taylor, in press). fluctuates in both sexes during the spawning season; mean GSI gradually Mummichogs may spawn eight or more declines as the season progresses times in a season. Each spawning peak (Taylor, in press). Although ripe ova may last five or more days and may be constantly available early in coincides with a high spring tide of the season (Wallace and Selman 1981>, the full or new moon (semilunar the number of eggs laid is always periodicity: Taylor and DiMichele greater during spring tides (Taylor et 1980). A circadian periodicity may be al. 1979). This semilunar cycle is superimposed on the semilunar rhythm in most pronounced late in the spawning some populations; maximal spawning then season in southern populations (Taylor, occurs when high spring tides are at in press). night (Taylor et al. 1979). Spawning
5 also occurs during the day (Hardy The pair quiver vigorously and shed 1978). their gametes several times in rapid succession (Keenleyside 1979). Spawning in separate genetic populations of mummichogs may be timed Spawning Site by different environmental stimuli (Wallace and Selman 1981). Spawning Mummichogs spawn - fresh rhythms may be timed by temperature brackish, and saltwater (Hgdy 1978): (Brummett 1966), tides, moonlight, and In a Delaware marsh, eggs were laid in salinity (Taylor et al. 1979). . Early clutches of 10-300, at levels only and late season peaks in spawning, such reached by high spring tides (1.2-1.4 m as Kneib and Stiven (1978) observed in above mean low water). Eggs are laid mummichogs, may be caused by a in empty shells of the ribbed mussel combination of moderate temperatures Geukensia demissa or inside the outer and shorter daylengths (Harrington dead leaves of smooth cordgrass 1959). Day and Taylor (1983) found (Spartina alterniflora) (Taylor- and that a photoreceptor other than the DiMichele 1983). EUCI deposition in pineal gland or retina of the eye shells has also been reported in influences seasonal reproduction in Virginia (Able and Castagna 1975), mummichogs. Female mummichogs respond North Carolina (Kneib and Stiven 1978), to photoperiod, but its effectiveness and Georgia (Taylor et al. 1981). has not been rigorously tested in males Other reported spawning sites include (Taylor, in press). In both sexes, low algal mats in Massachusetts (Taylor et temperature prevents and high al. 1981) and Connecticut (Pearcy and temperature permits gonadal development Richards 1962), shallow pits covered by (Taylor, in press). The semilunar the female (Newman 1907), and scattered cycle of oogenesis persists in the on the substrate (Keenleyside 1979). absence of lunar or tidal stimuli Selection of these different spawning (Taylor and Dimichele 1980). sites may reflect an inherited Schwassmann (1980) argued that we have preference or merely a response to no idea of the actual timing mechanism substrate availability (Taylor, in that entrains semilunar spawning in press). mummichogs. Striped killifish spawn in still, shallow water close to shore and, Spawning Behavior presumably, in small ponds. Active burying of eggs by females has been Rivalry is intense among breeding observed (Hardy 1978). male mummichogs, and size seems to be less important than nuptial coloration in defense of territories. Females with very ripe ova may solicit males by A key to eggs of cyprinodontids of turning on their sides to display their the Mid-Atlantic Bight appears in Hardy white bellies (Newman 1907). Adults (1978). Fertilized eggs of striped gather in shallow weedy areas, and'one killifish are spherical, 2.0-3.0 mm in or more males chase a female until she diameter, translucent yellow to amber, stops, at which time one male moves and slightly adhesive (Hardy 1978). beside her and pushes her against the vegetation (Keenleyside 1979). Newman Fertilized eggs of mummichogs are (1907) noted that courtship, consisting spherical, about 2 mm in diameter of tandem swimming, merges gradually (range: 1.5-2.5 mm), and transparent into spawning as the male's movements yellow to amber (Hardy 1978); dead or become progressively more excited. infertile eggs are opaque (Taylor, in Eventually the male clasps the female; press). Adhesive chorionic fibrils on their bodies form a shallow S-curve. eggs may be long and dense, short and 4 6 sparse, absent ’ (1978). Larvae of mummichogs may hatch populations':Hardy 1978; Mdrnin a~~r2\~~~ with the yolk already absorbed, if 1983). These fibrils may anchor eggs immersion of developed eggs has been to the substrate and/or retain moisture delayed (Taylor et al. 1977). At when eggs are stranded (Brummett and hatching mummichog embryos are 4-O-7.7 DuMont 1981). mm (mean 5.0 mm) long; striped killifish are 7.0-11.00 mm long (Hardy Mummichog eggs normally incubate 1978). In the laboratory at 20 OC, in air and are not submerged until the newly hatched mummichogs require 5.5 next spring tide after they are laid days to absorb the yolk. As the yolk (Taylor and DiMichele 1980). Eggs fail disappears, dorsal and ventral fins to develop if immersed for extended form, and the coordination of the lower periods in water with less than 1 ml/l jaw and operculum, undulating swimming, dissolved oxygen (DO) (Taylor, in and pectoral fin movements are press). Incubation of mummichog eggs perfected (Armstrong and Child 1965). in the field takes 7-8 days at 22-34 OC (Taylor et al. 1977); in the laboratory Larvae at 20 OC, hatching occurs in 10.5 days (Armstrong and Child 1965). In striped The larval stage lasts from killifish 50% of the eggs hatch by 41 yolk-sac absorption until the fish days at 16-20 OC, by 17 days at attains the characteristic shape of the 22-26 OC, and by 12 days at 28-32 'C species. During this stage the fin (Hardy 1978). rays and scales develop (Chuganova 1963). Striped killifish larvae are In mummichogs, hatching is 11.8-23.8 mm long. In mummichogs controlled by oxygen concentration of scales first appear above the pectoral the environment and hydration of the fin at about 12.5 mm, and are well egg (DiMichele and Taylor 1980); it can developed at 20.0 mm (Hardy 1978). be delayed 2 weeks or more if eggs are not immersed. Within seconds after Juveniles immersion of fully developed eggs, hatching begins. The egg swells, the Juveniles are also called fry or embryo's mouth opens, and heartbeat and young-of-the-year, and have fully respiration increase. Chorionase developed fins and a more or less (hatching enzyme) is released from distinct scale covering (Chuganova glands in the oral cavity and gills of 1963). The minimum described length of the embryo in response to hypoxia this stage in mummichogs is 25.0 mm; in ("respiratory distress syndrome" of striped killifish it is 26.0 mm (Hardy DiMichele and Taylor 1981). This 1978). For a detailed description of enzyme weakens the chorion so that body this stage for both species, see Hardy movements of the embryo can break it. (1978). Five minutes after immersion, the embryo begins to turn in the egg; the egg ruptures in 15-20 min (Taylor et Adults al. 1977). Female mummichogs are sexually Yolk-Sac Larvae mature at 38 mm and males at 32 mm; striped killifish females mature at The yolk-sac larval stage lasts 76 mm and males at 64 mm (Hildebrand from hatching until all yolk is and Schroeder 1928). Most members of absorbed, and may also be called the both species attain maturity during prolarva, free embryo (Chuganova 1963), their second year, although some or alevin (Tay and Garside 1978). A mummichogs may mature and spawn during key to yolk-sac larvae of fishes of the their first year (Hardy 1978; Kneib and Mid-Atlantic Bight appears in Hardy Stiven 1978). (See also
