REFERENCE COW 

Do Not Remove from the Librarl-1 

U. S. Fish and Wildlife Serv;~~ L I .- I \A we. L C.--L *.,,- -. . ,. 7 - 

Biological Report 82(11.40) 13U'1u1 Iu1 700 Cajun Dome Boule.,ar-1 TR EL-82-4 

June 1985 

Lafayette, Louisiana 70536 

Species Profiles: Life Histories and 

Environmental Requirements of Coastal Fishes 

and Invertebrates (Mid-Atlantic) 

MUMMICHOG AND STRIPED KILLIFISH 

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, pub1 ished by the Research 

and Development branch of the U.S. Fish and Wildlife Service, replaces 

the "FWS/OBS1' series published from 1976 to September 1984. The Biological 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 

Sl i dell , 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 Wildl ife Service. 1983-19-. Species profiles: 1 ife histories 

and environmental requirements of coastal 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: 1 i fe 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 envi ronmental changes caused by coastal development. Each 

profile has sections on taxonomy, 1 ife history, ecological role, environmental 

requirements, and economic importance, if appl icable. 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 foll owing addresses. 

Information Transfer Special i st 

National Coastal Ecosystems Team 

U.S. Fish and Wildlife Service 

NASA-Sl idel 1 Computer Complex 

1010 Gause Boulevard 

Slidell, LA 70458 

U. S. Army Engineer Waterways Experiment Station 

Attention: WESER-C 

Post Office Box 631 

Vicksburg, MS 39180 
CONVERSION TABLES 

Metric to U .S . Customary 

Mu1 tiply 

mil 1 imeters (mn) 

centimeters ( an) 

meters (m) 

ki 1 meters ( km) 

2 

square meters (m ) 10.76 

square ki 1 meters ( km2) 0.3861 

hectares (ha) 2.471 

liters (1) 

cubic meters (m3) 

cubic meters 

To Obtain 

Inches 

Inches 

feet 

mil es 

square feet 

square miles 

acres 

gal 1 ons 

cubic feet 

acre-feet 

mill igrams (mg) 0.00003527 ounces 

grams (g) 0.03527 ounces 

kilograms (k ) 2.205 pounds 

metric tons ft) 2205.0 pounds 

metric tons 1.102 short tons 

ki 1 ocal ories (kcal ) 3.968 British thermal units 

Cel sius degrees 1.8("C) + 32 Fahrenheit degrees 

U.S. Customary to Metric 

inches 25.40 

inches 2.54 

feet (ft) 0.3048 

fa thoms 1.829 

miles (mi) 1.609 

nautical miles (mi) 1.852 

square feet (ft2) 

acres 2 

square miles (mi ) 

gal 1 ons (gal ) 

cubic feet (ft3) 

acre-feet 

ounces (02) 23.35 

pounds (lb) 0.4536 

short tons (ton) 0.9072 

British thermal units (Biu) 0.2520 

mil 1 imeters 

centimeters 

meters 

meters 

kilometers 

ki 1 meters 

square meters 

hectares 

square kilometers 

1 i ters 

cubic meters 

cubic meters 

grams 

ki 1 ograms 

metric tons 

ki 1 ocal ories 

Fahrenheit degrees 0.5556("F - 32) Cel sius degrees 
CONTENTS 

Page 

PREFACE .................................................................. iii 

CONVERSION TABLE ........................................................... iv 

ACKNOWLEDGMENTS ............................................................ vi 

NOMENCLATURE/TAXONOMY/RANGE ................................................ 1 

MORPHOLOGY/IDENTIFICATION AIDS ............................................. 3 

REASON FOR INCLUSION IN SERIES ............................................ . 4 

LIFE HISTORY ............................................................... 5 

Reproductive Physiology .................................................. 5 

Spawning Season and Periodicity .......................................... 5 

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 ........................... 8 

GROWTH CHARACTERISTICS ..................................................... 8 

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 .............................................................. 12 

Salinity ................................................................. 13 

Temperature-Salinity Interactions ........................................ 14 

Contaminants ....................................................... 14 

Other Environmental Factors .............................................. 14 

LITERATURE CITED ........................................................... 
ACKNOWLEDGMENTS 

I am grateful to M.H. Taylor, University of Delaware, and two anonymous 

reviewers who commented on an early draft of the manuscript. 
Adult male killifish 

Adult female killifish 

Figure 1. Mummichog and striped ki 11 ifish . 

MLIMMICHOG AND STRIPED KILLIFISH 

NOMENC LATURE/TAXONOMY/RANGE Class .................... Osteichthyes 

Order .................. Atheriniformes 

Scientific mme ... Fundul us hetero- Family ................ Cypri nodontidae 

clitus (Linnaeus) (Robins m. 

1980) Geographic range ............. Gulf of 

Preferred common name ....... mummichog st. Lawrence south to northeastern 

(Figure 1, top) Florida (Figure 2). (F. grandls, 

Other common names ....... mud minnow, previously F. h. a, includes 

pike minnow, mud dabbler, gudgeon populations whit!% range from peninsu- 

(Hi 1 debrand 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 ki 1 1 i fish, (Rosen 1973). Salt, brackish, and 

brackish water chub (Northcutt and freshwater (Rosen 1973) ; characterDavis 1983) istically close to shore (Daiber 

1982). 

1 
A TLANTIC OCEAN 

-, BALTIMORE 

KILOMETERS 

Coastal dlstrlbutlon 

NORTH CAROLINA 

Figure 2. Mid-Atlantic coast distribution of mummichog and striped 

killifish. Both are found the entire length of the Mid-Atlantic region 

close to shore (within 110 m 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 far as 75 km beyond this boundary 

(Massman 1954). 
Recent systematic studies ..... On the 

basis of differences in egg morphology, gene frequencies , and spawning 

sites between populations north and 

south of northern New Jersey, Morin 

and Able (1983) proposed retaining 

the subspecies names F. h. heterocl itus for the southern population 

and F. - h. macrolepidotus for the 

northern. Populations in the upper 

Delaware and Chesapeake Bays seem' to 

be relicts of the northern subspecies. (These distributions do not 

match those of the subspecies as 

original ly named. ) 

Scientific name ..... Fundulus majalis 

(Wal baum) (Robins et a1 . 1980) 

Preferred common name ........ striped 

killifish (Figure 1, bottom) 

Other common names ...... bull minnow, 

gudgeon (Hildebrand and Schroeder 

1928) ; striped mummichog, banded 

killifish, killie, mummie (Ursin 

1977) 

Class .................... Osteichthyes 

Order .................. Atheriniformes 

Family ................ Cyprinodontidae 

Geographic range ....... New Hampshire 

south to northeastern Florida (Figure 

2). Mainly in saltwater, less often 

in brackish (Rosen 1973). Most 

authors do not record it from 

freshwater (but see Hardy 1978). 

