Overview

Brief Summary

Living Material

Material is best and most abundant, as a rule, during the first three weeks of June, but small numbers of fertilizable eggs have been procured through July 15 at Woods Hole, Mass.

  • Agassiz, A., and C. O. Whitman, 1885. The development of osseous fishes. I. The pelagic stages of young fishes. Mem. Mus. Comp. Zool., Harvard, 14: no. 1, part 1, pp. 1-56.
  • Agassiz, A., and C. O. Whitman, 1889. The development of osseous fishes. Ii. The preembryonic stages of development. Part First. The history of the egg from fertilization to cleavage. Mem. Mus. Comp. Zool., Harvard, 14: no. 2, part 2, pp. 1-40.
  • Breder, C. M., Jr., 1948. Field Book of Marine Fishes of the Atlantic Coast from Labrador to Texas. G. P. Putnam's Sons, New York. (Rev. ea.).
  • Clapp, C. M., 1891. Some points in the development of the toad-fish (Batrachus tau). J. Morph., 5: 494-501.
  • Clapp, C. M., 1898. Relation of the axis of the embryo to the first cleavage plane. Biol. Lectures M. B. L., Wood's Holl, Mass., pp. 139-151.
  • Newman, H. H., 1907. Spawning behavior and sexual dimorphism in Fundulus heteroclitus and allied fish. Biol. Bull., 12: 314-348.
  • Newman, H. H., 1915. Development and heredity in heterogenic teleost hybrids. J. Exp. Zool., 18: 511-576.
  • Newman, H. H., 1918. Hybrids between Fundulus and mackerel. A study of paternal heredity in heterogenic hybrids. J. Exp. Zool., 26: 391-421.
  • Nicholas, J. S., 1927. The application of experimental methods to the study of developing Fundulus embryos. Proc. Nat. acad. Sci., 13: 695-698.
  • Nicholas, J. S., and J. M. Oppenheimer, 1942. Regulation and reconstitution in Fundulus. J. Exp. Zool., 90: 127-157.
  • Oppenheimer, J. M., 1937. The normal stages of Fundulus heteroclitus. Anat. Rec., 68: 1-15.
  • Russell, A., 1939. Pigment inheritance in the Fundulus-Scomber hybrid. Biol. Bull.,., 77: 423-431.
  • Solberg, A. N., 1938. The development of a bony fish. Prog. Fish. Cult., no. 40, pp. 1-19.
  • Sumner, F. B., 1903. A study of early fish development. Experimental and morphological. Arch. f. Entw., 17: 92-149.
  • Trinkaus, J. P., and J. W. Drake, 1956. Exogenous control of morphogenesis in isolated Fundulus blastoderms by nutrient chemical factors. J. Exp. Zool., 132: 311-347.
  • Wilson, H. V., 1889. The embryology of the sea bass (Serranus atrarius). Bull. U. S. Fish Comm., 9: 209-278.
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Living Material

The pale olive F. majalis female has a pattern of heavy, black longitudinal stripes on the sides, and a non-pigmented dorsal fin. The sides of the somewhat darker male bear approximately 12 broad, dark transverse bars, and there is a striking black patch on the dorsal fin.

  • Agassiz, A., and C. O. Whitman, 1885. The development of osseous fishes. I. The pelagic stages of young fishes. Mem. Mus. Comp. Zool., Harvard, 14: no. 1, part 1, pp. 1-56.
  • Agassiz, A., and C. O. Whitman, 1889. The development of osseous fishes. Ii. The preembryonic stages of development. Part First. The history of the egg from fertilization to cleavage. Mem. Mus. Comp. Zool., Harvard, 14: no. 2, part 2, pp. 1-40.
  • Breder, C. M., Jr., 1948. Field Book of Marine Fishes of the Atlantic Coast from Labrador to Texas. G. P. Putnam's Sons, New York. (Rev. ea.).
  • Clapp, C. M., 1891. Some points in the development of the toad-fish (Batrachus tau). J. Morph., 5: 494-501.
  • Clapp, C. M., 1898. Relation of the axis of the embryo to the first cleavage plane. Biol. Lectures M. B. L., Wood's Holl, Mass., pp. 139-151.
  • Newman, H. H., 1907. Spawning behavior and sexual dimorphism in Fundulus heteroclitus and allied fish. Biol. Bull., 12: 314-348.
  • Newman, H. H., 1915. Development and heredity in heterogenic teleost hybrids. J. Exp. Zool., 18: 511-576.
  • Newman, H. H., 1918. Hybrids between Fundulus and mackerel. A study of paternal heredity in heterogenic hybrids. J. Exp. Zool., 26: 391-421.
  • Nicholas, J. S., 1927. The application of experimental methods to the study of developing Fundulus embryos. Proc. Nat. acad. Sci., 13: 695-698.
  • Nicholas, J. S., and J. M. Oppenheimer, 1942. Regulation and reconstitution in Fundulus. J. Exp. Zool., 90: 127-157.
  • Oppenheimer, J. M., 1937. The normal stages of Fundulus heteroclitus. Anat. Rec., 68: 1-15.
  • Russell, A., 1939. Pigment inheritance in the Fundulus-Scomber hybrid. Biol. Bull.,., 77: 423-431.
  • Solberg, A. N., 1938. The development of a bony fish. Prog. Fish. Cult., no. 40, pp. 1-19.
  • Sumner, F. B., 1903. A study of early fish development. Experimental and morphological. Arch. f. Entw., 17: 92-149.
  • Trinkaus, J. P., and J. W. Drake, 1956. Exogenous control of morphogenesis in isolated Fundulus blastoderms by nutrient chemical factors. J. Exp. Zool., 132: 311-347.
  • Wilson, H. V., 1889. The embryology of the sea bass (Serranus atrarius). Bull. U. S. Fish Comm., 9: 209-278.
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The mummichog, Fundulus heteroclitus, is a killifish in the family Fundulidae native to brackish and coastal waters along the Eastern North American seaboard from Florida to the Gulf of St. Lawrence. Known for its hardiness and tolerance of a wide range of salinity, oxygen levels, temperature and pollution in their environment, they also have a broad diet including diatoms, a range of invertebrates, fish and fish eggs and sea grass. Mummichogs have been used widely as an experimental fish since the end of the nineteenth century, in studies such as embryology, regeneration, developmental genetics, and starting in about 1930, in studies of endocrinology and toxicity. In addition, mummichogs have been included in biology experiments on space missions such as Spacelab 3 and Bion 3. In 1973 a pair of mummichogs were the first space fish on a voyage on Spacelab 3, investigating the role of otolith (earbone) organs. They are used to stock small ponds for mosquito control, and for bait.

(Atz 1986; Kraft, Carlson, and Carlson 2006; Webster’s online dictionary; Wikipedia 8 February 2012; Wikipedia 17 December 2011)

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Living Material

Material is best and most abundant, as a rule, during the first three weeks of June, but small numbers of fertilizable eggs have been procured through July 15 at Woods Hole, Mass.

  • Agassiz, A., and C. O. Whitman, 1885. The development of osseous fishes. I. The pelagic stages of young fishes. Mem. Mus. Comp. Zool., Harvard, 14: no. 1, part 1, pp. 1-56.
  • Agassiz, A., and C. O. Whitman, 1889. The development of osseous fishes. Ii. The preembryonic stages of development. Part First. The history of the egg from fertilization to cleavage. Mem. Mus. Comp. Zool., Harvard, 14: no. 2, part 2, pp. 1-40.
  • Breder, C. M., Jr., 1948. Field Book of Marine Fishes of the Atlantic Coast from Labrador to Texas. G. P. Putnam's Sons, New York. (Rev. ea.).
  • Clapp, C. M., 1891. Some points in the development of the toad-fish (Batrachus tau). J. Morph., 5: 494-501.
  • Clapp, C. M., 1898. Relation of the axis of the embryo to the first cleavage plane. Biol. Lectures M. B. L., Wood's Holl, Mass., pp. 139-151.
  • Newman, H. H., 1907. Spawning behavior and sexual dimorphism in Fundulus heteroclitus and allied fish. Biol. Bull., 12: 314-348.
  • Newman, H. H., 1915. Development and heredity in heterogenic teleost hybrids. J. Exp. Zool., 18: 511-576.
  • Newman, H. H., 1918. Hybrids between Fundulus and mackerel. A study of paternal heredity in heterogenic hybrids. J. Exp. Zool., 26: 391-421.
  • Nicholas, J. S., 1927. The application of experimental methods to the study of developing Fundulus embryos. Proc. Nat. acad. Sci., 13: 695-698.
  • Nicholas, J. S., and J. M. Oppenheimer, 1942. Regulation and reconstitution in Fundulus. J. Exp. Zool., 90: 127-157.
  • Oppenheimer, J. M., 1937. The normal stages of Fundulus heteroclitus. Anat. Rec., 68: 1-15.
  • Russell, A., 1939. Pigment inheritance in the Fundulus-Scomber hybrid. Biol. Bull.,., 77: 423-431.
  • Solberg, A. N., 1938. The development of a bony fish. Prog. Fish. Cult., no. 40, pp. 1-19.
  • Sumner, F. B., 1903. A study of early fish development. Experimental and morphological. Arch. f. Entw., 17: 92-149.
  • Trinkaus, J. P., and J. W. Drake, 1956. Exogenous control of morphogenesis in isolated Fundulus blastoderms by nutrient chemical factors. J. Exp. Zool., 132: 311-347.
  • Wilson, H. V., 1889. The embryology of the sea bass (Serranus atrarius). Bull. U. S. Fish Comm., 9: 209-278.
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Living Material

The pale olive F. majalis female has a pattern of heavy, black longitudinal stripes on the sides, and a non-pigmented dorsal fin. The sides of the somewhat darker male bear approximately 12 broad, dark transverse bars, and there is a striking black patch on the dorsal fin.