7 "MORPHOLOGY/IDENTIFICATION AIDS" and GROWTH CHARACTERISTICS "Population Size and Structure.") Length-Weight Relationship
Movement Patterns and Seasonal Length-weight equations have been Use of Habitats calculated for mummichogs:
The mummichog is "one of the most (1) log W = -5.232 + 3.172 log L stationary of fishes," according to where W = wt and L = Total Length Bigelow and Schroeder (1953). Breeding (Fritz and Garside 1975) migrations do not occur (Rosen 1973). (2) log Y = -2.06 + 3.27 log X Fish over 60 mm long maintain a summer X = Total Length (cm) and Y = wet home range of 36-38 m along one bank of wt (9) tidal creeks; however, some may move as (Meredith and Lotrich 1979) much as 375 m (Lotrich 1975). In some populations of mummichogs, the In winter, mummichogs in pools relation of weight to length may vary burrow 150-200 mm into the mud significantly through the year (Kneib (Chidester 1920; Hardy 1978). Others and Stiven 1978). Females are larger migrate to the mouth of the tidal than males of the same age, but males channel where they have been living; in are heavier than females of the same the subsequent spring they usually length (except during the spring return up the same channel (Butner and spawning peak). In the North Carolina Brattstrom 1960). Spring migration population studied, males averaged 251 begins when the water reaches 15 OC mg and females 245 mg. (Hardy 1978); feeding begins in New England marshes during the first week Seasonal and Annual Growth in April (Valiela et al. 1977). and Production
During the growing season, young Growth of first-season mummichogs mummichogs remain in the intertidal in mid-summer may average 5% of their zone for 6-8 weeks, inhabiting shallow total weight per day. In North pools on the marsh surface at low tide Carolina the major growing season is (Taylor et al. 1979). Juveniles of April to September. Both sexes grow 15-20 mm Standard Length (SL) begin to fastest during their first two growing move with the adults onto the marsh seasons. Females mature sexually at surface to feed (Kneib in press). To 30-35 mm SL in 5-6 months (Kneib and grow at a normal rate, mummichogs may Stiven 1978). need food from the marsh surface for at least part of their diet. Despite The age of mummichogs can be abundant food resources on the marsh determined by observing growth marks on surface, mummichogs may be food limited the otoliths after a fish has because tides limit foraging time there overwintered once. For a field (Weisberg and Lotrich 1980). population in Nova Scotia, Fritz and Garside (1975) observed the following Mummichogs maintain an endogeneous age-length correlations: circadian feeding pattern which, superimposed upon the tidal rhythm, AGE (yr): 1 2 3 4 allows maximal use of food resources LENGTH(mm): 35-50 51-74 68-83 78-95 available only at high tide (Weisberg et al. 1981). They feed primarily at Mummichogs reared in the laboratory at high tide during daylight (Weisberg and 13 OC have a maximum daily growth of Lotrich 1980; Reis and Dean 1981>, but 0.5% of their dry weight at a feeding also opportunistically (Weisberg et al. level of 6.0% of their body weight 1981). (Prinslow et al. 1974). There is no
8 stomach in mummichogs (i.e, no region Schroeder 1928). The density in summer of the gut secretes pepsin and of mummichogs longer than 40 mm may hydrochloric acid); digestion occurs range from 0.35 to 6.04/m2 in certain under alkaline conditions (Targett estuarine habitats (Kelso 1979). From 1979). Nevertheless, mummichogs have a 130,000 to 136,000 mummichogs longer higher than average assimilation than 30 mm have been reported along 3 (digestion) efficiency (87% on three km of a tidal creek in July, and from laboratory diets) compared to other 63,000 to 81,000 individuals a fishes (Weisberg and Lotrich 1982). September (Meredith and Lotrich 1979;: Metabolic costs account for 69% of ingested energy. Excretion exceeds Autumn, winter, and spring egested energy on some diets. populations of mummichogs are dominated by survivors of one growing season Gross growth efficiency (Kneib and Stiven 1978). During the (proportion of calories ingested that spawning season, synchronization of is incorporated into biomass x 100) has spawning and hatching with spring tides been calculated by several authors. results in pulses of larval and Larval mummichogs fed brine shrimp juvenile abundance on the marsh (Kneib nauplii were thought to be inefficient 1984). Later in the summer, the converters of food to biomass by Radtke population is overwhelmingly dominated and Dean (1979), but Kneib (1981), by young-of-the-year. In August, when using their data, calculated that gross all age classes are present, nearly 60% growth efficiency might be as high as of the population has not yet 25%-4l.%. Efficiency of adults was overwintered and less than 8% has determined to be 12% (Weisberg and completed two growing seasons (Kneib Lotrich 1982); this is at the low end and Stiven 1978). The number of fish of the normal range for carnivorous in the largest size class (longer than fishes. 70 mm) peaks in August and declines drastically by October (Meredith and Total production of mummichogs in Lotrich 1979). Migration, however, as salt marshes is among the highest well as mortality, may contribute to calculated for natural populations of this decline. In October and November, fishes: 160 kg dry wt/ha in New there are very few 3-year-olds (Valiela England (Valiela et al. 1977). et al. 1977). There is no evidence Meredith and Lotrich (1979) also that any members of this species calculated a high production (40.7 survive beyond their fourth growing g/m2/yr) and believed that local season (Kneib and Stiven 1978). conditions were the main reason that their figure varied from that of Mortality Valiela et al. (1977). Most mummichog eggs are fertile and survive to hatching; loss due to POPULATION DYNAMICS infertility, mortality, and predation is about 10% (Taylor, in press). Population Size and Structure Young-of-the-year may have an annual mortality of up to 99.5% and adults, Mummichogs are the most abundant 54% (Meredith and Lotrich 1979). member of their genus in some areas Female mortality increases dramatically (e.g., Beaufort, North Carolina: Huver after first reproduction (Kneib and 1973). Their name, in fact, is related Stiven 1978), perhaps because of to their abundance: "mummichog" is an detrimental after-effects of spawning Indian word meaning "going in crowds" (Meredith and Lotrich 1979). The (Nichols and Breder 1927). Schools may decline in the survivorship curve with number from a few fish to several age is not as marked as expected if hundred or more (Hildebrand and senescence were wholly responsible;