(See a1 so "Sal i ni ty. ") 

MORPHOLOGY/IDENTIFICATION AIDS 

The name "killifish" may be 

applied to members of three families: 

Cypri nodontidae, Anabl epidae, and 

Poeci 1 i dae (Rosen 1973). Ki 11 if i shes 

are soft-rayed, with no dorsal spine or 

spine preceding the anal fin. Most 

species are sexually dimorphic. Sexes 

are polymorphic with respect to pigment 

patterns and body and fin shape and 

size. Rosen (1973) suggested f i n-ray 

and scale counts to identify correctly 

males and females within species. His 

paper contains a key to species of 

these three families. 

In the Cyprinodontidae there is no 

lateral line and the upper surface of 

the head is conspicuously flattened 

(Hardy 1978). The anal fin in the male 

is not modified as an intromittent 

organ, and the third anal ray is 

branched. In the Fundulinae the jaw 

teeth are conical without cusps. Among 

members of the genus Fundulus, the body 

is not sharply angular or trapezoidal, 

and there is more than one series of 

jaw teeth (Rosen 1973). The mouth is 

terminal or the lower jaw projects 

slightly (Hildebrand and Schroeder 

1928). 

In both mummichogs and striped 

killifish, males are smaller than 

females after the first year. The 

origin of the dorsal fin in the male is 

directly above or in advance of the 

origin of the anal fin (Rosen 1973). 

The caudal fin is rounded in both 

species (Hardy 1978). 

Fundul us heterocl i tus: The snout is 

short, rounded, a little longer than 

diameter of eye in side view, with 8 

mandibular pores. A we1 1 -developed 

fleshy pouch is found at the anterior 

base of the anal fin. Scales along 

the lateral 1 ine number 31-35 (Rosen 

1973, but Hardy 1978 gives 31-39). 

There are 11-12 dorsal fin rays 

(Hi 1 debrand and Schroeder 1928). 

There is no conspicuous silvery sheen 

on the sides; adult females have no 

dark spot on dorsal fin (Rosen 1973), 

and are browni sh-green above, paler 

below. Small females have 13-15 dark 

vertical bars (Hi ldebrand and 

Schroeder 1928); adult females may be 

confused with those of F. confluentis (Rosen 1973). ~duit mzs 

are dark green or olive above, yellow 

beneath; have sides with about 15 

narrow silvery vertical bars and 

numerous white or ye1 1 owish spots; 

and may have a dark spot on posterior 

4-5 rays of dorsal fin. Sex-specific 

color patterns appear when fish are 

38-44 mm long (Hildebrand and 

Schroeder 1928) ; unti 1 then, young 

may be confused with adult female E. 

luciae (Rosen 1973). Adults are 
commonly 51-102 mm long (Armstrong 

and Child 1965). The largest 

specimen reported from Chesapeake Bay 

was 125 mm (Hildebrand and Schroeder 

1928). (Throughout this paper, 

length measurements are as noted by 

the original authors. Total length 

may be assumed where the authors did 

not specify standard 1 ength. ) 

Fundulus majalis: The snout is long, 

pointed, and a little less than two 

eye diameters in profile (Rosen 

1973). Overall body shape is slimmer 

than that of F. heteroclitus (Ursin 

1977). There -are 13-15 dorsal fin 

rays (Hildebrand and Schroeder 1928); 

and 32-37 scales in lateral series 

(Hardy 1978). A conspicuous si lvery 

sheen appears on the sides of young 

and adults of both sexes (Rosen 

1973); adults of both sexes have a 

black spot on the last rays of the 

dorsal fin. Adult females have one 

to several dark 1 ongi tudi nal 1 i nes 

and one or more disrupted vertical 

bars near the base of the tail; adult 

females are 01 ive above, white below. 

Adult males are dark olive on back, 

and their sides and belly are salmon 

ye1 low, with 15-20 vertical black 

stripes (Hi 1 debrand and Schroeder 

1928; Hardy 1978 gives 11-20 

stripes). Sex-speci fic color 

patterns appear when fish reach 38-51 

mm (Hi 1 debrand and Schroeder 1928) ; 

the young have bars similar to those 

of males (Rosen 1973). This species 

is the largest member of the genus in 

the study area; it frequently reaches 

152-178 mm. The record is 203 mm 

long (Ursin 1977). 

REASON FOR INCLUSION IN SERIES 

Although not valued as commercial 

or sport fishes, both ki 11 ifishes are 

important in food chains because of 

their distribution and abundance. (See 

a1 so "Predators. ") Because of their 

importance in marsh food chains, 

mummichogs may be instrumental in 

movement of organic material within and 

out of salt marsh ecosystems (Kneib et 

al. 1980). (See also "Movement 

Patterns" and "Food Habits. ") 

Mummichogs are sold as bait in 

sport fisheries for summer founder 

or fluke (Para1 ichthys dentatus) and 

young bl uef i sh or snappematomus 

saltatrix) in New York and New Jersey 

(Perlmutter 1961). . and for flounder in , el aware (M. H. Taylor, University of 

Delaware, Newark, pers. comm. ). 

Mummichogs are the primary 

predators in the Open Marsh Water 

Management mosquito control program 

currently being tested in New Jersey, 

Delaware, and Maryland (Winner et al. 

in press). A1 though Gambusia 

(mosquitofish, Fami ly Poecil iidae) is 

more often used as a biological control 

agent, the portion of the diet consisting of mosquito larvae is higher 

in studied species of Fundulus 

(grandis and conf 1 uentis). According 

to Harrington and Harrinaton (1961). 

the role if Gambusia affinis againkt 

Aedes mosquitoes has been overval ued 

because of its reputation as a predator 

on freshwater anophelines; in the 

control of sal tmarsh mosquitoes, 

primary consideration should be given 

to the 1 ocal endemic cyprinodontiform 

fishes rather than any single species. 

As a group, killifishes are used 

in research in experimental studies of 

embryo1 ogy, genetics, physiology, 

endocrinology, cytology, and behavior. 

Some of this research has important 

medical implications, such as the 

action of steroids in the regulation 

and reversal of sexual development, 

inheritance and devel opment of 

cancerous tissues, and the genetics of 

hi stocompatabi 1 i ty in tissue and organ 

transplants (Rosen 1973). 

Mummichogs are the nondomesticated 

fish most frequently used in research 

(Rosen 1973), including such disparate 

studies as bioassay for water pollution 

(Isai et al. 1979), effects of weightlessness in outer space (Hoffman et al. 

1978), ion transport in tissues (Evans 
1980), and cycling and biological 

magnification of radioisotopes (Huver 

1973). 

LIFE HISTORY 

Reproductive Physiology 

During the breeding season, males 

of mummichogs and striped killifish 

assume a brighter coloration (Ursin 

1977) and grow contact organs (Hardy 

1978). 

Fundulus spp. are oviparous. The 

ovary is single, and the number of ova 

produced depends upon the size of the 

fish. The largest number of ripe ova 

in a mummichog from Chesapeake Bay 

counted by Hildebrand and Schroeder 

(1928) was 460, in a female 98 mm long; 

the maximum for a striped killifish was 

540 (length not given). Hardy (1978) 

gave a range of 200-800 ova for 

mummichogs and 460-800 for striped 

killifish in the Mid-Atlantic region. 