  • Agassiz, A., and C. O. Whitman, 1885. The development of osseous fishes. I. The pelagic stages of young fishes. Mem. Mus. Comp. Zool., Harvard, 14: no. 1, part 1, pp. 1-56.
  • Agassiz, A., and C. O. Whitman, 1889. The development of osseous fishes. Ii. The preembryonic stages of development. Part First. The history of the egg from fertilization to cleavage. Mem. Mus. Comp. Zool., Harvard, 14: no. 2, part 2, pp. 1-40.
  • Breder, C. M., Jr., 1948. Field Book of Marine Fishes of the Atlantic Coast from Labrador to Texas. G. P. Putnam's Sons, New York. (Rev. ea.).
  • Clapp, C. M., 1891. Some points in the development of the toad-fish (Batrachus tau). J. Morph., 5: 494-501.
  • Clapp, C. M., 1898. Relation of the axis of the embryo to the first cleavage plane. Biol. Lectures M. B. L., Wood's Holl, Mass., pp. 139-151.
  • Newman, H. H., 1907. Spawning behavior and sexual dimorphism in Fundulus heteroclitus and allied fish. Biol. Bull., 12: 314-348.
  • Newman, H. H., 1915. Development and heredity in heterogenic teleost hybrids. J. Exp. Zool., 18: 511-576.
  • Newman, H. H., 1918. Hybrids between Fundulus and mackerel. A study of paternal heredity in heterogenic hybrids. J. Exp. Zool., 26: 391-421.
  • Nicholas, J. S., 1927. The application of experimental methods to the study of developing Fundulus embryos. Proc. Nat. acad. Sci., 13: 695-698.
  • Nicholas, J. S., and J. M. Oppenheimer, 1942. Regulation and reconstitution in Fundulus. J. Exp. Zool., 90: 127-157.
  • Oppenheimer, J. M., 1937. The normal stages of Fundulus heteroclitus. Anat. Rec., 68: 1-15.
  • Russell, A., 1939. Pigment inheritance in the Fundulus-Scomber hybrid. Biol. Bull.,., 77: 423-431.
  • Solberg, A. N., 1938. The development of a bony fish. Prog. Fish. Cult., no. 40, pp. 1-19.
  • Sumner, F. B., 1903. A study of early fish development. Experimental and morphological. Arch. f. Entw., 17: 92-149.
  • Trinkaus, J. P., and J. W. Drake, 1956. Exogenous control of morphogenesis in isolated Fundulus blastoderms by nutrient chemical factors. J. Exp. Zool., 132: 311-347.
  • Wilson, H. V., 1889. The embryology of the sea bass (Serranus atrarius). Bull. U. S. Fish Comm., 9: 209-278.
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Comprehensive Description

Biology

Occurs in saltwater marshes, tidal creeks and nearby fresh water (Ref. 86798). A resident intertidal species with homing behavior (Ref. 32612). Adults are mainly found in saltwater marshes and in tidal creeks. They may leave tide pools if aquatic conditions become inhospitable (Ref. 31184). They also enter fresh water to a limited extent (Ref. 7251). Not a seasonal killifish. They breathe air when out of water (Ref. 31184). Difficult to maintain in aquariums (Ref. 27139). Introduction has caused the decline of native species and near extinction of Aphanius baeticus in southwestern Spain (Ref. 59043).
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Distribution

Global Range: Atlantic Coast from Gulf of St. Lawrence to northeastern. Florida (Page and Burr 2011).

Subspecies macrolepidotus: Newfoundland south to Connecticut, with disjunct populations in upper Chesapeake and Delaware bays. Subspecies heteroclitus: New Jersey south to Florida, including lower southern Chesapeake and Delaware bays. (Morin and Able 1983, Able and Felley 1986). These distributions are supported by morphological (Able and Felley 1986, Morin and Able 1983) and nuclear gene data (see Brown and Chapman 1991).

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Range Description

Atlantic Coast from Gulf of St. Lawrence to northeastern. Florida (Page and Burr 2011).

Subspecies macrolepidotus: Newfoundland south to Connecticut, with disjunct populations in upper Chesapeake and Delaware bays. Subspecies heteroclitus: New Jersey south to Florida, including lower southern Chesapeake and Delaware bays. (Morin and Able 1983, Able and Felley 1986). These distributions are supported by morphological (Able and Felley 1986, Morin and Able 1983) and nuclear gene data (see Brown and Chapman 1991).
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Gulf of St. Lawrence to northeastern Florida
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Gulf of St. Lawrence to northeastern Florida
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Fundulus heteroclitus, a species of killifish commonly known as the mummichog, occurs along the Atlantic coast of North America. They extend from the Gulf of St. Lawrence all of the way to the gulf coast of Texas. The waters of Sable Island, southeast of Halifax, Canada, has also been known to be inhabited by Fundulusheteroclitus. These remarkable fish also live inland in tidal creeks and lagoons (Rutherford, 1996).

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Western Atlantic: Gulf of St. Lawrence to northeast Florida, USA. Two subspecies were previously recognized: Fundulus heteroclitus heteroclitus and Fundulus heteroclitus macrolepidotus (Ref. 86798). Introduced to southern Portugal and southern Spain (Ref. 59043).
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National Distribution

Canada

Origin: Native

Regularity: Regularly occurring

Currently: Present

Confidence: Confident

Type of Residency: Year-round

United States

Origin: Native

Regularity: Regularly occurring

Currently: Present

Confidence: Confident

Type of Residency: Year-round

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Eastern Canada and eastern U.S.A.; introduced elsewhere.
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Coast of North America,from the Gulf of St.Lawrence to Texas.
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Physical Description

Morphology

As adults, Fundulus heteroclitus range between 12.7 and 17.8 centimeters in length, the females growing larger than the males. They have flattened heads and the mouth is turned upward, clearly an adaptation to feeding at the surface of the water. This attractive fish is dimorphic, meaning that males and females have different physical characteristics. The males are darker in color than the females and exhibit blue or orange markings during the breeding season (Save the Bay, date unknown). Males are dark olive green on the dorsal side and lighter yellow on the ventral side. They also display vertical stripes along their sides. Females are silverish yellow on the ventral side and that color gradually fades to a more distinct yellow on the dorsal side. They also lack the stripes that male Fundulus heteroclitus display. All Fundulus heteroclitus have a single soft dorsal fin and their pelvic fins are located close to the rear fin (National Oceanic and Atmospheric Administration Coastal Services Center, 2001).

Other Physical Features: bilateral symmetry

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Size

Max. size

15.0 cm TL (male/unsexed; (Ref. 27139)); max. reported age: 4 years (Ref. 59043)
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Length: 13 cm

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Maximum size: 125 mm TL
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to 15.0 cm TL (male/unsexed).
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Diagnostic Description

Distinguished from nearly identical species Fundulus grandis by having the following characters: more convex upper profile; dark bars alternating with silvery interspaces on side; small ocellus at rear of dorsal fin of male; and each mandible with 4 pores (Ref. 86798).
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Ecology

Habitat

Habitat and Ecology

Habitat and Ecology
Mummichogs are common in salt marsh flats, estuaries, and tidal creeks, especially where there is abundant submergent and emergent vegetation. Adults use intertidal zone only when it is flooded; young remain on marsh even at low tide, inhabiting shallow puddles (Kneib 1986). They occasionally enter freshwater streams and rivers (Lee et al. 1980, Page and Burr 2011). Individuals may burrow into bottom mud in winter. Spawning occurs in fresh, brackish, or saltwater; generally in estuarine and salt marsh environments. Eggs are laid in various sites at levels reached only by high spring tides; usually in sand in New England populations and in Spartina alterniflora or empty Geukensia demissa shells in Middle Atlantic and southern populations (Taylor 1986). Eggs normally incubate in air (aerial incubation apparently is essential for survival), not submerged until next spring tide. Abrupt decreases in salinity (e.g. due to spring freshets) may decrease fertilization success and increase larval mortality in local populations (Able and Palmer 1988).

Systems
  • Freshwater
  • Marine
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nektonic
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Benthopelagic species, found in saltwater marshes and tidal creeks, occasionally enters fresh water.
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nektonic
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Benthopelagic species, found in saltwater marshes and tidal creeks, occasionally enters fresh water.
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Tidal creeks are the habitat of choice for Fundulus heteroclitus. They are also found in saltwater marshes, estuaries, and in sheltered shores where tides flow over eelgrass. The common feature between all of these habitats is that there is submerged vegetation where the fish can spawn and feed (Rutherford, 1996).

Fundulus heteroclitus is a remarkable fish. It has proven to be one of the most hardy and adaptable fish known. Most fish cannot survive for any period of time in waters as warm as 34° C. However, the mummichog can survive in this temperature for up to 63 minutes before falling victim to heat shock. It can also withstand temperature fluctuations from 6° C to 35° C (Abraham, 1985).

The mummichog also has a great tolerance to changes in salinity. Some mummichogs, such as the ones inhabiting the Chesapeake Bay area, prefer to live in freshwater and rarely, if ever, find themselves in salt water. Other mummichogs live along the coast in bays filled with seawater. The fish's upper limit for salinity is 106 - 120.3 ppt, while the average salinity of sea water is 32-33 ppt. This demonstrates the huge range of salinities that the mummichog can survive in (Abraham, 1985).