9 predation may be an important source of virens) isopods, ostracods, snails, mortality (Kneib and Stiven 1978). insects 1 bivalves, algae, fishes, fish During winter on the marsh, predation eggs, beetles and shrimps, and is low (Valiela et al. 1977); less than hymenopterans. In one North Carolina half as many large fish are lost marsh, the major prey of mummichogs between October and April as between were fiddler crabs, polychaetes, August and October (Meredith and tanaids, and other small crustaceans. Lotrich 1979). Fish less than 30 mm SL primarily ate small crustaceans (amphipods, tanaids, copepods); larger fish consumed crabs, ECOLOGICAL ROLE detritus, and algae more often (Kneib and Stiven 1978; Kneib et al. 1980). Feeding Behavior/Food Habits Although mummichogs may ingest large quantities of detritus while feeding on The protruding lower jaw and the surface of the substrate, detritus tilted mouth of cyprinodontids are is not a significant energy source for well-adapted to surface feeding (Eddy them because it is not assimilated 1957). Fundulus, however, does not (Prinslow et al. 1974). Mummichogs hesitate to feed in mid-water or on the cannot subsist on a diet of plant bottom (Huver 1973). Grass shrimp material or detritus (Katz 1975). (Palaemonetes pugio) swimming near the surface are often consumed, perhaps Mummichogs swallow their because they are silhouetted against prey intact, so mouth gape limits prey size. the light (Heck and Thoman 1981). Large mummichogs will also eat small Vision plays an important role in but larger are more feeding of mummichogs, but swallowing prey3 prey important in their diet (Vince et al. depends upon another sense, probably 1976). The numerical response of most olfactory (Hara 1971). Mummichogs are infaunal invertebrates to mummichog attracted to the amino acid predation depends on fish size more gamma-aminobutyric acid (GABA), but than fish density. Kneib and Stiven will not swallow substrate containing (1982) attributed this to very small GABA (Jonsson 1980). infauna (organisms in bottom sediments) not being readily available to larger Mummichogs use all potential food fish. sources: organisms in the water column, subtidal benthos, and intertidal benthos (Weisberg and Mummichog populations are large Lotrich 1982). Recent radioisotope enough to influence their prey's tracer studies have shown that 56% of distribution and abundance (Valiela et mummichogs' body carbon is derived from al. 1977), but few studies have algal (benthic and planktonic) food convincingly demonstrated this result chains and 44% from the Spartina food (Kneib 1984). Some prey species that chain (Hughes and Sherr 1983). may be regulated by mummichogs are the amphipod Gammarus palustris (Van Dolah Baker-Dittus (1978) decided that 1978), the pulmonate snail Melampus Fundulus uses all available food items bidentatus (Vince et al. 1976), and the except detritus. She found small soft-shelled clam Mya arenaria (Kelso crustaceans and polychaetes to be the 1979). Kneib (in press) points out most frequent food of both mummichogs that, by controlling smaller predators, and striped killifish. Fritz (1974) mummichogs may indirectly increase ranked food items found in the guts of densities of some infaunal marsh mummichogs 22-101 mm long. Copepods invertebrates. Also, predation by were the most common, followed in order larval and juvenile mummichogs, which by flies (mostly larvae and pupae), has previously been overlooked, may amphipods, polychaetes (mostly Nereis affect the patchy distribution patterns
10 of some small invertebrates (e.g., harpacticoid copepods) in salt marshes.
Predators
Small tidal marsh fishes, such as killifishes, are the major prey for wading birds, aerial searching birds, piscivorous ducks, and many predatory fishes (Peterson and Peterson 1979). These predators include herons, egrets, terns, gulls, striped bass, and bluefish (Valiela et al. 1977). The diet of nestling herons and egrets may contain up to 95% killifishes (including Gambusia), and up to 30% Fundulus spp. (Jenni 1969). Least and common terns eat Fundulus spp. in pools when the tide goes out (Butner and Brattstrom 1960). Selective predation by visual predators is suggested by increased mortality in male mummichogs Sheepshead minnow after they achieve sex-specific coloration (Kneib and Stiven 1978). Figure 3. Silhouettes of the three most common killifishes on the Mid-Atlantic From late August to early coast of the United States (from Rosen September in Delaware, American eels 1973). (Anguilla rostrata) preyed heavily on mummichogs (Lotrich 1975). Other fish that prey on Fundulus spp. are white Derch (Morone americana: Miller 1963). unvegetated tidal flats (Peterson and summer‘flounder (Paralichthys dentatusl Peterson 1979). All three species use Meredith and Lotrich 1979), and red the marsh surface at high tide, but drum (Sciaenops ocellata: Peterson and only mummichogs and sheepshead minnows Peterson 1979). Fundulus spp. are also do most of their feeding at this time. eaten by crabs (Libinia, Callinectes, These two species consume more Uca: Butner and Brattstrom 1960; epiphytic algae than do striped Eurytium limosum: Kneib in press): killifish, which eat more benthic Predation by blue crabs produces size- invertebrates. Sheepshead minnows are specific losses in experimental field herbivores, but mummichogs and striped populations of mummichogs (Kneib 1982). killifish seek invertebrates in the Another example of size-specific preda- algal mats (Werme 1981). tion is cannibalism of their own eggs by spawning mummichogs (Able and In the laboratory, sheepshead Castagna 1975). minnows chase mummichogs from their territories more often than they chase Competitors rainwater killifish (Lucania parva) or mosquitofish (Gambusia affinis). This Three species of killifishes mav be due to similar spawning (Figure 3)--mummichog, striped behavior. All four of these species killifish, and sheepshead minnow may occur together in nature, but in (Cyprinodon variegatus)--may occur ponds in the field, mummichogs tend to together in permanent ponds of the high remain in water deeper than 10 cm, and marsh (Nixon 1982), tidal creeks, the only briefly visit sheepshead minnow low marsh surface (Daiber 1982), and territories (Itzkowitz 1974).