According to Taylor (in press), 

production of 100-300 eggs per day for 

3-5 days is not unusual for Delaware 

mummichogs early in the spawning 

season. However, even early in the 

spawning season in North Carolina, up 

to 50% of the ova may be reabsorbed 

(Kneib and Stiven 1978). 

In male and female mummichogs, 

the gonadosomatic index (GSI = gonad 

wt/body wt x 100) rises steeply in the 

spring and declines steeply in the 

autumn (Kneib and Stiven 1978). GSI 

fluctuates in both sexes during the 

spawning season; mean GSI gradually 

declines as the season progresses 

(Taylor, in press). Although ripe ova 

may be constantly available early in 

the season (Wallace and Selman 1981), 

the number of eggs laid is always 

greater during spring tides (Taylor et 

al. 1979). This semilunar cycle is 

most pronounced late in the spawning 

season in southern populations (Taylor, 

in press). 

Food availability may be the 

ultimate control for egg production in 

mummi chogs. If feedi ng ceases, 

vi tel logenesi s (yo1 k formation) ceases 

and maturational stages are fl ushed 

from the reproductive tract; renewed 

feeding causes reappearance of 

maturational stages (Wal lace and Selman 

1980). Fritz and Garside (1975) 

reported that in an estuarine 

population, spawning mummichog females 

averaged 65 mm long and 243 ova; in a 

population in less brackish water 

(0.6-15.5 ppt), averages were 60 mm and 

161 ova. The difference was attributed 

to lower food density in the less 

brackish area. Weisberg (1981) a1 so 

attributed the reduced female GSI 

observed in crowded experimental 

populations to insufficient food 

avai labi 1 i ty. Taylor (in press), 

however, cautioned that additional 

sources of stress may contribute to 

reduced fecundity in fish populations 

in nature. 

Spawning Season and Periodicity 

The spawning seasons of mummichogs 

and striped ki 11 ifish vary with 

latitude. In Chesapeake Bay striped 

ki 11 if ish spawn from April through 

September; from New Jersey northward, 

the season lasts from June through 

August (Hardy 1978). Spawning of 

mummichogs usually begins in spring 

(March to May) and ends in later summer 

or early autumn (July to September) 

(Hardy 1978; Kneib, in press). Although 

timing and duration of spawning seasons 

differ , the water temperature range 

over which spawning occurs is similar 

(Taylor, in press). 

Mummichogs may spawn eight or more 

times in a season. Each spawning peak 

may last five or more days and 

coincides with a high spring tide of 

the full or new moon (semilunar 

periodicity: Taylor and DiMichele 

1980). A circadian periodicity may be 

superimposed on the semilunar rhythm in 

some populations; maximal spawning then 

occurs when high spring tides are at 

night (Taylor et al. 1979). Spawning 
also occurs during the day (Hardy 

1978). 

Spawning in separate genetic 

populations of mummichogs may be timed 

by different environmental stimuli 

(Wall ace and Selman 1981). Spawning 

rhythms may be timed by temperature 

(Brummett 1966), tides, moon1 ight, and 

sal inity (Taylor et a1 . 1979). . Early 

and late season peaks in spawning, such 

as Kneib and Stiven (1978) observed in 

mummichogs, may be caused by a 

combination of moderate temperatures 

and shorter day1 engths (Harri ngton 

1959). Day and Taylor (1983) found 

that a photoreceptor other than the 

pineal gland or retina of the eye 

i nf 1 uences seasonal reproduction in 

mummichogs. Female mummichogs respond 

to photoperiod, but its effectiveness 

has not been rigorously tested in males 

(Taylor, in press). In both sexes, low 

temperature prevents and high 

temperature permits gonadal development 

(Taylor, in press). The semilunar 

cycle of oogenesis persists in the 

absence of lunar or tidal stimuli 

(Taylor and Dimichele 1980). 

Schwassmann (1980) argued that we have 

no idea of the actual timing mechanism 

that entrains semi 1 unar spawning in 

mummichogs. 

Spawning Behavior 

Rivalry is intense among breeding 

male mummichogs, and size seems to be 

1 ess important than nuptial coloration 

in defense of territories. Females 

with very ripe ova may solicit males by 

turning on their sides to display their 

white be1 1 ies (Neman 1907). Adults 

gather in shallow weedy areas, and'one 

or more males chase a female until she 

stops, at which time one male moves 

beside her and pushes her against the 

vegetation (Keen1 eyside 1979). Newman 

(1907) noted that courtship, consisting 

of tandem swimming, merges gradual ly 

into spawning as the male's movements 

become progressively more excited. 

Eventually the male clasps the female; 

their bodies form a shallow S-curve. 

The pair quiver vigorously and shed 

their gametes several times in rapid 

succession (Keenleyside 1979). 

Spawning Site 

Mummichogs spawn in fresh, 

brackish, and saltwater (Hardy 1978). 

In a Delaware marsh, eggs were laid in 

clutches of 10-300, at levels only 

reached by high spring tides (1.2-1.4 m 

above mean low water). Eggs are laid 

in empty shells of the ribbed mussel 

~eukensia demissa or inside the outer 

dead leaves of smooth cordarass 4 

(Sparti na a1 terni f 1 ora) (Taylor and 

DiMichele 1983). Egg deposition in 

she1 1s has a1 so been reported in 

Virginia (Able and Castagna 1975), 

North Carolina (Kneib and Stiven 1978), 

and Georgia (Taylor et al. 1981). 

Other reported spawni ng sites i ncl ude 

algal mats in Massachusetts (Taylor et 

al. 1981) and Connecticut (Pearcy and 

Richards 1962), shallow pits covered by 

the female (Newman 1907), and scattered 

on the substrate (Keenleyside 1979). 

Selection of these different spawning 

sites may reflect an inherited 

preference or merely a response to 

substrate availability (Taylor, in 

press). 

Striped killifish spawn in still, 

shallow water close to shore and, 

presumably, in small ponds. Active 

burying of eggs by females has been 

observed (Hardy 1978). 

A key to eggs of cyprinodontids of 

the Mid-Atlantic Bight appears in Hardy 

(1978). Ferti 1 ized eggs of striped 

killifish are spherical, 2.0-3.0 mm in 

diameter, trans1 ucent ye1 low to amber, 

and slightly adhesive (Hardy 1978). 

Fertilized eggs of mummichogs are 

spherical, about 2 mm in diameter 

(range: 1.5-2.5 mm), and transparent 

yellow to amber (Hardy 1978); dead or 

infertile eggs are opaque (Taylor, in 

press). Adhesive chorionic fibrils on 

eggs may be long and dense, short and 
sparse, or absent in various 

populations (Hardy 1978; Morin and Able 

1983). 'These fibrils may anchor eggs 

to the substrate and/or retain moisture 

when eggs are stranded (Brummett and 

DuMont 1981). 