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Habitat Type: Freshwater

Comments: Mummichogs are common in salt marsh flats, estuaries, and tidal creeks, especially where there is abundant submergent and emergent vegetation. Adults use intertidal zone only when it is flooded; young remain on marsh even at low tide, inhabiting shallow puddles (Kneib 1986). They occasionally enter freshwater streams and rivers (Lee et al. 1980, Page and Burr 2011). Individuals may burrow into bottom mud in winter. Spawning occurs in fresh, brackish, or saltwater; generally in estuarine and salt marsh environments. Eggs are laid in various sites at levels reached only by high spring tides; usually in sand in New England populations and in Spartina alterniflora or empty Geukensia demissa shells in Middle Atlantic and southern populations (Taylor 1986). Eggs normally incubate in air (aerial incubation apparently is essential for survival), not submerged until next spring tide. Abrupt decreases in salinity (e.g. due to spring freshets) may decrease fertilization success and increase larval mortality in local populations (Able and Palmer 1988).

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Environment

benthopelagic; non-migratory; freshwater; brackish; marine
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Benthopelagic; freshwater; brackish; marine. A resident intertidal species with homing behavior. Mainly found in saltwater marshes and in tidal creeks. May leave tidepools if aquatic conditions become inhospitable. Enters fresh water to limited extent. Breathes air when out of water.
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Migration

Non-Migrant: Yes. At least some populations of this species do not make significant seasonal migrations. Juvenile dispersal is not considered a migration.

Locally Migrant: Yes. At least some populations of this species make local extended movements (generally less than 200 km) at particular times of the year (e.g., to breeding or wintering grounds, to hibernation sites).

Locally Migrant: No. No populations of this species make annual migrations of over 200 km.

May migrate to mouth of tidal channel for winter, return up same channel in spring (Abraham 1985).

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Trophic Strategy

Mummichogs primarily feed at the surface of the water (Government of Newfoundland and Labrador, date unknown). This feeding occurs predominantly at high tide during the daytime. They are however somewhat opportunistic feeders and will feed at all levels of the aquatic zone as long as there is food available. Mummichogs feed on a large variety of organisms. Some of the things that they eat include phytoplankton, mollusks, crustaceans, insect larvae, eggs of their own species, and vegetation such as eel grass. These fish have also been known to eat other smaller fish (Rutherford, 1996).

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A resident intertidal species with homing behavior (Ref. 32612). Mainly found in saltwater marshes and in tidal creeks. May leave tide pools if aquatic conditions become inhospitable (Ref. 31184). Enters fresh water to limited extent (Ref. 7251). Omnivorous feeder, food includes small crustaceans, polychaetes, insect larvae and vegetable matter. Preyed upon by kingfishers, small mammals, brook trout and bullfrogs.
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Comments: Feeds at surface, mid-water, and off bottom mainly on various invertebrates, also algae and detritus. Feeds mainly at high tide during daylight, but also feeds opportunistically (Abraham 1985).

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Omnivorous; vegetable matter, foraminifera, shrimps and other small crustacea, small mollusks; small fish.
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Associations

Known predators

Fundulus heteroclitus (common mummichog) is prey of:
Anatidae
Anguilliformes
Homo sapiens
Laridae

Based on studies in:
USA: Rhode Island (Marine)

This list may not be complete but is based on published studies.
  • S. W. Nixon and C. A. Oviatt, Ecology of a New England salt marsh, Ecol. Monog. 43:463-498, from p. 491 (1973).
  • S. W. Nixon and C. A. Oviatt, Ecology of a New England salt marsh, Ecol. Monogr. 43:463-498, from p. 491 (1973).
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Known prey organisms

Fundulus heteroclitus (common mummichog) preys on:
detritus
Nematoda
Ostracoda
Copepoda
Amphipoda
Decapoda

Based on studies in:
USA: Rhode Island (Marine)

This list may not be complete but is based on published studies.
  • S. W. Nixon and C. A. Oviatt, Ecology of a New England salt marsh, Ecol. Monog. 43:463-498, from p. 491 (1973).
  • S. W. Nixon and C. A. Oviatt, Ecology of a New England salt marsh, Ecol. Monogr. 43:463-498, from p. 491 (1973).
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Population Biology

Number of Occurrences

Note: For many non-migratory species, occurrences are roughly equivalent to populations.

Estimated Number of Occurrences: > 300

Comments: This species is represented by a large number of occurrences (subpopulations).

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Global Abundance

>1,000,000 individuals

Comments: Total adult population size is unknown but very large. This species is common in much of its range.

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General Ecology

Summer density of individuals longer than 40 mm may range from 0.35-6.04/ sq m in certain estuaries. Individuals longer than 60 mm maintained summer range of 36-38 m along bank of tidal creek; some moved up to 375 m (Abraham 1985). Preyed on by many species of fishes and wading birds; blue crab is a major predator of adults in some salt marshes. Predation by adult mummichogs and xanthid crabs may contribute to the high mortality of larvae and juveniles (Kneib 1986). See Weisburg (1986) for a discussion of competition and coexistence among this and other FUNDULUS species.

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Life History and Behavior

Behavior

Diet

Feeds on crustaceans, polychaetes, insect larvae and vegetables
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Diet

Feeds on crustaceans, polychaetes, insect larvae and vegetables
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Perception Channels: tactile ; chemical

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Cyclicity

Comments: May become inactive in winter (Abraham 1985).

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Life Cycle

Later Stages of Development

The following schedule is based on observations made at room temperatures which approximated 22-25° C. Times are recorded from insemination.

4-1/2 - 5-1/2 hours
StageTime
Blastodisc formation25-35 minutes
First cleavage2-3 hours
Four-cell stage2-1/2-3-1/2 hours
Eight-cell stage4-5 hours
Sixteen-cell stage
Early high blastula (Oppenheimer Stage 8)10 hours
Late blastula (Oppenheimer Stage 9)12 hours
Expanding blastula (Oppenheimer Stage 11)17 hours
Early gastrula; embryonic shield (Oppenheimer Stage 12)1 day
Middle gastrula; keel (Oppenheimer Stage 13)2 days
Late gastrula, closure of blastopore (Oppenheimer Stages 14-15)2-1/2 - 3 days
Formation of brain and auditory capsules; 4-14 somites (Oppenheimer Stage 18)3-1/2 days
Heart-beat, embryonic circulation (Oppenheimer Stage 20)4 days

  • Agassiz, A., and C. O. Whitman, 1885. The development of osseous fishes. I. The pelagic stages of young fishes. Mem. Mus. Comp. Zool., Harvard, 14: no. 1, part 1, pp. 1-56.
  • Agassiz, A., and C. O. Whitman, 1889. The development of osseous fishes. Ii. The preembryonic stages of development. Part First. The history of the egg from fertilization to cleavage. Mem. Mus. Comp. Zool., Harvard, 14: no. 2, part 2, pp. 1-40.
  • Breder, C. M., Jr., 1948. Field Book of Marine Fishes of the Atlantic Coast from Labrador to Texas. G. P. Putnam's Sons, New York. (Rev. ea.).
  • Clapp, C. M., 1891. Some points in the development of the toad-fish (Batrachus tau). J. Morph., 5: 494-501.
  • Clapp, C. M., 1898. Relation of the axis of the embryo to the first cleavage plane. Biol. Lectures M. B. L., Wood's Holl, Mass., pp. 139-151.
  • Newman, H. H., 1907. Spawning behavior and sexual dimorphism in Fundulus heteroclitus and allied fish. Biol. Bull., 12: 314-348.
  • Newman, H. H., 1915. Development and heredity in heterogenic teleost hybrids. J. Exp. Zool., 18: 511-576.
  • Newman, H. H., 1918. Hybrids between Fundulus and mackerel. A study of paternal heredity in heterogenic hybrids. J. Exp. Zool., 26: 391-421.
  • Nicholas, J. S., 1927. The application of experimental methods to the study of developing Fundulus embryos. Proc. Nat. acad. Sci., 13: 695-698.
  • Nicholas, J. S., and J. M. Oppenheimer, 1942. Regulation and reconstitution in Fundulus. J. Exp. Zool., 90: 127-157.
  • Oppenheimer, J. M., 1937. The normal stages of Fundulus heteroclitus. Anat. Rec., 68: 1-15.
  • Russell, A., 1939. Pigment inheritance in the Fundulus-Scomber hybrid. Biol. Bull.,., 77: 423-431.
  • Solberg, A. N., 1938. The development of a bony fish. Prog. Fish. Cult., no. 40, pp. 1-19.
  • Sumner, F. B., 1903. A study of early fish development. Experimental and morphological. Arch. f. Entw., 17: 92-149.
  • Trinkaus, J. P., and J. W. Drake, 1956. Exogenous control of morphogenesis in isolated Fundulus blastoderms by nutrient chemical factors. J. Exp. Zool., 132: 311-347.
  • Wilson, H. V., 1889. The embryology of the sea bass (Serranus atrarius). Bull. U. S. Fish Comm., 9: 209-278.
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Later Stages of Development and Metamorphosis

The sexes of both species of Fundulus are quite easily identified and obtained. The mature F. heteroclitus female is pale olive in color and usually has no definite bars or spots, although young females may have indistinct, dark, transverse bars on the sides; the dorsal fin is non-pigmented. The adult male of this species is a dull, dark green color, with narrow, ill-defined transverse bars composed of silvery spots; the dorsal fin is black-pigmented, in a mottled pattern.