11 Mummichogs, striped killifish, and Pathogenicity of parasites varies. Fundulus diaphanus were found in Oodinium cyprinodontiform is believed heterotypic schools in the intertidal to be a "symphoriont ectocommensal," region of a Maryland estuary by Baker- deriving most of its energy from Dittus (1978). Mummichogs were by far photosynthesis, but the other the most abundant in May and June, but dinoflagellate recorded from Fundulus striped killifish slightly dominated in SPP. 3 Amylodinium, seriously disrupts August and September. Diets of these gill epithelium (Lom and Lawler 1971). two species differed in only one Although trematode metacercarial cysts aspect: during July and August striped in their brains must cause compression killifish continued to eat many and tissue damage, the behavior of polychaetes, while mummichogs ate more infected mummichogs remains normal plant material. Greatest dietary (Abbott 1968). Because of its large overlap coincided with both the highest size relative to the host, larval density of all three fish species and Eustronglides spp. are a substantial the highest food density. nutritional drain; they also reduce host gonadal weight (Hirschfield et al. Parasites 1983). Larvae of Eustrongylides ignotus, when eaten in infected Parasites of Fundulus spp. are mummichogs, may cause high mortality in listed by Dillon (1966);fman (1967) nestling herons and egrets (Wiese et listed additional species and noted al. 1977). larval stages. Parasites of mummichogs and striped killifish include dinoflagellates, sporozoans, monogenetic flukes, digenetic flukes, ENVIRONMENTAL REQUIREMENTS/TOLERANCES tapeworms, roundworms, spiny-headed worms and copepods. Mummichogs are Temperature also susceptible to lymphocystis (caused by a virus) and Chondrococcus Temperate marine fishes do not columnaris, a myxobacterium infecting normally survive water temperatures the skin, gills, and fins (Sinderman greater than 34 OC (de Silva 1969). 1970). However, several species of Fundulus can recover from exposures to 40-42 OC water (Altman and Dittmer 1966). In Since 1967 additional parasites laboratory experiments, mummichogs and have been recorded from mummichogs, striped killifish died of heat shock in including Eimeria funduli, a sporozoan 63 min at 34 OC, 28 min at 36 OC, 9 min unusual in that it requires grass at 37 OC, and 2 min at 42 OC (Orr shrimp (Palaemonetes pugioj as a second 1955). host (Upton and Duszynski 1982); Trypanoplasma bullocki, a flagellate Mummichogs are eurythermal (de vectored by a leech in Chesapeake Bay Silva 1969). In Delaware salt marshes, (Burreson and Zwerner 1980); and mummichogs experience a temperature Dichelne bullocki, a roundworm- whose fluctuation of 6-35 OC (Schmelz 1964). larvae 1 ive in the anterior. and adults In Maine salt marshes, summer tidal the posterior, intestine (Kuzia cycles expose mummichogs to rapid ;:78). temperature changes from 15 to 30 OC. Swimming ability is maintained in Add itional parasites reported from nature during temperature fluctuations both species of ‘Fundulus are Swingelus which would substantially impair a monogenetic trematode (Billeter contractile function in other fishes. and Cryptobia bullocki a In the laboratory, low ATPase activity bifla:ellate kinetoDlastid found ii the at very cold temperatures supports bloodd(Nigrelli et al. 1975). field observations of a relatively
12 torpid state for overwintering unstable at 30 OC or above (DiMichele mummichogs (Side11 et al. 1983). et al. 1981).
Although assimilation (digestion) efficiency of mummichogs varies significantly with temperature in the Salinity laboratory, this is probably of little ecological significance. Maximum Although mummichogs are efficiency occured from 13 to 29 OC in physiologically euryhaline, Fritz and the laboratory; at lower temperatures Garside (1974) considered salinity fish simply took longer to digest their preference to be the most important food (Targett 1979). In the environmental factor influencing their laboratory, rate of feeding of young distribution in Nova Scotia. mummichogs depended on temperature and Laboratory studies supported this increased rapidly above 24 OC (Radtke conclusion by showing that mummichogs and Dean 1979). strongly preferred salinity of 20 ppt over 8 ppt. Oxygen consumption is relatively independent of acclimation temperature Striped killifish rarely if ever from 15 to 25 OC (Moerland and Side11 enter freshwater (Rosen 1973; Robins et 1981). Targett (1979) reported a al. 1980, but see Hardy 1978). In the similar relationship for the Chesapeake Bay area mummichogs are respiratory metabolic rate: rarely taken in full saltwater and are temperature independence from 13 to found more often in freshwater than are 29 OC and temperature-sensitive striped killifish, although the reduction from 5 to 13 OC. habitats of these species overlap ?L (Hildebrand and Schroeder 1928). In Mummichogs spawn in water from North Carolina, striped killifish also 16.5 to 25 OC (Hardy 1978). Eggs tend to occur in higher salinities, develop at temperatures of 12-27 OC while mummichogs tolerate lower with less than a 2% incidence of salinities (Peterson and Peterson abnormality (Smith 1982). In the 1979). These studies support the laboratory, mortality was low (7%) to long-held notion of the greater ability moderate (29%) in striped killifish of mummichogs to live in a range of embryos transferred from an inter- salinities, perhaps selected in a given mediate, fluctuating temperature regime area to avoid competition with (22-26 "C) to higher (28-32 "C) or congeners. lower (16-20 "C) fluctuating tempera- tures (Fahy 1976). The number of Mummichogs are especially tolerant vertebrae and dorsal fin rays developed of abrupt salinity changes. They adapt by embryos was inversely correlated so quickly that prior salinity with temperature, and was influenced acclimation has no effect on laboratory more by cold water than by warm (Fahy testing of response to salinity 1979). Thermal effluents from electric (Garside and Chin-Yuen-Kee 1972). power plants caused an elevated frequency of vertebral abnormalities in The ability of ova from northern mummichog embryos (Mitton and Koehn and southern populations of mummichogs 1976). to be fertilized at various salinities was examined by Bush and Weis (1983). The median time to hatching is There was no difference in the ability inversely related to water temperature of sperm to fertilize ova at 30 ppt. (Tay and Garside 1975). Although However, at 15 ppt there was a hatching is not affected by the difference, which may mean that egg normally encountered range of response to salinity varies temperatures, the hatching enzyme is geographically.