Mummichog eggs normal ly i ncubate 

in air and are not submerged until the 

next spring tide after they are laid 

(Taylor and DiMichele 1980). Eggs fail 

to develop if immersed for extended 

periods in water with less than 1 ml/l 

dissolved oxygen (DO) (Taylor, in 

press). Incubation of mummichog eggs 

in the field takes 7-8 days at 22-34 OC 

(Taylor et al. 1977); in the laboratory 

at 20 OC, hatching occurs in 10.5 days 

(Armstrong and Child 1965). In striped 

killifish 50% of the eggs hatch by 41 

days at 16-20 OC, by 17 days at 

22-26 OC, and by 12 days at 28-32 OC 

(Hardy 1978). 

In mummichogs, hatching is 

control led by oxygen concentration of 

the environment and hydration of the 

egg (DiMichele and Taylor 1980); it can 

be delayed 2 weeks or more if eggs are 

not immersed. Within seconds after 

immersion of ful ly developed eggs, 

hatching begins. The egg swells, the 

embryo's mouth opens, and heartbeat and 

respiration increase. Chorionase 

(hatching enzyme) is released from 

glands in the oral cavity and gills of 

the embryo in response to hypoxia 

("respiratory distress syndrome" of 

DiMichele and Taylor 1981). This 

enzyme weakens the chorion so that body 

movements of the embryo can break it. 

Five minutes after immersion, the 

embryo begins to turn in the egg; the 

egg ruptures in 15-20 min (Taylor et 

al. 1977). 

Yo1 k-Sac Larvae 

The yo1 k-sac larval stage lasts 

from hatching until all yolk is 

absorbed, and may also be called the 

prolarva, free embryo (Chuganova 1963), 

or alevin (Tay and Garside 1978). A 

key to yo1 k-sac larvae of fishes of the 

Mid-Atlantic Bight appears in Hardy 

(1978). Larvae of mummichogs may hatch 

with the yolk already absorbed, if 

immersion of devel oped eggs has been 

delayed (Taylor et al. 1977). At 

hatching mummichog embryos are 4.0-7.7 

mm (mean 5.0 mm) long; striped 

kill ifish are 7.0-11.00 mm long (Hardy 

1978). In the laboratory at 20 OC, 

newly hatched mummichogs require 5.5 

days to absorb the yolk. As the yo1 k 

disappears, dorsal and ventral fins 

form, and the coordination of the lower 

jaw and opercul um, undulating swimming, 

and pectoral fin movements are 

perfected (Armstrong and Chi 1 d 1965). 

Larvae 

The larval stage lasts from 

yo1 k-sac absorption until the fish 

attains the characteristic shape of the 

species. During this stage the fin 

rays and scal es devel op (Chuganova 

1963). Striped ki 11 ifi sh larvae are 

11.8-23.8 mm long. In mummichogs 

scales first appear above the pectoral 

fin at about 12.5 mm, and are well 

developed at 20.0 mm (Hardy 1978). 

Juveni 1 es 

Juveniles are also called fry or 

young-of-the-year , and have ful ly 

developed fins and a more or less 

distinct scale covering (Chuganova 

1963). The minimum described length of 

this stage in mummichogs is 25.0 mm; in 

striped killifish it is 26.0 mm (Hardy 

1978). For a detailed description of 

this stage for both species, see Hardy 

(1978). 

Adults 

Female mummi chogs are sexual ly 

mature at 38 mm and males at 32 mm; 

striped killifish females mature at 

76 mm and males at 64 mm (Hi ldebrand 

and Schroeder 1928). Most members of 

both species attain maturity during 

their second year, although some 

mummichogs may mature and spawn during 

their first year (Hardy 1978; Kneib and 

Stiven 1978). (See a1 so 
"MORPHOLOGY/IDENTIFICATION AIDS" and GROWTH CHARACTERISTICS 

"Population Size and Structure.") 

Movement Patterns and Seasonal 

Use of Habitats 

The mummichog is "one of the most 

stationary of fishes ,I1 according to 

Bigelow and Schroeder (1953). Breeding 

migrations do not occur (Rosen 1973). 

Fish over 60 mm long maintain a summer 

home range of 36-38 m along one bank of 

tidal creeks; however, some may move as 

much as 375 m (Lotrich 1975). 

In winter, mummichogs in pools 

burrow 150-200 mm into the mud 

(Chidester 1920; Hardy 1978). Others 

migrate to the mouth of the tidal 

channel where they have been living; in 

the subsequent spring they usual ly 

return up the same channel (Butner and 

Brattstrom 1960). Spring migration 

begins when the water reaches 15 OC 

(Hardy 1978); feeding begins in New 

England marshes during the first week 

in April (Valiela et al. 1977). 

During the growing season, young 

mummichogs remain in the intertidal 

zone for 6-8 weeks, inhabiting shallow 

pools on the marsh surface at low tide 

(Taylor et al. 1979). Juveniles of 

15-20 mm Standard Length (SL) begin to 

move with the adults onto the marsh 

surface to feed (Kneib in press). To 

grow at a normal rate, mummichogs may 

need food from the marsh surface for at 

least part of their diet. Despite 

abundant food resources on the marsh 

surface, mummichogs may be food limited 

because tides limit foraging time there 

(Weisberg and Lotrich 1980). 

Mummichogs maintain an endogeneous 

circadian feeding pattern which, 

superimposed upon the tidal rhythm, 

allows maximal use of food resources 

available only at high tide (Weisberg 

et al. 1981). They feed primarily at 

high tide during daylight (Weisberg and 

Lotrich 1980; Reis and Dean 1981), but 

also opportunistically (Weisberg et al. 

1981). 

Length-Weight Relationship 

Length-weight equations have been 

calculated for mummichogs: 

(1) log W = -5.232 + 3.172 log L 

where W = wt and L = Total Length 

(Fritz and Garside 1975) 

(2) log Y = -2.06 + 3.27 log X 

X = Total Length (cm) and Y = wet 

wt (g) (Meredith and Lotrich 1979) 

In some populations of mummichogs, the 

relation of weight to length may vary 

significantly through the year (Kneib 

and Stiven 1978). Females are larger 

than males of the same age, but males 

are heavier than females of the same 

length (except during the spring 

spawning peak). In the North Carolina 

population studied, males averaged 251 

mg and females 245 mg. 

Seasonal and Annual Growth 

and Production 

Growth of fi rst-season mummichogs 

in mid-summer may average 5% of their 

total weight per day. In North 

Carolina the major growing season is 

April to September. Both sexes grow 

fastest during their first two growing 

seasons. Females mature sexual ly at 

30-35 mm SL in 5-6 months (Kneib and 

Stiven 1978). 

The age of mummichogs can be 

determined by observing growth marks on 

the otoliths after a fish has 

overwintered once. For a field 

population in Nova Scotia, Fritz and 

Garside (1975) observed the following 

age-1 ength correlations: 

AGE (yr): 1 2 3 4 

LENGTH(mm): 35-50 51-74 68-83 78-95 

Mummichogs reared in the laboratory at 

13 OC have a maximum daily growth of 

0.5% of their dry weight at a feeding 

level of 6.0% of their body weight 

(Prinslow et al. 1974). There is no 
stomach in mummichogs (i.e, no region 

of the gut secretes pepsin and 

hydrochloric acid); digestion occurs 

under a1 kal i ne conditions (Targett 

1979). Nevertheless , mummi chogs have a 

higher than average assimilation 

(digestion) efficiency (87% on three 

laboratory diets) compared to other 

fishes (Weisberg and Lotrich 1982). 