  • Agassiz, A., and C. O. Whitman, 1885. The development of osseous fishes. I. The pelagic stages of young fishes. Mem. Mus. Comp. Zool., Harvard, 14: no. 1, part 1, pp. 1-56.
  • Agassiz, A., and C. O. Whitman, 1889. The development of osseous fishes. Ii. The preembryonic stages of development. Part First. The history of the egg from fertilization to cleavage. Mem. Mus. Comp. Zool., Harvard, 14: no. 2, part 2, pp. 1-40.
  • Breder, C. M., Jr., 1948. Field Book of Marine Fishes of the Atlantic Coast from Labrador to Texas. G. P. Putnam's Sons, New York. (Rev. ea.).
  • Clapp, C. M., 1891. Some points in the development of the toad-fish (Batrachus tau). J. Morph., 5: 494-501.
  • Clapp, C. M., 1898. Relation of the axis of the embryo to the first cleavage plane. Biol. Lectures M. B. L., Wood's Holl, Mass., pp. 139-151.
  • Newman, H. H., 1907. Spawning behavior and sexual dimorphism in Fundulus heteroclitus and allied fish. Biol. Bull., 12: 314-348.
  • Newman, H. H., 1915. Development and heredity in heterogenic teleost hybrids. J. Exp. Zool., 18: 511-576.
  • Newman, H. H., 1918. Hybrids between Fundulus and mackerel. A study of paternal heredity in heterogenic hybrids. J. Exp. Zool., 26: 391-421.
  • Nicholas, J. S., 1927. The application of experimental methods to the study of developing Fundulus embryos. Proc. Nat. acad. Sci., 13: 695-698.
  • Nicholas, J. S., and J. M. Oppenheimer, 1942. Regulation and reconstitution in Fundulus. J. Exp. Zool., 90: 127-157.
  • Oppenheimer, J. M., 1937. The normal stages of Fundulus heteroclitus. Anat. Rec., 68: 1-15.
  • Russell, A., 1939. Pigment inheritance in the Fundulus-Scomber hybrid. Biol. Bull.,., 77: 423-431.
  • Solberg, A. N., 1938. The development of a bony fish. Prog. Fish. Cult., no. 40, pp. 1-19.
  • Sumner, F. B., 1903. A study of early fish development. Experimental and morphological. Arch. f. Entw., 17: 92-149.
  • Trinkaus, J. P., and J. W. Drake, 1956. Exogenous control of morphogenesis in isolated Fundulus blastoderms by nutrient chemical factors. J. Exp. Zool., 132: 311-347.
  • Wilson, H. V., 1889. The embryology of the sea bass (Serranus atrarius). Bull. U. S. Fish Comm., 9: 209-278.
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Deposits eggs in the shells of Modiolus demissus (Ref. 26281).
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Later Stages of Development

The following schedule is based on observations made at room temperatures which approximated 22-25° C. Times are recorded from insemination.

4-1/2 - 5-1/2 hours
StageTime
Blastodisc formation25-35 minutes
First cleavage2-3 hours
Four-cell stage2-1/2-3-1/2 hours
Eight-cell stage4-5 hours
Sixteen-cell stage
Early high blastula (Oppenheimer Stage 8)10 hours
Late blastula (Oppenheimer Stage 9)12 hours
Expanding blastula (Oppenheimer Stage 11)17 hours
Early gastrula; embryonic shield (Oppenheimer Stage 12)1 day
Middle gastrula; keel (Oppenheimer Stage 13)2 days
Late gastrula, closure of blastopore (Oppenheimer Stages 14-15)2-1/2 - 3 days
Formation of brain and auditory capsules; 4-14 somites (Oppenheimer Stage 18)3-1/2 days
Heart-beat, embryonic circulation (Oppenheimer Stage 20)4 days

  • Agassiz, A., and C. O. Whitman, 1885. The development of osseous fishes. I. The pelagic stages of young fishes. Mem. Mus. Comp. Zool., Harvard, 14: no. 1, part 1, pp. 1-56.
  • Agassiz, A., and C. O. Whitman, 1889. The development of osseous fishes. Ii. The preembryonic stages of development. Part First. The history of the egg from fertilization to cleavage. Mem. Mus. Comp. Zool., Harvard, 14: no. 2, part 2, pp. 1-40.
  • Breder, C. M., Jr., 1948. Field Book of Marine Fishes of the Atlantic Coast from Labrador to Texas. G. P. Putnam's Sons, New York. (Rev. ea.).
  • Clapp, C. M., 1891. Some points in the development of the toad-fish (Batrachus tau). J. Morph., 5: 494-501.
  • Clapp, C. M., 1898. Relation of the axis of the embryo to the first cleavage plane. Biol. Lectures M. B. L., Wood's Holl, Mass., pp. 139-151.
  • Newman, H. H., 1907. Spawning behavior and sexual dimorphism in Fundulus heteroclitus and allied fish. Biol. Bull., 12: 314-348.
  • Newman, H. H., 1915. Development and heredity in heterogenic teleost hybrids. J. Exp. Zool., 18: 511-576.
  • Newman, H. H., 1918. Hybrids between Fundulus and mackerel. A study of paternal heredity in heterogenic hybrids. J. Exp. Zool., 26: 391-421.
  • Nicholas, J. S., 1927. The application of experimental methods to the study of developing Fundulus embryos. Proc. Nat. acad. Sci., 13: 695-698.
  • Nicholas, J. S., and J. M. Oppenheimer, 1942. Regulation and reconstitution in Fundulus. J. Exp. Zool., 90: 127-157.
  • Oppenheimer, J. M., 1937. The normal stages of Fundulus heteroclitus. Anat. Rec., 68: 1-15.
  • Russell, A., 1939. Pigment inheritance in the Fundulus-Scomber hybrid. Biol. Bull.,., 77: 423-431.
  • Solberg, A. N., 1938. The development of a bony fish. Prog. Fish. Cult., no. 40, pp. 1-19.
  • Sumner, F. B., 1903. A study of early fish development. Experimental and morphological. Arch. f. Entw., 17: 92-149.
  • Trinkaus, J. P., and J. W. Drake, 1956. Exogenous control of morphogenesis in isolated Fundulus blastoderms by nutrient chemical factors. J. Exp. Zool., 132: 311-347.
  • Wilson, H. V., 1889. The embryology of the sea bass (Serranus atrarius). Bull. U. S. Fish Comm., 9: 209-278.
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Later Stages of Development and Metamorphosis

The sexes of both species of Fundulus are quite easily identified and obtained. The mature F. heteroclitus female is pale olive in color and usually has no definite bars or spots, although young females may have indistinct, dark, transverse bars on the sides; the dorsal fin is non-pigmented. The adult male of this species is a dull, dark green color, with narrow, ill-defined transverse bars composed of silvery spots; the dorsal fin is black-pigmented, in a mottled pattern.

  • Agassiz, A., and C. O. Whitman, 1885. The development of osseous fishes. I. The pelagic stages of young fishes. Mem. Mus. Comp. Zool., Harvard, 14: no. 1, part 1, pp. 1-56.
  • Agassiz, A., and C. O. Whitman, 1889. The development of osseous fishes. Ii. The preembryonic stages of development. Part First. The history of the egg from fertilization to cleavage. Mem. Mus. Comp. Zool., Harvard, 14: no. 2, part 2, pp. 1-40.
  • Breder, C. M., Jr., 1948. Field Book of Marine Fishes of the Atlantic Coast from Labrador to Texas. G. P. Putnam's Sons, New York. (Rev. ea.).
  • Clapp, C. M., 1891. Some points in the development of the toad-fish (Batrachus tau). J. Morph., 5: 494-501.
  • Clapp, C. M., 1898. Relation of the axis of the embryo to the first cleavage plane. Biol. Lectures M. B. L., Wood's Holl, Mass., pp. 139-151.
  • Newman, H. H., 1907. Spawning behavior and sexual dimorphism in Fundulus heteroclitus and allied fish. Biol. Bull., 12: 314-348.
  • Newman, H. H., 1915. Development and heredity in heterogenic teleost hybrids. J. Exp. Zool., 18: 511-576.
  • Newman, H. H., 1918. Hybrids between Fundulus and mackerel. A study of paternal heredity in heterogenic hybrids. J. Exp. Zool., 26: 391-421.
  • Nicholas, J. S., 1927. The application of experimental methods to the study of developing Fundulus embryos. Proc. Nat. acad. Sci., 13: 695-698.
  • Nicholas, J. S., and J. M. Oppenheimer, 1942. Regulation and reconstitution in Fundulus. J. Exp. Zool., 90: 127-157.
  • Oppenheimer, J. M., 1937. The normal stages of Fundulus heteroclitus. Anat. Rec., 68: 1-15.
  • Russell, A., 1939. Pigment inheritance in the Fundulus-Scomber hybrid. Biol. Bull.,., 77: 423-431.
  • Solberg, A. N., 1938. The development of a bony fish. Prog. Fish. Cult., no. 40, pp. 1-19.
  • Sumner, F. B., 1903. A study of early fish development. Experimental and morphological. Arch. f. Entw., 17: 92-149.
  • Trinkaus, J. P., and J. W. Drake, 1956. Exogenous control of morphogenesis in isolated Fundulus blastoderms by nutrient chemical factors. J. Exp. Zool., 132: 311-347.
  • Wilson, H. V., 1889. The embryology of the sea bass (Serranus atrarius). Bull. U. S. Fish Comm., 9: 209-278.
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Reproduction

Fertilization and Cleavage

The periblast appears 16-24 hours after fertilization. The uncleaved protoplasm around the margin of the group of blastomeres is called the marginal periblast, while that beneath the blastodisc (visible only in sections) is the central periblast. At about this same time, the large, pinkish periblast nuclei may be visible. The nuclei of the marginal row of cells gradually become free of cell outlines, continue their divisions and migrate into the marginal periblast, converting it into a nucleated but non-cellular structure. Subsequent to the nucleation of the periblast, the blastoderm changes in form and size, and the embryo is now referred to as a blastula. Soon the margin of the blastodisc thickens (due both to a peripheral increase in cells and to a thinning of the central part of the disc), to form the germ-ring; this structure is best observed in eggs of F. majalis.