13 Mummichog larvae survive in water a solution of 15% water soluble between 0.39 ppt (fresh, by definition) fraction (WSF) of No. 2 fuel oil, which and IO0 PPt, although growth is contains 0.28 ppm naphthalenes, was retarded at salinities greater than only mildly toxic to mummichog embryos. that of seawater (32-33 ppt). Larvae Concentrations of 25% WSF (0.47 ppm are active and feed in water under 1 naphthalenes) were required to attain PPt, but do not survive more than 11 >50% mortality. However, under days in tap water (Joseph and Saksena suboptimal conditions, especially 1966). When mummichog embryos are extreme temperatures, even the 15% incubated in salinities of (0.5 ppt to exposure resulted in ~50% mortality 60 ppt, those in 20 ppt are always the (Linden et al. 1979). Adult mummichogs longest and those in freshwater are under thermal or osmotic stress are always the shortest (Tay and Garside also more sensitive to the lethal 1978). effect of emulsions of fuel oil (Butler et al. 1982). Temperature-Salinity Interactions Teratogenicity (ability to cause Salinity affects response to water birth defects) of a pollutant may also temperature in mummichogs. For be higher under suboptimal salinities example, the role of increased serum (Weis et al. 1979). For example, glucose in low temperature survival is teratogenicity of methylmercury is affected by salinity (Leach and Taylor higher at salinities of 10 ppt than at 1977). The upper lethal temperature is 30 PPt, and at temperatures of 16 OC highest (34 "C) in an isosmotic (14 than of 25 OC (Weis et al. 1979). ppt) medium. The lethal temperature is Sensitivity to radiation also depends depressed 6 OC in freshwater (0 ppt), on temperature and salinity (Baptist et but only 2.5 OC in seawater (32 ppt) al. 1972). In both mummichogs and (Garside and Chin-Yuen-Kee 1972). In striped killifish, salinity plays an the laboratory, mummichogs preferred important role in copper uptake which 24 OC in seawater but 22 OC in differs for each species and life cycle freshwater. Stress imposed by unusual stage (Bennett and Dooley 1982). salinity does not allow complete expression of thermal acclimation Other Environmental Factors (Garside and Morrison 1977). Low dissolved oxygen is a necessary hatching stimulus for Contaminants mummichog eggs (DiMichele and Taylor 1980). Adult mummichogs survive Mummichogs are considered overnight in anoxic tidal waters by particularly stress- and pollution- gaping at the surface. Presumably tolerant (Huver 1973), although they splashing oxygenates the surface water are more susceptible to chlorine and that reaches the gills (M.H. Taylor, chloramine than are winter flounder University of Delaware, Newark; pers. (Pseudopleuronectes americanus) and comm.). - scup (Stenotomus chrysops) (Capuzzo et al. 1977). Response to pollutants may Although found on many substrates, depend upon temperature, salinity, and mummichogs prefer mud (Hildebra;ld97;;d presence or absence of certain Schroeder 1928). Fritz chemicals in the water. The same however, was unable to detect A concentration of a pollutant that is substrate preference. Mummichoqs seem only mildly toxic to fish under optimal to prefer areas with grass (Spartina environmental conditions may be lethal patens) over areas without (mud to fish under thermal or osmotic substrate), possibly because vegetation stress. For example, under optimal allows them to evade predators conditions of temperature and salinity, (confirmed in the laboratory by Ordzie
14 1978). Striped killifish tend to occur than two fathoms (Bigelow and Schroeder over sandy sediments more often than do 1953). Striped killifish may be found mummichogs, and are by far the most in water only a few centimeters in important killifish on the unvegetated depth. This species is concentrated intertidal flats of North Carolina along the shoreline during flood tides, (Peterson and Peterson 1979). In the and, if washed ashore, can reenter the laboratory, both species respond more water (as can the mummichog) by a strongly to substrate and cover than to series of jumps (Hildebrand and current (Rosen 1973). Schroeder 1928; Rosen 1973). Mummichogs and striped killifish are The maximum depth at which positively rheotactic (face into a mummichogs are found is seldom more water current) (Rosen 1973).
15
LITERATURE CITED
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Able, K.W., and M. Castagna. 1975. Brummett, A.R. 1966. Observations on Aspects of an undescribed the eggs and breeding season of reproductive behavior in Fundulus Fundulus heteroclitus at Beaufort, heteroclitus (Pisces: Cyprinodon- North Carolina. Copeia 1966:616-620. tidae) from Virginia. Chesapeake Sci. 16(4):282-284. Brummett, A.R., and J.N. DuMont. 1981. A comparison of chorions from eggs of Altman, P-L., and D.S. Dittmer, eds. northern and southern populations of 1966. Environmental biology. Fundulus heteroclitus. Copeia Federation of American Societies for 1981(3):607-614. Experimental Biology, Bethesda, Md. 694 pp. Burreson, E.M., and D.E. Zwerner. 1980., Host, range, life cycle, and Armstrong, P.B., and J.S. Child. 1965. pathology of Trypanoplasma bullocki Stages in the normal development of in lower Chesapeake Bay fishes. J. Fundulus heteroclitus. Bioi. Bull. Protozool. 27(3):23A-24A. (Woods Hole) 128:143-168. Bush, C..P., and J.S. Weis. 1983. Baker-Dittus, A.M. 1978. Foraging Effects of salinity on fertilization patterns of three sympatric success in two populations of killifish. Copeia 1978(3):383-389. Fundulus heteroclitus. Biol. Bull. (Woods Hole) 164(3):406-417. Baptist, J.P., T.R. Rice, F.A. Cross, and T.W. Duke. 1972. Potential Butler, R.G., W. Trivelpiece, and D.S. hazards from radioactive pollution of Miller. 1982. The effects of oil the estuary. Mar. Pollut. Sea Life dispersant and emulsions on the FA0:272-276. survival and behavior of an estuarine teleost and an intertidal amphipod. Bennett, R.O., and J.K. Dooley. 1982. Environ. Res. 27(2):266-276. Copper uptake bv two svmpatric species 'of kiliifish, iundulus Butner, A., and B.H. Brattstrom. 1960. heteroclitus and Fundulus majalis. Local movement in Menidia and J. Fish Biol. 21(4):381-398. Fundulus. Copeia 1960(2):139-141.