Metabolic costs account for 69% of 

ingested energy. Excretion exceeds 

egested energy on some diets. 

Gross growth efficiency 

(proportion of calories ingested that 

is incorporated into biomass x 100) has 

been calculated by several authors. 

Larval mummichogs fed brine shrimp 

naupl i i were thought to be inefficient 

converters of food to biomass by Radtke 

and Dean (1979), but Kneib (1981), 

using their data, calculated that gross 

growth efficiency might be as high as 

25%-4%. Efficiency of adults was 

determined to be 12% (Weisberg and 

Lotrich 1982); this is at the low end 

of the normal range for carnivorous 

fishes. 

Total production of mummichogs in 

salt marshes is among the highest 

calculated for natural populations of 

fishes: 160 kg dry wt/ha in New 

England (Valiela et al. 1977). 

Meredith and Lotrich (1979) also 

calculated a high production (40.7 

g/m2/yr) and be1 ieved that local 

conditions were the main reason that 

their figure varied from that of 

Valiela et al. (1977). 

POPULATION DYNAMICS 

Population Size and Structure 

Mummichogs are the most abundant 

member of their genus in some areas 

(e.g. , Beaufort, North Carolina: Huver 

1973). Their name, in fact, is related 

to their abundance: "mummichog" is an 

Indian word meaning "going in crowds" 

(Nichols and Breder 1927). Schools may 

number from a few fish to several 

hundred or more (Hildebrand and 

Schroeder 1928). The density in summer 

of mummichogs longer than 40 mm may 

range from 0.35 to 6.04/m2 in certain 

estuarine habitats (Kel so 1979). From 

130,000 to 136,000 mummichogs longer 

than 30 mm have been reported along 3 

km of a tidal creek in July, and from 

63,000 to 81,000 individuals in 

September (Meredith and Lotrich 1979). 

Autumn, winter, and spring 

populations of mummichogs are dominated 

by survivors of one growing season 

(Kneib and Stiven 1978). During the 

spawning season, synchronization of 

spawning and hatching with spring tides 

results in pulses of larval and 

juvenile abundance on the marsh (Kneib 

1984). Later in the summer, the 

population is overwhelmingly dominated 

by young-of-the-year. In August, when 

a1 1 age classes are present, nearly 60% 

of the population has not yet 

overwintered and less than 8% has 

completed two growing seasons (Knei b 

and Stiven 1978). The number of fish 

in the largest size class (longer than 

70 mm) peaks in August and decl ines 

drastical ly by October (Meredith and 

Lotrich 1979). Migration, however, as 

well as mortality, may contribute to 

this decline. In October and November, 

there are very few 3-year-olds (Val iela 

et al. 1977). There is no evidence 

that any members of this species 

survive beyond their fourth growing 

season (Kneib and Stiven 1978). 

Mortal i ty 

Most mummichog eggs are fertile 

and survive to hatching; loss due to 

i nferti 1 i ty, mortality, and predation 

is about 10% (Taylor, in press). 

Young-of-the-year may have an annual 

mortality of up to 99.5% and adults, 

54% (Meredith and Lotrich 1979). 

Female mortal i ty increases dramatical ly 

after first reproduction (Kneib and 

Stiven 1978), perhaps because of 

detrimental after-effects of spawning 

(Meredith and Lotrich 1979). The 

decline in the survivorship curve with 

age is not as marked as expected if 

senescence were wholly responsible; 
predation may be an important source of 

mortal i ty (Knei b and Stiven 1978). 

During winter on the marsh, predation 

is low (Valiela et al. 1977); less than 

half as many large fish are lost 

between October and April as between 

August and October (Meredith and 

Lotrich 1979). 

ECOLOGICAL ROLE 

Feeding Behavior/Food Habits 

The protruding lower jaw and 

ti 1 ted mouth of cyprinodontids are 

well-adapted to surface feeding (Eddy 

1957). Fundulus, however, does not 

hesitate to feed in mid-water or on the 

bottom (Huver 1973). Grass shrimp 

(Palaemonetes pugio) swimming near the 

surface are often consumed. perhaps 

because they are si 1 houetted againkt 

the light (Heck and Thoman 1981). 

Vision plays an important role in 

feeding of mummichogs, but swal lowing 

depends upon another sense, probably 

01 factory (Hara 1971). Mummichogs are 

attracted to the amino acid 

gamma-ami nobutyric acid (GABA) , but 

wi 11 not swal low substrate containing 

GABA (Jonsson 1980). 

Mummichogs use all potential food 

sources: organisms in the water 

column, subtidal benthos, and 

intertidal benthos (Wei sberg and 

Lotrich 1982). Recent radioisotope 

tracer studies have shown that 56% of 

mummichogs' body carbon is derived from 

algal (benthic and planktonic) food 

chains and 44% from the Spartina food 

chain (Hughes and Sherr 1983). 

Baker-Di ttus (1978) decided that 

Fundulus uses a1 1 available food items 

except detritus. She found small 

crustaceans and polychaetes to be the 

most frequent food of both mummichogs 

and striped killifish. Fritz (1974) 

ranked food items found in the guts of 

mummichogs 22-101 mm long. Copepods 

were the most common, followed in order 

by flies (mostly larvae and pupae), 

amphi pods, polychaetes (mostly Nereis 

virens), isopods, ostracods, snails, 

insects, bivalves, algae, fishes, fish 

eggs, beetles and shrimps, and 

hymenopterans. In one North Carolina 

marsh, the major prey of mummichogs 

were fiddler crabs, polychaetes, 

tanai ds , and other small crustaceans. 

Fish less than 30 mm SL primarily ate 

small crustaceans (amphipods, tanaids, 

copepods) ; 1 arger fish consumed crabs, 

detritus, and algae more often (Kneib 

and Stiven 1978; K-neib et al. 1980). 

A1 though mummi chogs may ingest 1 arge 

quantities of detritus while feeding on 

the surface of the substrate, detritus 

is not a significant energy source for 

them because it is not assimilated 

(Prinslow et al. 1974). Mummichogs 

cannot subsist on a diet of plant 

material or detritus (Katz 1975). 

Mummichogs swallow their prey 

intact, so mouth gape limits prey size. 

Large mummichogs will also eat small 

prey, but larger prey are more 

important in their diet (Vince et al. 

1976). The numerical response of most 

infaunal invertebrates to mummichog 

predation depends on fish size more 

than fish density. Kneib and Stiven 

(1982) attributed this to very small 

infauna (oraanisms . - in bottom sediments) 

not being readily available to larger 

fish. 