During the next few hours, the germ-ring grows completely over the surface of the yolk mass, so that the uncovered portion of the egg (the blastopore) is finally covered. This process of blastopore closure occurs after the first stages of formation of the embryonic axis. Under favorable conditions, it is sometimes possible to observe the beginning of gastrulation; a slight indentation appears at the edge of the germ-ring, usually when the yolk is about one-fourth covered. Staining with neutral red (one or two drops of a 0.5% solution in a Syracuse dish of sea water) may make easier the identification of the germ-ring and periblast.

While the germ-ring is extending around the yolk, the embryonic axis is being established. The first indication of this process is a cellular thickening, the embryonic shield, resulting from a more active movement of cells in one region of the germ-ring It is usually initiated when the blastoderm has covered from onequarter to one-third the surface of the yolk. When the blastoderm has spread to cover approximately one-half the yolk, the embryonic shield has become a bluntly triangular area, extending from the margin of one portion of the germ-ring almost to the center of the blastoderm. The shield can best be identified in profile view. As the blastoderm spreads over the surface of the yolk, the embryo grows rapidly in length, and becomes segmented; this segmentation is confined to the mesoderm. It is suggested that embryos be removed from the chorion for observation of the later developmental stages. Although this de-chorionation is rather difficult at early stages, it can readily be accomplished later, with the use of sharpened forceps or beading needles. Injury to the yolk sac should be avoided.

After hatching, the young fish may be studied in detail if they are anaesthetized with chloretone. The paper by Oppenheimer (1937) contains further details of developmental stages.

  • Agassiz, A., and C. O. Whitman, 1885. The development of osseous fishes. I. The pelagic stages of young fishes. Mem. Mus. Comp. Zool., Harvard, 14: no. 1, part 1, pp. 1-56.
  • Agassiz, A., and C. O. Whitman, 1889. The development of osseous fishes. Ii. The preembryonic stages of development. Part First. The history of the egg from fertilization to cleavage. Mem. Mus. Comp. Zool., Harvard, 14: no. 2, part 2, pp. 1-40.
  • Breder, C. M., Jr., 1948. Field Book of Marine Fishes of the Atlantic Coast from Labrador to Texas. G. P. Putnam's Sons, New York. (Rev. ea.).
  • Clapp, C. M., 1891. Some points in the development of the toad-fish (Batrachus tau). J. Morph., 5: 494-501.
  • Clapp, C. M., 1898. Relation of the axis of the embryo to the first cleavage plane. Biol. Lectures M. B. L., Wood's Holl, Mass., pp. 139-151.
  • Newman, H. H., 1907. Spawning behavior and sexual dimorphism in Fundulus heteroclitus and allied fish. Biol. Bull., 12: 314-348.
  • Newman, H. H., 1915. Development and heredity in heterogenic teleost hybrids. J. Exp. Zool., 18: 511-576.
  • Newman, H. H., 1918. Hybrids between Fundulus and mackerel. A study of paternal heredity in heterogenic hybrids. J. Exp. Zool., 26: 391-421.
  • Nicholas, J. S., 1927. The application of experimental methods to the study of developing Fundulus embryos. Proc. Nat. acad. Sci., 13: 695-698.
  • Nicholas, J. S., and J. M. Oppenheimer, 1942. Regulation and reconstitution in Fundulus. J. Exp. Zool., 90: 127-157.
  • Oppenheimer, J. M., 1937. The normal stages of Fundulus heteroclitus. Anat. Rec., 68: 1-15.
  • Russell, A., 1939. Pigment inheritance in the Fundulus-Scomber hybrid. Biol. Bull.,., 77: 423-431.
  • Solberg, A. N., 1938. The development of a bony fish. Prog. Fish. Cult., no. 40, pp. 1-19.
  • Sumner, F. B., 1903. A study of early fish development. Experimental and morphological. Arch. f. Entw., 17: 92-149.
  • Trinkaus, J. P., and J. W. Drake, 1956. Exogenous control of morphogenesis in isolated Fundulus blastoderms by nutrient chemical factors. J. Exp. Zool., 132: 311-347.
  • Wilson, H. V., 1889. The embryology of the sea bass (Serranus atrarius). Bull. U. S. Fish Comm., 9: 209-278.
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The Unfertilized Ovum

In order to follow all the pre-cleavage changes, it is important to (1) record the exact time of insemination, and (2) transfer the eggs immediately to a slide (see above) for observation. Polar bodies have not been described for Fundulus eggs, and it is not certain what stage the egg nucleus is in at the time of fertilization. No fertilization membrane is given off.

There is a gradual accumulation of the egg protoplasm at one pole of the egg, 25-35 minutes after fertilization, to form the blastodisc or germ-disc. A groove on the surface of this blastodisc is the first indication of cleavage; it usually occurs two to three hours after fertilization. The cleavages continue for a considerable period without much change in the over-all form from that of the original blastodisc; this is called the period of the high blastula. Details of the process of cleavage are given by Oppenheimer (1937).

  • Agassiz, A., and C. O. Whitman, 1885. The development of osseous fishes. I. The pelagic stages of young fishes. Mem. Mus. Comp. Zool., Harvard, 14: no. 1, part 1, pp. 1-56.
  • Agassiz, A., and C. O. Whitman, 1889. The development of osseous fishes. Ii. The preembryonic stages of development. Part First. The history of the egg from fertilization to cleavage. Mem. Mus. Comp. Zool., Harvard, 14: no. 2, part 2, pp. 1-40.
  • Breder, C. M., Jr., 1948. Field Book of Marine Fishes of the Atlantic Coast from Labrador to Texas. G. P. Putnam's Sons, New York. (Rev. ea.).
  • Clapp, C. M., 1891. Some points in the development of the toad-fish (Batrachus tau). J. Morph., 5: 494-501.
  • Clapp, C. M., 1898. Relation of the axis of the embryo to the first cleavage plane. Biol. Lectures M. B. L., Wood's Holl, Mass., pp. 139-151.
  • Newman, H. H., 1907. Spawning behavior and sexual dimorphism in Fundulus heteroclitus and allied fish. Biol. Bull., 12: 314-348.
  • Newman, H. H., 1915. Development and heredity in heterogenic teleost hybrids. J. Exp. Zool., 18: 511-576.
  • Newman, H. H., 1918. Hybrids between Fundulus and mackerel. A study of paternal heredity in heterogenic hybrids. J. Exp. Zool., 26: 391-421.
  • Nicholas, J. S., 1927. The application of experimental methods to the study of developing Fundulus embryos. Proc. Nat. acad. Sci., 13: 695-698.
  • Nicholas, J. S., and J. M. Oppenheimer, 1942. Regulation and reconstitution in Fundulus. J. Exp. Zool., 90: 127-157.
  • Oppenheimer, J. M., 1937. The normal stages of Fundulus heteroclitus. Anat. Rec., 68: 1-15.
  • Russell, A., 1939. Pigment inheritance in the Fundulus-Scomber hybrid. Biol. Bull.,., 77: 423-431.
  • Solberg, A. N., 1938. The development of a bony fish. Prog. Fish. Cult., no. 40, pp. 1-19.
  • Sumner, F. B., 1903. A study of early fish development. Experimental and morphological. Arch. f. Entw., 17: 92-149.
  • Trinkaus, J. P., and J. W. Drake, 1956. Exogenous control of morphogenesis in isolated Fundulus blastoderms by nutrient chemical factors. J. Exp. Zool., 132: 311-347.
  • Wilson, H. V., 1889. The embryology of the sea bass (Serranus atrarius). Bull. U. S. Fish Comm., 9: 209-278.
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Breeding Season

Fish are usually delivered by the M. B. L. Supply Department in mixed lots, but it is advisable to segregate the sexes, to prevent spawning. Males and females should be placed in separate aquaria until needed, and after they have been stripped, they should be removed to a discard tank. An adequate supply of running sea water is, of course, essential.