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19 Joseph, E.B., and V.P. Saksena. 1966. Kneib, R.T., and A.E. Stiven. 1982. Determination of salinity tolerances Benthic invertebrate responses to in mummichog (Fundulus heteroclitus) size and density manipulations of the larvae obtained from hormone-induced common mummichog, Fundulus spawning. Chesapeake Sci. 7(4): heteroclitus, e intertidal 193-197. saltmarsh. Ecolb;y S&:1518-1532.
Katz, L.M. 1975. Laboratory studies Kneib, R.T., A.E. Stiven, and E.B. on diet, growth, and energy Haines. 1980. Stable carbon isotope requirements of Fundulus heteroclitus ratios in Fundulus heteroclitus (Linnaeus). Ph.D. Dissertation. muscle tissue and gut contents from a University of Delaware, Newark. 81 North Carolina Spartina alterniflora PP. marsh. J. EXD. Mar. Biol. Ecol. 46(1):89-98. ’ Keenleyside, M.H.A. 1979. Diversity and adaptation in fish behavior. Kuzia, E.J. 1978. Studies on the life Springer-Verlag, New York. 208 pp. history, season1 periodicity and histopathology of Dichelyne bullocki Kelso, W.E. 1979. Predation on Stromberg and Crites (Nematoda: soft-shell clams, Mya arenaria, by Cucullanidae), a parasite of Fundulus the common mummichog, Fundulus heteroclitus (L.). Ph.D. heteroclitus. Estuaries 20:249- Dissertation. University of New 254. Hampshire, Durham. 104 pp.
Kneib, R.T. 1981. Re-analysis of Leach, G.J., and M.H. Taylor. 1977. conversion efficiencies for larval Seasonal measurements of serum Fundulus heteroclitus. Mar. Biol. glucose and serum cortisol in a (Berl.)63(2):213-215. natural population of Fundulus heteroclitus. Comp. Biochem. Physiol. 56(2):217-223. Kneib, R.T. 1982. The effects of predation by wading birds (Ardeidae) and blue crabs (Callinectes sapidus) Linden, O., R. Laughlin, Jr., J.R. on the population size structure of Sharp, and J.M. Neff. 1979. The the common mummichoa (Fundulus combined effect of salinity, heteroclitus). Estuarine . Coastal temperature and oil on the growth Shelf Sci. 14(2):159-166. pattern of embryos of the killifish (Fundulus heteroclitus). Mar. Environ. Res. 3(2):129-144. Kneib, R.T. 1984. Patterns in the utilization of the intertidal Lom, J., and A.R. Lawler. 1971. Mode saltmarsh bv larvae and iuveniles of of attachment and relation to host Fundulus h&eroclitus (Ljnnaeus) and tissue in two dinoflagellates from Fundulus luciae (Baird). J. Exp. gills of cyprinodonts of Virginia. Mar. Biol. Ecol. 83:41-51. J. Protozool. 18(Suppl.):43.
Kneib, R.T. In press. The roie of Lotrich, V.A. 1975. Summer home range Fundulus heteroclitus in saltmarsh and movements of Fundulus trophic dynamics. Am. 2001. heteroclitus (Pisces: Cyprinodon- tidae) in a tidal creek. Ecology Kneib, R.T., and A.E. Stiven. 1978. 56(1):191-198. Growth, reproduction, and feeding of Fundulus heteroclitus (L.) on a North Massman, W.H. 1954. Marine fishes in Carolina saltmarsh. J. Exp. Mar. fresh and brackish waters of Virginia Biol. Ecol. 31(2):121-140. rivers. Ecology 35(1):75-78.
20 Meredith, W.H., and V.A. Lotrich. Serv. Biol. Serv. Program. 1979. Production dynamics of a tidal FWS/OBS-81/55. 70 pp. creek population of Fundulus heteroclitus (Linnaeus). Estuarine Northcutt, R.G., and R.E. Davis, eds. Coastal Mar. Sci. 8(2):99-118. 1983. Fish neurobiology. Vol. 1. University of Michigan Press, Ann Miller, L.W. 1963. Growth, Arbor. 414 pp. reproduction and food habits of the white perch, Roccus americanus (Gmelin) in the Delaware River Ordzie, C.J. 1978. Habitat selection Estuary. M.S. Thesis. University of by Fundulus heteroclitus. Ph.D. Delaware, Newark. 62 pp. Dissertation. University of Rhode Island, Kingston. 114 pp: Mitton, J.B., and R.K. Koehn. 1976. Morphological adaptation to thermal Orr, P.R. 1955. Heat death: I. stress in a marine fish, Fundulus Time-temperature relationships in heteroclitus. Biol. Bull. (Woods marine animals. Physiol. Zool. Hole) 151(3):548-559. 28(4): 290-294.