Mummichog populations are large 

enough to influence their prey's 

distribution and abundance (Val iela et 

al. 1977), but few studies have 

convincingly demonstrated this result 

(Kneib 1984). Some prey species that 

may be regulated by mummichogs are the 

amphi pod Gammarus pal ustris (Van Do1 ah 

1978), the pulmonate snai 1 Melampus 

bidentatus (Vince et al. 1976), and the 

soft-shelled clam Mya arenaria (Kelso 

1979). Kneib (in press) points out 

that, by controlling smaller predators, 

mummichogs may i ndi rectly increase 

densities of some infaunal marsh 

invertebrates. A1 so, predation by 

larval and juvenile mummichogs, which 

has previously been overlooked, may 

affect the patchy distribution patterns 
of some small invertebrates (e. g. , harpacticoid copepods) in salt marshes. 

Predators 

Small tidal marsh fishes, such as Mummic hog 

killifishes, are the major prey for 

wadirlg 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 sppmi 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 mortal i tv in ma1 e mummichoas 

after they a&ieve sex-specific 

coloration (Kneib and Stiven 1978). 

From late August to early 

September in Delaware, American e e l s 

(Angui 11 a rostrata) preyed heavi ly on 

mummichogs (Lotrich 1975). Other fish 

that prey on Fundulus spp. are white 

perch (Morone ameri cana: Mi 11 er 1963), 

summer flounder (Paralichthys dentatus: 

Meredith and Lotrich 1979), and red 

drum (Sciaeno~s ocellata: Peterson and 

~eterGon 19793. Fundul us spp. are also 

eaten by crabs (Libinia, Call inectes, 

- Uca: Butner and Brattstrom 1960: 

Eurytium 1 imosum: Kneib in press).' 

Predation by blue crabs produces sizespecific losses in experimental field 

populations of mummichogs (Knei b 1982). 

Another example of size-specific predation is cannibalism of their own eggs 

by spawning mummichogs (Able and 

Castagna 1975). 

Competitors 

Three species of ki 11 if ishes 

(Figure 3)--mummichog, striped 

ki 11 if i sh, and sheepshead minnow 

(Cypri nodon vari egatus)--may occur 

toaether in permanent ponds of the high 

marsh (~ixon 1982), tidal creeks, the 

low marsh surface (Daiber 1982), and 

Sheepshead minnow 

Figure 3. Silhouettes of the three most 

common ki 11 ifishes on the Mid-Atlantic 

coast of the United States (from Rosen 

1973). 

unvegetated tidal flats (Peterson and 

Peterson 1979). All three species use 

the marsh surface at high tide, but 

only mummichogs and sheepshead minnows 

do most of their feeding at this time. 

These two species consume more 

epiphytic algae than do striped 

ki 11 ifish, which eat more benthic 

invertebrates. Sheepshead minnows are 

herbivores, but mummichogs and striped 

killifish seek invertebrates in the 

algal mats (Werme 1981). 

In the laboratory, sheepshead 

minnows chase mummichogs from their 

territories more often than they chase 

rainwater ki 11 if i sh (Lucania parva) or 

mosquitofish (Gambusia affinis). This 

may be due to similar spawning 

behavior. All four of these species 

may occur together in nature, but in 

ponds in the field, mummichogs tend to 

remain in water deeper than 10 cm, and 

only briefly visit sheepshead minnow 

territories (Itzkowitz 1974). 
Mummichogs, striped killifish, and 

Fundulus diaphanus were found in 

heterotypic schools in the intertidal 

region of a Maryland estuary by BakerDittus (1978). Mummichogs were by far 

the most abundant in May and June, but 

striped killifish slightly dominated in 

August and September. Diets of these 

two species differed in only one 

aspect: during July and August striped 

killifish continued to eat many 

polychaetes, while mummichogs ate more 

plant material. Greatest dietary 

overlap coincided with both the highest 

density of all three fish species and 

the highest food density. 

Parasites 

Parasites of Fundulus spp. are 

1 i sted by Di 11 on (1966);foff (1967) 

1 isted additional species and noted 

larval stages. Parasites of mummichogs 

and striped kill ifish include 

dinoflagellates, sporozoans, 

monogenetic flukes, digenetic flukes, 

tapeworms, roundworms, spiny-headed 

worms and copepods. Mummichogs are 

also susceptible to lymphocystis 

(caused by a virus) and ~hondrococcus 

col umnari s, a myxobacterium infecting 

the skin, gills, and fins (Sinderman 

1970). 

Since 1967 additional parasites 

have been recorded from mummichogs, 

including Eimeria funduli, a sporozoan 

unusual in that it requires grass 

shrimp (Palaemonetes pugio) as a second 

host (Upton and Duszvnski 1982); 

~r~~ano~lasma bull ocki , -a fl agel 1 ate 

vectored bv a leech in Chesa~eake Bav 

(Burreson "and Zwerner 1980); anld 

dichel ne bull ocki , a roundworm. whose 

larvae 1 ive in the anterior. and adults 

in the posterior, intestine (Kuzia 

1978). 

Additional parasites reported from 

both species of .~undulus are Swingel us 

SP. , a monoaenetic trematode (Bi 11 eter 

1974), and- Cryptobia bullocki, a 

biflagel late kinetoplastid found in the 

blood (Nigrelli et al. 1975). 

Pathogenicity of parasites varies. 

Oodinium cyprinodontiform is believed 

to be a "symphoriont ectocommensal ," 

deriving most of its energy from 

photosynthesis, but the other 

dinofl agel 1 ate recorded from Fundul us 

spp. , Amylodinium, seriously disrupts 

gill epithelium (Lom and Lawler 1971). 

Although trematode metacercarial cysts 

in their brains must cause compression 

and tissue damage, the behavior of 

infected mummichogs remains normal 

(Abbott 1968). Because of its large 

size relative to the host, larval 

Eustronglides spp. are a substantial 

nutritional drain; they also reduce 

host gonadal weight (Hirschfield et a1 . 1983). Larvae of Eustrongyl ides 

ignotus, when eaten in infected 

mummichogs, may cause high mortality in 

nestling herons and egrets (Wiese et 

a1 . 1977). 

ENVIRONMENTAL REQUI REMENTS/TOLERANCES 

Temperature 

Temperate marine fishes do not 

normally survive water temperatures 

greater than 34 OC (de Silva 1969). 

However, several species of Fundul us 

can recover from exposures to 40-42 OC 

water (Altman and Dittmer 1966). In 

laboratory experiments, mummichogs and 

striped killifish died of heat shock in 

63 min at 34 OC, 28 min at 36 OC, 9 min 

at 37 OC, and 2 min at 42 OC (Orr 

1955). 

Mummichogs are eurythermal (de 

Silva 1969). In Delaware salt marshes, 

mummichogs experience a temperature 

fluctuation of 6-35 OC (Schmelz 1964). 

In Maine salt marshes, summer tidal 

cycles expose mummichogs to rapid 

temperature changes from 15 to 30 OC. 

Swimming ability is maintained in 

nature during temperature fluctuations 

which would substantially impair 

contractile function in other fishes. 

In the laboratory, low ATPase activity 

at very cold temperatures supports 

field observations of a relatively 
torpid state for overwintering 

mummichogs (Sidell et al. 1983). 