  • Agassiz, A., and C. O. Whitman, 1885. The development of osseous fishes. I. The pelagic stages of young fishes. Mem. Mus. Comp. Zool., Harvard, 14: no. 1, part 1, pp. 1-56.
  • Agassiz, A., and C. O. Whitman, 1889. The development of osseous fishes. Ii. The preembryonic stages of development. Part First. The history of the egg from fertilization to cleavage. Mem. Mus. Comp. Zool., Harvard, 14: no. 2, part 2, pp. 1-40.
  • Breder, C. M., Jr., 1948. Field Book of Marine Fishes of the Atlantic Coast from Labrador to Texas. G. P. Putnam's Sons, New York. (Rev. ea.).
  • Clapp, C. M., 1891. Some points in the development of the toad-fish (Batrachus tau). J. Morph., 5: 494-501.
  • Clapp, C. M., 1898. Relation of the axis of the embryo to the first cleavage plane. Biol. Lectures M. B. L., Wood's Holl, Mass., pp. 139-151.
  • Newman, H. H., 1907. Spawning behavior and sexual dimorphism in Fundulus heteroclitus and allied fish. Biol. Bull., 12: 314-348.
  • Newman, H. H., 1915. Development and heredity in heterogenic teleost hybrids. J. Exp. Zool., 18: 511-576.
  • Newman, H. H., 1918. Hybrids between Fundulus and mackerel. A study of paternal heredity in heterogenic hybrids. J. Exp. Zool., 26: 391-421.
  • Nicholas, J. S., 1927. The application of experimental methods to the study of developing Fundulus embryos. Proc. Nat. acad. Sci., 13: 695-698.
  • Nicholas, J. S., and J. M. Oppenheimer, 1942. Regulation and reconstitution in Fundulus. J. Exp. Zool., 90: 127-157.
  • Oppenheimer, J. M., 1937. The normal stages of Fundulus heteroclitus. Anat. Rec., 68: 1-15.
  • Russell, A., 1939. Pigment inheritance in the Fundulus-Scomber hybrid. Biol. Bull.,., 77: 423-431.
  • Solberg, A. N., 1938. The development of a bony fish. Prog. Fish. Cult., no. 40, pp. 1-19.
  • Sumner, F. B., 1903. A study of early fish development. Experimental and morphological. Arch. f. Entw., 17: 92-149.
  • Trinkaus, J. P., and J. W. Drake, 1956. Exogenous control of morphogenesis in isolated Fundulus blastoderms by nutrient chemical factors. J. Exp. Zool., 132: 311-347.
  • Wilson, H. V., 1889. The embryology of the sea bass (Serranus atrarius). Bull. U. S. Fish Comm., 9: 209-278.
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The breeding patterns of Fundulus heteroclitus have been studied intensely. Fundulus heteroclitus have the ability to spawn up to eight times in one season (Rutherford, 1996). During spawning season, males become increasingly aggressive and they begin to display bright colors on their rear fins and bright spotting along the sides of their bodies. The spawning season begins in the spring and lasts until fall. Spawning takes place when the tides are highest during the new or full moon. This is because the eggs develop out of the water. They are laid on almost any surface around the spawning site. Common places for mummichog eggs are in empty mussel shells, on aquatic plants, in pits dug and covered by the female, and even directly on the bottom. The eggs are laid in the shallow area during high tide so when the tide goes out, they will be exposed to the air in which they develop. After the following monthly high tide, they are submerged in water again and begin to hatch (National Oceanic and Atmospheric Administration Coastal Services Center, 2001). This process takes approximately 24 days to complete. Females can release up to 460 eggs at one time and when the eggs are released, they affix themselves to whatever object they first contact (Government of Newfoundland and Labrador, date unknown).

When hatched, the larva of Fundulus heteroclitus are approximately seven millimeters long. They remain in the intertidal zone for six to eight weeks after hatching. Here they live on the outskirts of the marsh during high tides and in shallow pools during low tides. Once the larva are about 15-20 mm in length, they begin to move and swim with the adults in schools. When tides are low, these juveniles no longer stay in the shallow pools but move to subtidal marsh creeks and deep intertidal pools. Full physical maturity is reached in about two years (Rutherford, 1996).

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Spawns spring through summer or early fall. May spawn 8 or more times during season; peaks coincide with high spring tides. Eggs hatch only when eggs are inundated, usually on spring tides (in about 7-8 days). Usually sexually mature in 2nd year, some in 1st year (Abraham 1985).

On the Atlantic coast of Nova Scotia, hybrids of F. DIAPHANUS and F. HETEROCLITUS are unisexual diploid gynogens; sperm from males probably is required to stimulate embryogenesis (Dawley et al. 2000).

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Oviparous. Eggs are about 2mm in diameter, colorless or pale yellowish and are surrounded by firm capsules that sink and become sticky on contact with the water. Capsules mass together in clumps or stick fast to sand grains or other substrate. Incubation; 9 to 18 days. Upon hatching, larvae is 7 to 7.7 long, yolk is fully absorbed, and pectoral and caudal fins are fully formed.
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Fertilization and Cleavage

The periblast appears 16-24 hours after fertilization. The uncleaved protoplasm around the margin of the group of blastomeres is called the marginal periblast, while that beneath the blastodisc (visible only in sections) is the central periblast. At about this same time, the large, pinkish periblast nuclei may be visible. The nuclei of the marginal row of cells gradually become free of cell outlines, continue their divisions and migrate into the marginal periblast, converting it into a nucleated but non-cellular structure. Subsequent to the nucleation of the periblast, the blastoderm changes in form and size, and the embryo is now referred to as a blastula. Soon the margin of the blastodisc thickens (due both to a peripheral increase in cells and to a thinning of the central part of the disc), to form the germ-ring; this structure is best observed in eggs of F. majalis.

During the next few hours, the germ-ring grows completely over the surface of the yolk mass, so that the uncovered portion of the egg (the blastopore) is finally covered. This process of blastopore closure occurs after the first stages of formation of the embryonic axis. Under favorable conditions, it is sometimes possible to observe the beginning of gastrulation; a slight indentation appears at the edge of the germ-ring, usually when the yolk is about one-fourth covered. Staining with neutral red (one or two drops of a 0.5% solution in a Syracuse dish of sea water) may make easier the identification of the germ-ring and periblast.

While the germ-ring is extending around the yolk, the embryonic axis is being established. The first indication of this process is a cellular thickening, the embryonic shield, resulting from a more active movement of cells in one region of the germ-ring It is usually initiated when the blastoderm has covered from onequarter to one-third the surface of the yolk. When the blastoderm has spread to cover approximately one-half the yolk, the embryonic shield has become a bluntly triangular area, extending from the margin of one portion of the germ-ring almost to the center of the blastoderm. The shield can best be identified in profile view. As the blastoderm spreads over the surface of the yolk, the embryo grows rapidly in length, and becomes segmented; this segmentation is confined to the mesoderm. It is suggested that embryos be removed from the chorion for observation of the later developmental stages. Although this de-chorionation is rather difficult at early stages, it can readily be accomplished later, with the use of sharpened forceps or beading needles. Injury to the yolk sac should be avoided.

After hatching, the young fish may be studied in detail if they are anaesthetized with chloretone. The paper by Oppenheimer (1937) contains further details of developmental stages.

  • Agassiz, A., and C. O. Whitman, 1885. The development of osseous fishes. I. The pelagic stages of young fishes. Mem. Mus. Comp. Zool., Harvard, 14: no. 1, part 1, pp. 1-56.
  • Agassiz, A., and C. O. Whitman, 1889. The development of osseous fishes. Ii. The preembryonic stages of development. Part First. The history of the egg from fertilization to cleavage. Mem. Mus. Comp. Zool., Harvard, 14: no. 2, part 2, pp. 1-40.
  • Breder, C. M., Jr., 1948. Field Book of Marine Fishes of the Atlantic Coast from Labrador to Texas. G. P. Putnam's Sons, New York. (Rev. ea.).
  • Clapp, C. M., 1891. Some points in the development of the toad-fish (Batrachus tau). J. Morph., 5: 494-501.
  • Clapp, C. M., 1898. Relation of the axis of the embryo to the first cleavage plane. Biol. Lectures M. B. L., Wood's Holl, Mass., pp. 139-151.
  • Newman, H. H., 1907. Spawning behavior and sexual dimorphism in Fundulus heteroclitus and allied fish. Biol. Bull., 12: 314-348.
  • Newman, H. H., 1915. Development and heredity in heterogenic teleost hybrids. J. Exp. Zool., 18: 511-576.
  • Newman, H. H., 1918. Hybrids between Fundulus and mackerel. A study of paternal heredity in heterogenic hybrids. J. Exp. Zool., 26: 391-421.
  • Nicholas, J. S., 1927. The application of experimental methods to the study of developing Fundulus embryos. Proc. Nat. acad. Sci., 13: 695-698.
  • Nicholas, J. S., and J. M. Oppenheimer, 1942. Regulation and reconstitution in Fundulus. J. Exp. Zool., 90: 127-157.
  • Oppenheimer, J. M., 1937. The normal stages of Fundulus heteroclitus. Anat. Rec., 68: 1-15.
  • Russell, A., 1939. Pigment inheritance in the Fundulus-Scomber hybrid. Biol. Bull.,., 77: 423-431.
  • Solberg, A. N., 1938. The development of a bony fish. Prog. Fish. Cult., no. 40, pp. 1-19.
  • Sumner, F. B., 1903. A study of early fish development. Experimental and morphological. Arch. f. Entw., 17: 92-149.
  • Trinkaus, J. P., and J. W. Drake, 1956. Exogenous control of morphogenesis in isolated Fundulus blastoderms by nutrient chemical factors. J. Exp. Zool., 132: 311-347.
  • Wilson, H. V., 1889. The embryology of the sea bass (Serranus atrarius). Bull. U. S. Fish Comm., 9: 209-278.
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The Unfertilized Ovum

In order to follow all the pre-cleavage changes, it is important to (1) record the exact time of insemination, and (2) transfer the eggs immediately to a slide (see above) for observation. Polar bodies have not been described for Fundulus eggs, and it is not certain what stage the egg nucleus is in at the time of fertilization. No fertilization membrane is given off.