Moerland, T.S., and B.D. Sidell. 1981. Pearcy, W.G., and S.W. Richards. 1962. Characterization of metabolic carbon Distribution and ecology of fishes of flow in hepatocytes isolated from the Mystic River Estuary, thermally acclimated killifish Connecticut. Ecology 43~248-259. (Fundulus heteroclitus). Physiol. Zool.54(3) :379-389. Perlmutter, A. 1961. Guide to marine fishes. New York University Press, Morin, R.P., and K.W. Able. 1983. New York. 431 pp. Patterns of geographic variation in the egg morphology of the fundulid Peterson, C.H., and N.M. Peterson. fish Fundulus heteroclitus. Copeia 1979. The ecology of intertidal 1983(3):726-740. flats of North Carolina: a community profile. U.S. Fish Wildl. Serv. Newman, H.H. 1907. Spawning behavior Biol. Serv. Program FWS/OBS-79/39. and sexual dimorphism in Fundulus 73 PP. heteroclitus and allied fish. Biol. Bull. (Woods Hole) 12:314-345. Prinslow, T.E., I. Valiela, and J.M. Teal. 1974. The effect of detritus and ration size on the growth of Nichols, J.T., and C.M. Breder, Jr. Fundulus heteroclitus. J. Exp. Mar. 1927. The marine fishes of New York Biol. Ecol. 16(1):1-10. and southern New England. Zoologica 9(1):1-192. Radtke, R.L., and J.M. Dean. 1979. Feeding, conversion efficiencies, and Nigrelli, R.F., K.S. Pokorny, and G.D. growth-of larval mummichogs, Fundulus Ruggieri. 1975. Studies heteroclitus. Mar. Biol. (Berlt) parasitic kinetoplastids. Part ? 55(3):231-237. Occurrence of a biflagellate kinetoplastid in the blood of Reis, R.R., and J.M. Dean. 1981. Opsansus tau (toadfish) transmitted Temporal variations in the by the lea Piscicola funduli. J. utilization of an intertidal creek bv Protozool. 220:43A. the bay anchovy, Anchoa mitchilli: Estuaries 4(1):16-23. Nixon, S.W. 1982. The ecology of New England high salt marshes: Robins, C.R., R.M. Baily, C.E. Bond, community profile. U.S. Fish Wildly J.R. Brooker, E.A. Lachner, R.N. Lea,
21 and W.B. Scott. 1980. A list of the (Cyprinodontidae), to continuous common and scientific names of fishes incubation in various combinations of from the United States and Canada, temperature and salinity. Canad. J. 4th ed. Am. Fish. Sot. Spec. Publ. Zool. 53(7):920-933. 12. Bethesda, Md. 174 pp. Tay, K.L., and E.T. Garside. 1978. Rosen, D.E. 1973. Suborder Compensatory embryogenic responses in Cyprinodontoidei. Pages 229-262 in osmoregulative structures of D.M. Cohen, N.B. Marshall, A.K mummichog Fundulus heteroclitus L. Ebeling, D.E. Rosen, T. Iwamoto, P. incubated at constant temperature and Sonoda, S.B. McDowell, W.H. Weed III, various constant levels of salinity. and L.P. Woods. Fishes of the Canad. J. Zool. 56(4):613-623. western North Atlantic. Sears Found. Mar. Res. Mem. 1, Pt. 6, Yale Taylor, M.H. In press. Environmental University. and endocrine influences on repro- duction of Fundulus heteroclitus. Schmelz, G.W. 1964. A natural history Am. Zool. of the mummichog, Fundulus heteroclitus (Linnaeus), in Canary Taylor, M-H., and L. DiMichele. 1983. Creek marsh. M.S. Thesis. Spawning site utilization in a University of Delaware, Newark. 65 Delaware population of Fundulus PP. Cyprinodon- 1983(3):719-725. Schwassmann, H.O. 1980. Biological rhythms: their adaptive Taylor, M-H., and L. DiMichele. 1980. significance. Pages 613-630 in M.A. Ovarian changes during the lunar Ali, ed. Environmental physio%gy of spawning cycle Fundulus fishes. Plenum Press, New York. heteroclitus. Copeiaof1980(1):118- 125. Sidell, B.D., I.A. Johnston, T.S. Moerland, and G. Goldspink. 1983. Taylor, M.H., L. DiMichele, R.T. Kneib, The eurythermal myofibrillar protein and S. Bradford. 1981. Comparison complex of the mummichog Fundulus of reproductive strategies in Georgia heteroclitus: adaptation to a and Massachusetts populations of fluctuating thermal environment. 3. Fundulus heteroclitus. Am. Zool. Comp. Physiol. 153(2):167-174. 21(4):921.
Sinderman, C.J. 1970. Principal Taylor, M.H., L. DiMichele, and G.J. diseases of marine fish and Leach. 1977. Egg stranding in the shellfish. Academic Press, New York. life cycle of the mummichog Fundulus 369 pp. heteroclitus. Copeia 1977(2):397- 399. Smith, L.S. 1982. Introduction to fish physiology. T.F.H. Publ., Inc., Taylor, M.H., G.J. Leach, L. DiMichele, Neptune, N.J. 352 pp. W.H. Levitan, and W.F. Jacob. 1979. Lunar spawnina cycle in the Targett, T.E. 1979. The effect of mummichog; Fundulu; heteroclitus temperature and body size on (Pisces: Cyprinodontidae). Copeia digestive efficiency in Fundulus 1979(2):291-297. heteroclitus. J. Exp. Mar. Biol. Ecol. 38(2):179-186. Teal, J.M., and K.A. Burns. 1979. The West Falmouth oil spill: hydro- Tay, K.L., and E.T. Garside. 1975. carbons in the saltmarsh ecosystem. Some embryogenic responses of the Estuarine Coastal Mar. Sci. 8(4): mummichog, Fundulus heteroclitus (L.) 349-360.