A1 though assimilation (digestion) 

efficiency of mummichogs varies 

significantly with temperature in the 

laboratory, this is probably of 1 ittle 

ecological significance. Maxi mum 

efficiency occured from 13 to 29 OC in 

the laboratory; at 1 ower temperatures 

fish simply took longer to digest their 

food (Targett 1979). In the 

laboratory, rate of feeding of young 

mummichogs depended on temperature and 

increased rapidly above 24 OC (Radtke 

and Dean 1979). 

Oxygen consumption is relatively 

independent of acclimation temperature 

from 15 to 25 OC (Moerland and Sidell 

1981). Targett (1979) reported a 

similar relationship for the 

respiratory metabol ic rate: 

temperature independence from 13 to 

29 OC and temperature-sensitive 

reduction from 5 to 13 OC. 

Mummichogs spawn in water from 

16.5 to 25 OC (Hardy 1978). Eggs 

develop at temperatures of 12-27 OC 

with less than a 2% incidence of 

abnormality (Smith 1982). In the 

laboratory, mortality was low (7%) to 

moderate (29%) i n striped ki 11 i f i sh 

embryos transferred from an intermediate, fluctuating temperature regime 

(22-26 OC) to higher (28-32 OC) or 

lower (16-20 OC) f 1 uctuati ng temperatures (Fahy 1976). The number of 

vertebrae and dorsal fin rays developed 

by embryos was inversely correlated 

with temperature, and was i nf 1 uenced 

more by cold water than by warm (Fahy 

1979). Thermal effluents from electric 

power plants caused an elevated 

frequency of vertebral abnormal i ties in 

mummichog embryos (Mitton and Koehn 

1976). 

The median time to hatching is 

inversely related to water temperature 

(Tay and Garside 1975). Although 

hatching is not affected by the 

normal ly encountered range of 

temperatures, the hatching enzyme is 

unstable at 30 OC or above (DiMichele 

et al. 1981). 

Sal i ni ty 

A1 though mummichogs are 

physiologically euryhal ine, Fritz and 

Garside (1974) considered sal inity 

preference to be the most important 

environmental factor influencing their 

distribution in Nova Scotia. 

Laboratory studies supported this 

conclusion by showing that mummichogs 

strongly preferred salinity of 20 ppt 

over 8 ppt. 

Striped kill ifish rarely if ever 

enter freshwater (Rosen 1973; Robins et 

al. 1980, but see Hardy 1978). In the 

Chesapeake Bay area mummichogs are 

rarely taken in full saltwater and are 

found more often in freshwater than are 

striped killifish, although the 

habitats of these species over1 ap 

(Hi ldebrand and Schroeder 1928). In 

North Carolina, striped killifish also 

tend to occur in higher salinities, 

whi le mummichogs to1 erate 1 ower 

sal i ni ties (Peterson and Peterson 

1979). These studies support the 

long-held notion of the greater ability 

of mummichogs to live in a range of 

salinities, perhaps selected in a given 

area to avoid competition with 

congeners. 

Mummichogs are especially tolerant 

of abrupt salinity changes. They adapt 

so quickly that prior salinity 

acclimation has no effect on laboratory 

testing of response to sal i ni ty 

(Garside and Chi n-Yuen-Kee 1972). 

The ability of ova from northern 

and southern populations of mummichogs 

to be fertilized at various salinities 

was examined by Bush and Weis (1983). 

There was no difference in the ability 

of sperm to fertilize ova at 30 ppt. 

However, at 15 ppt there was a 

difference, which may mean that egg 

response to sal i ni ty varies 

geographically. 
Mummichog larvae survive in water 

between 0.39 ppt (fresh, by definition) 

and 100 ppt, a1 though growth is 

retarded at salinities greater than 

that of seawater (32-33 ppt). Larvae 

are active and feed in water under 1 

ppt, but do not survive more than 11 

days in tap water (Joseph and Saksena 

1966). When mummichog embryos are 

incubated in salinities of (0.5 ppt to 

60 ppt, those in 20 ppt are always the 

longest and those in freshwater are 

always the shortest (Tay and Garside 

1978). 

Temperature-Salinity Interactions 

Salinity affects response to water 

temperature in mummichogs. For 

example, the role of increased serum 

glucose in low temperature survival is 

affected by salinity (Leach and Taylor 

1977). The upper lethal temperature is 

highest (34 OC) in an isosmotic (14 

ppt) medium. The lethal temperature is 

depressed 6 OC in freshwater (0 ppt), 

but only 2.5 OC in seawater (32 ppt) 

(Garside and Chin-Yuen-Kee 1972). In 

the laboratory, mummichogs preferred 

24 OC in seawater but 22 OC in 

freshwater. Stress imposed by unusual 

salinity does not a1 low complete 

expression of thermal acclimation 

(Garside and Morrison 1977). 

Contaminants 

Mummi chogs are considered 

particularly stress- and pol 1 utiontolerant (Huver 1973), a1 though they 

are more susceptible to chlorine and 

chl orami ne than are winter f 1 ounder 

(Pseudopl euronectes americanus) and 

scup (Stenotomus chrysops) (Capuzzo et 

al. 1977). Response to pol 1 utants may 

depend upon temperature, sal i ni ty , and 

presence or absence of certain 

chemicals in the water. The same 

concentration of a pol 1 utant that is 

only mildly toxic to fish under optimal 

environmental conditions may be lethal 

to fish under thermal or osmotic 

stress. For example, under optimal 

conditions of temperature and sal i ni ty, 

a solution of 15% water soluble 

fraction (WSF) of No. 2 fuel oil, which 

contains 0.28 ppm naphthalenes, was 

only mildly toxic to mummichog embryos. 

Concentrations of 25% WSF (0.47 ppm 

naphtha1 enes) were required to attain 

>50% mortal i ty. However, under 

suboptimal conditions, especial ly 

extreme temperatures, even the 15% 

exposure resulted in >50% mortal i ty 

(Linden et a1 . 1979). Adult mummi chogs 

under thermal or osmotic stress are 

also more sensitive to the lethal 

effect of emulsions of fuel oil (Butler 

et al. 1982). 

Teratogenici ty (abi 1 i ty to cause 

birth defects) of a pollutant may also 

be higher under suboptimal sal ini ties 

(Weis et al. 1979). For example, 

teratogenicity of methylmercury is 

higher at salinities of 10 ppt than at 

30 ppt, and at temperatures of 16 OC 

than of 25 OC (Weis et al. 1979). 

Sensitivity to radiation also depends 

on temperature and salinity (Baptist et 

al. 1972). In both mummichogs and 

striped killifish, salinity plays an 

important role in copper uptake which 

differs for each species and life cycle 

stage (Bennett and Dooley 1982). 

Other Environmental Factors 

Low dissolved oxygen is a 

necessary hatching stimulus for 

mumyichog eggs (DiMichele and Taylor 

1980). Adult mummichogs survive 

overnight in anoxic tidal waters by 

gaping at the surface. Presumably 

splashing oxygenates the surface water 

that reaches the gills (M.H. Taylor, 

University of Delaware, Newark; pers. 

comm. 1. 