There is a gradual accumulation of the egg protoplasm at one pole of the egg, 25-35 minutes after fertilization, to form the blastodisc or germ-disc. A groove on the surface of this blastodisc is the first indication of cleavage; it usually occurs two to three hours after fertilization. The cleavages continue for a considerable period without much change in the over-all form from that of the original blastodisc; this is called the period of the high blastula. Details of the process of cleavage are given by Oppenheimer (1937).

  • Agassiz, A., and C. O. Whitman, 1885. The development of osseous fishes. I. The pelagic stages of young fishes. Mem. Mus. Comp. Zool., Harvard, 14: no. 1, part 1, pp. 1-56.
  • Agassiz, A., and C. O. Whitman, 1889. The development of osseous fishes. Ii. The preembryonic stages of development. Part First. The history of the egg from fertilization to cleavage. Mem. Mus. Comp. Zool., Harvard, 14: no. 2, part 2, pp. 1-40.
  • Breder, C. M., Jr., 1948. Field Book of Marine Fishes of the Atlantic Coast from Labrador to Texas. G. P. Putnam's Sons, New York. (Rev. ea.).
  • Clapp, C. M., 1891. Some points in the development of the toad-fish (Batrachus tau). J. Morph., 5: 494-501.
  • Clapp, C. M., 1898. Relation of the axis of the embryo to the first cleavage plane. Biol. Lectures M. B. L., Wood's Holl, Mass., pp. 139-151.
  • Newman, H. H., 1907. Spawning behavior and sexual dimorphism in Fundulus heteroclitus and allied fish. Biol. Bull., 12: 314-348.
  • Newman, H. H., 1915. Development and heredity in heterogenic teleost hybrids. J. Exp. Zool., 18: 511-576.
  • Newman, H. H., 1918. Hybrids between Fundulus and mackerel. A study of paternal heredity in heterogenic hybrids. J. Exp. Zool., 26: 391-421.
  • Nicholas, J. S., 1927. The application of experimental methods to the study of developing Fundulus embryos. Proc. Nat. acad. Sci., 13: 695-698.
  • Nicholas, J. S., and J. M. Oppenheimer, 1942. Regulation and reconstitution in Fundulus. J. Exp. Zool., 90: 127-157.
  • Oppenheimer, J. M., 1937. The normal stages of Fundulus heteroclitus. Anat. Rec., 68: 1-15.
  • Russell, A., 1939. Pigment inheritance in the Fundulus-Scomber hybrid. Biol. Bull.,., 77: 423-431.
  • Solberg, A. N., 1938. The development of a bony fish. Prog. Fish. Cult., no. 40, pp. 1-19.
  • Sumner, F. B., 1903. A study of early fish development. Experimental and morphological. Arch. f. Entw., 17: 92-149.
  • Trinkaus, J. P., and J. W. Drake, 1956. Exogenous control of morphogenesis in isolated Fundulus blastoderms by nutrient chemical factors. J. Exp. Zool., 132: 311-347.
  • Wilson, H. V., 1889. The embryology of the sea bass (Serranus atrarius). Bull. U. S. Fish Comm., 9: 209-278.
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Breeding Season

Fish are usually delivered by the M. B. L. Supply Department in mixed lots, but it is advisable to segregate the sexes, to prevent spawning. Males and females should be placed in separate aquaria until needed, and after they have been stripped, they should be removed to a discard tank. An adequate supply of running sea water is, of course, essential.

  • Agassiz, A., and C. O. Whitman, 1885. The development of osseous fishes. I. The pelagic stages of young fishes. Mem. Mus. Comp. Zool., Harvard, 14: no. 1, part 1, pp. 1-56.
  • Agassiz, A., and C. O. Whitman, 1889. The development of osseous fishes. Ii. The preembryonic stages of development. Part First. The history of the egg from fertilization to cleavage. Mem. Mus. Comp. Zool., Harvard, 14: no. 2, part 2, pp. 1-40.
  • Breder, C. M., Jr., 1948. Field Book of Marine Fishes of the Atlantic Coast from Labrador to Texas. G. P. Putnam's Sons, New York. (Rev. ea.).
  • Clapp, C. M., 1891. Some points in the development of the toad-fish (Batrachus tau). J. Morph., 5: 494-501.
  • Clapp, C. M., 1898. Relation of the axis of the embryo to the first cleavage plane. Biol. Lectures M. B. L., Wood's Holl, Mass., pp. 139-151.
  • Newman, H. H., 1907. Spawning behavior and sexual dimorphism in Fundulus heteroclitus and allied fish. Biol. Bull., 12: 314-348.
  • Newman, H. H., 1915. Development and heredity in heterogenic teleost hybrids. J. Exp. Zool., 18: 511-576.
  • Newman, H. H., 1918. Hybrids between Fundulus and mackerel. A study of paternal heredity in heterogenic hybrids. J. Exp. Zool., 26: 391-421.
  • Nicholas, J. S., 1927. The application of experimental methods to the study of developing Fundulus embryos. Proc. Nat. acad. Sci., 13: 695-698.
  • Nicholas, J. S., and J. M. Oppenheimer, 1942. Regulation and reconstitution in Fundulus. J. Exp. Zool., 90: 127-157.
  • Oppenheimer, J. M., 1937. The normal stages of Fundulus heteroclitus. Anat. Rec., 68: 1-15.
  • Russell, A., 1939. Pigment inheritance in the Fundulus-Scomber hybrid. Biol. Bull.,., 77: 423-431.
  • Solberg, A. N., 1938. The development of a bony fish. Prog. Fish. Cult., no. 40, pp. 1-19.
  • Sumner, F. B., 1903. A study of early fish development. Experimental and morphological. Arch. f. Entw., 17: 92-149.
  • Trinkaus, J. P., and J. W. Drake, 1956. Exogenous control of morphogenesis in isolated Fundulus blastoderms by nutrient chemical factors. J. Exp. Zool., 132: 311-347.
  • Wilson, H. V., 1889. The embryology of the sea bass (Serranus atrarius). Bull. U. S. Fish Comm., 9: 209-278.
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Video of spawning in Provincetown Harbor

Spawning mummichogs (fundulus heteroclitus) observed in Provincetown harbor crossing the West End breakwater. It was about 1 hour past high tide and about 3 days after the full moon. The water was about a foot deep at the time. See http://www.youtube.com/watch?v=IxEVxdwDBWE

(Note, I'm new to EOL, I'm not sure how to upload media, feel free to edit this article.)

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Molecular Biology and Genetics

Molecular Biology

Barcode data: Fundulus heteroclitus

The following is a representative barcode sequence, the centroid of all available sequences for this species.


There are 11 barcode sequences available from BOLD and GenBank.

Below is a sequence of the barcode region Cytochrome oxidase subunit 1 (COI or COX1) from a member of the species.

See the BOLD taxonomy browser for more complete information about this specimen and other sequences.

ACACGTTGATTTTTCTCAACTAACCACAAAGATATCGGTACCCTTTATTTAGTATTTGGTGCCTGAGCCGGTATAGTAGGTACAGCTCTT---AGCCTTCTTATTCGGGCGGAACTAAGCCAACCAGGCTCCCTCCTAGGGGAT---GACCAAATTTATAATGTAATCGTTACAGCACATGCATTTGTAATAATCTTTTTTATAGTTATGCCTATTATAATTGGTGGTTTTGGAAATTGACTAGTCCCTCTTATG---ATTGGTGCCCCAGACATAGCTTTTCCTCGAATAAATAATATAAGCTTCTGACTACTCCCACCCTCATTTTTACTTCTTTTAGCCTCTTCCGGTGTTGAAGCCGGGGCTGGTACAGGTTGAACAGTCTATCCCCCTCTAGCAGGTAATTTAGCTCATGCTGGGGCTTCTGTAGATTTA---ACTATTTTTTCCCTTCACTTAGCTGGTATTTCATCAATTTTAGGTGCTATTAATTTTATTACAACTATTATTAACATAAAACCTCCAGCTATCTCCCAATACCAAACCCCTCTGTTCGTCTGAGCTGTCTTAATTACTGCTGTACTTCTTCTACTTTCCTTACCAGTTCTTGCTGCA---GGAATTACAATACTGTTAACTGACCGAAATTTAAATACTACATTTTTTGATCCGGCAGGCGGAGGAGATCCAATTCTATACCAACATTTATTCTGATTCTTTGGTCACCCAGAAGTTTATATTCTCATTCTACCAGGCTTTGGTATGATTTCACATATTGTAGCATACTATTCTGGTAAAAAA---GAACCGTTTGGATATATGGGTATAGTATGAGCAATAATAGCAATTGGTCTTCTCGGTTTTATTGTTTGAGCCCATCACATATTTACAGTCGGAATAGACGTAGACACTCGAGCTTACTTTACATCTGCTACTATAATTATTGCTATTCCAACAGGAGTGAAAGTGTTTAGCTGATTA---GCTACTCTCCATGGAGGA---TCTATTAAATGAGAAACCCCTTTACTCTGAGCATTAGGATTCATTTTCCTATTTACAGTAGGGGGACTTACAGGAATTGTTTTAGCTAATTCATCCTTAGATATTGTGCTCCACGATACTTATTATGTAGTTGCTCACTTCCATTATGTT---TTATCCATGGGAGCCGTATTTGCAATTATCGCTGCCTTTGTTCATTGATTCCCTCTGTTCTCAGGTTACACCCTTCATAGCACATGAACTAAAATTCATTTTGGTATTATGTTTGTAGGCGTTAATTTAACCTTTTTCCCACAACATTTCCTTGGATTAGCAGGTATACCTCGA---CGATATTCTGATTATCCAGATGCCTATACC---CTTTGAAACACAGTGTCTTCTATTGGGTCATTAATTTCCCTTGTAGCAGTAATCATGTTTTTATTTATTATCTGAGAAGCATTCGCTGCTAAACGTGAAGTA---TTATCTGTTGAAATAACAGCAACTAAT
-- end --

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Statistics of barcoding coverage: Fundulus heteroclitus

Barcode of Life Data Systems (BOLDS) Stats
Public Records: 12
Specimens with Barcodes: 23
Species With Barcodes: 1
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Conservation

Conservation Status

IUCN Red List Assessment


Red List Category
LC
Least Concern

Red List Criteria

Version
3.1

Year Assessed
2013

Assessor/s
NatureServe

Reviewer/s
Smith, K. & Darwall, W.R.T.