22 Upton, S.J., and S.W. Duszynski. 1982. heteroclitus). Pages 64-70 in R. Development of Eimeria funduli in Lasker and K. Sherman, eds. 3981. Fundulus heteroclitus. J. Protozool. The early life history of fish: 29(1):66-71. recent studies. ICES Symp. Early Life Hist. Fish. Woods Hole, Mass. Ursin, M.J. 1977. A guide to the fishes of the temperate Atlantic Weisberg, S.B. 1981. Food coast. E.P. Dutton, New York. 269 availability and utilization by the PP. mummichog, Fundulus heteroclitus (L.). Ph.D. Dissertation. Valiela, I., J.E. Wright, J.M. Teal, University of Delaware, Newark. and S.B. Volkman. 1977. Growth, production and enerqy transformations Weisberg, S.B., and V.A. Lotrich. in the saltmarsh kiilifish, Fundulus 1980. Food limitation of the heteroclitus. Mar. Biol. '(Berl.) mummichog Fundulus heteroclitus in a 40(2):135-144. Delaware saltmarsh. Am. Zool. 20(4): 880. Van Dolah, R.F. 1978. Factors regulating the distribution and Weisberg, S.B., and V.A. Lotrich. population dynamics of the amphipod 1982. The importance of an Gammarus palustris in an intertidal infrequently flooded intertidal marsh saltmarsh community. Ecol. Monogr. surface as an enerqy source for the 48(2):191-218. mummichog Fundulus -heteroclitus: an experimental approach. Mar. Biol. Vince, S., I. Valiela, N. Backus, and (Berl.) 66(3):307-310. J.M. Teal. 1976. Predation by the saltmarsh killifish Fundulus Weisberg, S.B., R. Whalen, and V.A. heteroclitus (L.) in relation to prey Lotrich. 1981. Tidal and diurnal size and habitat structure: influence on food consumption of a consequences for prey distribution saltmarsh killifish, Fundulus and abundance. J. Exp. Mar. Biol. heteroclitus. Mar. Biol. (Berl.) Ecol. 23(3):255-266. 61(2-3):243-246.
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Wallace, R.A., and K. Selman. 1981. Wiese, J.H., W.R. Davidson, and V.F. Reproductive activity of Fundulus Nettles. 1977. Large-scale heteroclitus females from Woods Hole, mortality of nestling ardeids caused Massachusetts, as compared with more by nematode infection. J. Wildl. southern locations. Copeia Dis. 13(4):376-382. 1981:212-215. Winner, S.M., J.R. Day, J.A. Bartley, Weis, J.S., P. Weis, and J.L. Ricci. and M.H. Taylor. In press. 1979. Effects of cadmium, zinc, Reproduction in Fundulus heteroclitus salinity, and temperature on the populations in OMWM habitats. Proc. teratogenicity of methylmercury to 71st Annu. Mtg. N. J. Mosq. Contr. the killifish (Fundulus Assoc.
23
REFORT WCUMENTAflON I 1. ?C’Orn.-- PAGE i Biological Report 82(11.40)* ' L_wdtiW Species Profiles: Life Histories and Environmental Require- ments of Coastal Fishes and Invertebrates (Mid-Atlantic)-- Mummichoq and StriDed Killifish 7.Mhar(9l Barbara J. Abraham i.hr(orml~o~lnlXmion Warm lnd IddrnB
0 l2. Soomodw Ocgnlmtbn Name and Addmu 1%T~dR*oa+tL~rlod~nc National Coastal Ecosystems Team U.S. Army Corps of Engineers Fish and Wildlife Service Waterways Experiment Station U.S. Dep. of the Interior P.O. Box 631 14. Washington, DC 20240 Vicksburg, MS 39180
c 1S. Sap lmwtwy Notes *U.S. Army Corps of Engineers Report No. TR EL-82-4
Species profiles are literature summaries of the taxonomy, morphology, range, life history, and environmental requirements of coastal aquatic species. They are designed to assist in environmental impact assessment. Mummichogs and striped killifish are not commercial or sport fish; however, their distribution and abundance make them important in nearshore and estuarine food webs. These are two of the three most common killifishes on the Mid-Atlantic coast; summer density of mummichogs in estuaries may be as high as six per square meter. Most members of both species attain sexual maturity and spawn during their second year. Mummichogs have a semilunar spawning periodicity during the spawning season; eggs incubate in air and are not submerged until the next spring tide after they are laid. Young mummichogs remain on the marsh for 6-8 weeks, then begin to move off with the tides, with the adults. Mummichogs and striped killifish are euryphagous predators. Both species are tolerant of temperature and salinity fluctuations. Mummichogs are rarely taken in full seawater; striped killifish rarely, if ever, enter freshwater. Between these extremes, the habitats of the two species overlap.
17. Docummt Analysis l . Descriptors Parasites Feeding Fishes Salinity Growth Tidal marshes Estuaries tdmtw n/olmn.End*dT*rmr r~mm,CliOQ Life historv Predators Fundulus-heteroclitus Spawning u Competitors Striped killifish Temperature requirements Fundulus majalis Salinity requirements
L COSATI Field/Group h Av~lbbltity Statmmnt 19. kcunty CIJSS (This Repott) 21. No. ol PIICS Unlimited Unclassified 73 -- 20.kcurity Class (This Pap) 22 Pricm lassified OPTIONAL FORU 272 W-7 (Corlnerly f4119-31) oapmtmwlt of commwa
species profiles : 25234 ., ,_. . . . Abraham, Barbara J, ULl!JS ,sb3 no.82
.es~wch Library .J. S. Departrrrent of the Interlc, Ziatinnal Biological Survey Genre Cj?ZQter
I 1 / \ 7 -- - \ - \ I \ \ I
Se&s. Washmgton. DC - Eastern Energy and Land “se Team
Locarions of Regtonal 0th~~
REGION 1 REGION 2 REGION 3 Regional Director Regional Director Regional Director U.S. Fish and Wildlife Service U.S. Fish and Wildlife Service U.S. Fish and Wildlife Service Lloyd Five Hundred Building, Suite 1692 P.O. Box 1306 Federal Building, Fort Snelling 500 N.E. Multnomah Street Albuquerque, New Mexico 87 103 Twin Cities, Minnksota 55 I I I Portland, Oregon 97232
REGION 4 REGION 5 REGION 6 Regional Director Regional Director Regional Director U.S. Fish and Wildlife Service U.S. Fish and Wildlife Service U.S. Fish and Wildlife Service Richard B. Russell Building One Gateway Center P.O. Box 25486 75 Spring Street, S.W. Newton Corner, Massachusetts 02158 Denver Federal Center Atlanta, Georgia 30303 Denver. Colorado 80225
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As the Nation’s principal conservation agency, the Department of the Interior has respon- sibility for most of our nationally owned public lands and natural resources. This includes fostering the wisest use of our land and water resources, protecting our fish and wildlife, preserving theenvironmental and cultural values of our national parks and historical places, and providing for the enjoyment of life through outdoor recreation. The Department as- sesses our energy and mineral resources and works to assure that their development is in the best interests of all our people. The Department also has a major responsibility for American Indian reservation communities and for people who live in island territories under U.S. administration.