Although found on many substrates, 

mummichogs prefer mud (Hi ldebrand and 

Schroeder 1928). Fritz (1973), 

however, was unable to detect a 

substrate preference. Mummichogs seem 

to prefer areas with grass (S % artina 

patens) over areas without 

substrate), possibly because vegetation 

allows them to evade predators 

(confirmed in the laboratory by Ordzie 
1978). Striped killifish tend to occur 

over sandy sediments more often than do 

mummichogs, and are by far the most 

important ki 1 lifish on the unvegetated 

intertidal flats of North Carolina 

(Peterson and Peterson 1979). In the 

laboratory, both species respond more 

strongly to substrate and cover than to 

current (Rosen 1973). 

The maximum depth at which 

mummichogs are found is seldom more 

than two fathoms (Bigelow and Schroeder 

1953). Striped killifish may be found 

in water only a few centimeters in 

depth. This species is concentrated 

along the shoreline during flood tides, 

and, if washed ashore, can reenter the 

water (as can the mummichog) by a 

series of jumps (Hildebrand and 

Schroeder 1928 ; Rosen 1973). 

Mummichogs and striped kill ifish are 

positively rheotactic (face into a 

water current) (Rosen 1973). 

LITERATURE CITED 

Abbott, F.S. 1968. Metacercariae of a 

trematode in the brain of Fundulus 

heterocl i tus (L. ) Can. J. Zoo1 . 46(6): 1205-1206. 

Able, K.W. , and M. Castagna. 1975. 

Aspects of an undescri bed 

reproductive behavior in Fundul us 

heterocl i tus (Pisces: Cyprinodontidae) from Virginia. Chesapeake 

Sc i . 16 (4) : 282- 284. 

Altman, P. L., and D.S. Dittmer, eds. 

1966. Environmental biology. 

Federation of American Societies for 

Experimental Biology, Bethesda, Md. 

694 pp. 

Armstrong, P. B. , and J. S. Chi 1 d. 1965. 

Staaes in the normal develo~ment of 

~uniul us heterocl i tus. ~ioi . Bull. 

(Woods Hole) 128: 143-168. 

Baker-Di ttus, A.M. 1978. Foraging 

patterns of three sympatric 

ki 11 ifish. Copeia 1978(3): 383-389. 

Baptist, J.P., T.R. Rice, F.A. Cross, 

and T.W. Duke. 1972. Potential 

hazards from radioactive pollution of 

the estuary. Mar. Pollut. Sea Life 

FAO: 272-276. 

Bennett, R. 0. , and J. K. Do01 ey. 1982. 

Copper uptake by two sympatric . . 

species -of killifish, Fundulus 

heterocl itus and Fundulus majal is. 

J. Fish Biol. 21(4): 381-398. 

Billeter, P.A. 1974. New host and 

locality records for the genus 

Swi ngel us. J. Parasi to1 . 69(6): 

1041. 

Brummett. A. R. 1966. Observations on 

the eigs and breeding season of 

~undul us heterocl i tus at Beaufort, 

North Carol ina. Copeia 1966: 616-620. 

Brummett, A. R., and J.N. DuMont. 1981. 

A comparison of chorions from eggs of 

northern and southern populations of 

Fundul us heterocl i tus, Copei a 

1981(3): 607-614. 

Burreson, E.M., and D.E. Zwerner. 

1980. Host. ranae. life cycle, and 

pathology of ~rfiano~lasma- bul locki 

in ... lower .. - Chesa~eake Bav fishes. J. 

Protozool . 27('3): 23~-24~. 

Bush, C.. P. , and J. S. Weis. 1983. 

Effects of sal inity on ferti 1 ization 

success in two populations of 

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17 
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Assoc. 
Species Profiles: Life Histories and Envi ronmental Requirements of Coastal Fishes and Invertebrates (Mid-Atlantic)-- 

sosn -lo1 

June 1985 

& I 

ncmm .m. 

lrron DauMrNTAT'oN /8iologlcal ~e~oit 82(11.40)*1 & 

PAGE 

LTnlb.nllv#l(k 

I. nbClPbn('. kCmS).. NO. 

na0on oat0 

If Soonwving O~..nWlon Namb and Addmu 

Murnmichoq and Striped Killifish 

7. krckr(r) 

Barbara J. Abraham 

j. W-lng Oqbnlratlon Nanm and Mdmm 

National Coastal Ecosystems Team U.S. Army Corps of Engineers 

Fish and Wildlife Service Waterways Experifflent Station 

& Padormin# manlrmtion 11.4. b. 

10. Wut/Task/worh Unn 

U.S. Dep. of the Interior P.O. B~X 631' 

Washington, DC 20240 Vicksburg, MS 39180 

1% Sup amantav Not.. *u.?. 

Species 

Army 

profiles 

Corps of 

are 

Engineers 

1 i terature 

Report 

summaries 

No. TR 

of the 

EL-82-4 

taxonomy, morphology, 

-

range, 1 ife 

history, and environmental requirements of coastal aquatic species. They are designed to 

assist in environmental impact assessment. Mummichogs and striped kill ifish 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 kil lifishes 

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 semi1 unar 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. 

Parasites Feed i ng 

Fishes Salinity 

Growth Tidal marshes 

Estuaries 

Yantlt nlOp.n.Endad Tanns thmmi dog Life history Predators 

Fundul us heterocl i tus Spawning Competi tors 

Stri~ed killifish Temperature requirements 

I ~und"1us majal is Salinity requirements 

21. NO. d panes 

73 

1& Avbilbbllity Stbtamant 

Unlimited 

19. Sccunty CIbss (This ReDorl) 

Unclassified 
REGION 1 

Regional Director 

U.S. Fish and Wildlife Service 

Lloyd Five Hundred Building, Suite 1692 

500 N.E. Multnornah Street 

Portland, Oregon 97232 

REGION 4 

Regional Director 

U.S. Fish and Wildlife Service 

Richard B. Russell Building 

75 Spring Street, S.W. 

Atlanta, Georgia 30303 

REGION 2 REGION 3 

Regional Director Regional Director 

U.S. Fish and Wildlife Service U.S. Fish and Wildlife Service 

P.O. Box 1306 Federal Building, Fort Snelling 

Albuquerque, New Mexico 87 103 Twin Cities, Minnesota 55 1 1 1 

REGION 5 REGION 6 

Regional Director Regional Director 

U.S. Fish and Wildlife Service U.S. Fish and Wildlife Service 

One Gateway Center P.O. Box 25486 

Newton Corner, Massachusetts 02158 Denver Federal Center 

Denver, Colorado 80225 

REGION 7 

Regional Director 

U.S. Fish and Wildlife Service 

101 1 E. Tudor Road 

Anchorage, Alaska 99503 
DEPARTMENT OF THE INTERIOR 

U.S. FISH AID WILDLIFE SERVICE 

As the Nation's principal conservation agency, the Department of the Interior has responsibility 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 assesses 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.