Contributor/s

Justification
Listed as Least Concern in view of the large extent of occurrence, large number of subpopulations, large population size, apparently stable trend, and lack of major threats.
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US Federal List: no special status

CITES: no special status

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National NatureServe Conservation Status

Canada

Rounded National Status Rank: NNR - Unranked

United States

Rounded National Status Rank: N5 - Secure

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NatureServe Conservation Status

Rounded Global Status Rank: G5 - Secure

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Population

Population
This species is represented by a large number of occurrences (subpopulations).

Total adult population size is unknown but very large. This species is common in much of its range.

Trend over the past 10 years or three generations is uncertain but likely relatively stable.

Population Trend
Stable
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Global Short Term Trend: Relatively stable (=10% change)

Comments: Trend over the past 10 years or three generations is uncertain but likely relatively stable.

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Threats

Major Threats
No major threats are known.
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Least Concern (LC)
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Comments: No major threats are known.

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Management

Conservation Actions

Conservation Actions
Currently, this species is of relatively low conservation concern and does not require significant additional protection or major management, monitoring, or research action.
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Relevance to Humans and Ecosystems

Benefits

Preparation of Eggs for Sectioning

Eggs stripped from a female fish into diluted sea water (70% fresh water, 30% sea water) retain the morphological characteristics of freshly-extruded eggs, including the yolk platelets, oil drops, membranes, etc. A micropyle is present, but it must be observed before removal of the chorionic jelly.

  • Agassiz, A., and C. O. Whitman, 1885. The development of osseous fishes. I. The pelagic stages of young fishes. Mem. Mus. Comp. Zool., Harvard, 14: no. 1, part 1, pp. 1-56.
  • Agassiz, A., and C. O. Whitman, 1889. The development of osseous fishes. Ii. The preembryonic stages of development. Part First. The history of the egg from fertilization to cleavage. Mem. Mus. Comp. Zool., Harvard, 14: no. 2, part 2, pp. 1-40.
  • Breder, C. M., Jr., 1948. Field Book of Marine Fishes of the Atlantic Coast from Labrador to Texas. G. P. Putnam's Sons, New York. (Rev. ea.).
  • Clapp, C. M., 1891. Some points in the development of the toad-fish (Batrachus tau). J. Morph., 5: 494-501.
  • Clapp, C. M., 1898. Relation of the axis of the embryo to the first cleavage plane. Biol. Lectures M. B. L., Wood's Holl, Mass., pp. 139-151.
  • Newman, H. H., 1907. Spawning behavior and sexual dimorphism in Fundulus heteroclitus and allied fish. Biol. Bull., 12: 314-348.
  • Newman, H. H., 1915. Development and heredity in heterogenic teleost hybrids. J. Exp. Zool., 18: 511-576.
  • Newman, H. H., 1918. Hybrids between Fundulus and mackerel. A study of paternal heredity in heterogenic hybrids. J. Exp. Zool., 26: 391-421.
  • Nicholas, J. S., 1927. The application of experimental methods to the study of developing Fundulus embryos. Proc. Nat. acad. Sci., 13: 695-698.
  • Nicholas, J. S., and J. M. Oppenheimer, 1942. Regulation and reconstitution in Fundulus. J. Exp. Zool., 90: 127-157.
  • Oppenheimer, J. M., 1937. The normal stages of Fundulus heteroclitus. Anat. Rec., 68: 1-15.
  • Russell, A., 1939. Pigment inheritance in the Fundulus-Scomber hybrid. Biol. Bull.,., 77: 423-431.
  • Solberg, A. N., 1938. The development of a bony fish. Prog. Fish. Cult., no. 40, pp. 1-19.
  • Sumner, F. B., 1903. A study of early fish development. Experimental and morphological. Arch. f. Entw., 17: 92-149.
  • Trinkaus, J. P., and J. W. Drake, 1956. Exogenous control of morphogenesis in isolated Fundulus blastoderms by nutrient chemical factors. J. Exp. Zool., 132: 311-347.
  • Wilson, H. V., 1889. The embryology of the sea bass (Serranus atrarius). Bull. U. S. Fish Comm., 9: 209-278.
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Permanent Total Preparations

Eggs to be sectioned must be dechorionated before fixation, so that fluids can penetrate to the interior. (For details of this process, see the paper by Nicholas, 1927.) The following schedule for dehydration and embedding is useful.

1. Fix in Bouin's or Zenker's solution, 12-24 hours.

2. Dehydrate as usual through the alcohol series (up to and including 95% alcohol), leaving the eggs in each for one hour.

3. Absolute alcohol, two hours&emdash;use several changes.

4. Equal parts absolute alcohol and amyl acetate, two hours. 5. Amyl acetate, 24-48 hours.

6. Equal parts amyl acetate and paraffin, 12 hours (incubate at 30°).

7. Three changes of infiltrating paraffin (15 minutes in each); embed in 56-58° paraffin.

  • Agassiz, A., and C. O. Whitman, 1885. The development of osseous fishes. I. The pelagic stages of young fishes. Mem. Mus. Comp. Zool., Harvard, 14: no. 1, part 1, pp. 1-56.
  • Agassiz, A., and C. O. Whitman, 1889. The development of osseous fishes. Ii. The preembryonic stages of development. Part First. The history of the egg from fertilization to cleavage. Mem. Mus. Comp. Zool., Harvard, 14: no. 2, part 2, pp. 1-40.
  • Breder, C. M., Jr., 1948. Field Book of Marine Fishes of the Atlantic Coast from Labrador to Texas. G. P. Putnam's Sons, New York. (Rev. ea.).
  • Clapp, C. M., 1891. Some points in the development of the toad-fish (Batrachus tau). J. Morph., 5: 494-501.
  • Clapp, C. M., 1898. Relation of the axis of the embryo to the first cleavage plane. Biol. Lectures M. B. L., Wood's Holl, Mass., pp. 139-151.
  • Newman, H. H., 1907. Spawning behavior and sexual dimorphism in Fundulus heteroclitus and allied fish. Biol. Bull., 12: 314-348.
  • Newman, H. H., 1915. Development and heredity in heterogenic teleost hybrids. J. Exp. Zool., 18: 511-576.
  • Newman, H. H., 1918. Hybrids between Fundulus and mackerel. A study of paternal heredity in heterogenic hybrids. J. Exp. Zool., 26: 391-421.
  • Nicholas, J. S., 1927. The application of experimental methods to the study of developing Fundulus embryos. Proc. Nat. acad. Sci., 13: 695-698.
  • Nicholas, J. S., and J. M. Oppenheimer, 1942. Regulation and reconstitution in Fundulus. J. Exp. Zool., 90: 127-157.
  • Oppenheimer, J. M., 1937. The normal stages of Fundulus heteroclitus. Anat. Rec., 68: 1-15.
  • Russell, A., 1939. Pigment inheritance in the Fundulus-Scomber hybrid. Biol. Bull.,., 77: 423-431.
  • Solberg, A. N., 1938. The development of a bony fish. Prog. Fish. Cult., no. 40, pp. 1-19.
  • Sumner, F. B., 1903. A study of early fish development. Experimental and morphological. Arch. f. Entw., 17: 92-149.
  • Trinkaus, J. P., and J. W. Drake, 1956. Exogenous control of morphogenesis in isolated Fundulus blastoderms by nutrient chemical factors. J. Exp. Zool., 132: 311-347.
  • Wilson, H. V., 1889. The embryology of the sea bass (Serranus atrarius). Bull. U. S. Fish Comm., 9: 209-278.
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Methods of Observation

Fix the eggs in Stockard's solution (formalin, 5 parts; glacial acetic acid, 4 parts; glycerine, 6 parts; distilled water, 85 parts). This turns the protoplasm white but leaves the yolk transparent. The fixative may be used as a preservative, or the material can be transferred to 10% formalin after two days.

  • Agassiz, A., and C. O. Whitman, 1885. The development of osseous fishes. I. The pelagic stages of young fishes. Mem. Mus. Comp. Zool., Harvard, 14: no. 1, part 1, pp. 1-56.
  • Agassiz, A., and C. O. Whitman, 1889. The development of osseous fishes. Ii. The preembryonic stages of development. Part First. The history of the egg from fertilization to cleavage. Mem. Mus. Comp. Zool., Harvard, 14: no. 2, part 2, pp. 1-40.
  • Breder, C. M., Jr., 1948. Field Book of Marine Fishes of the Atlantic Coast from Labrador to Texas. G. P. Putnam's Sons, New York. (Rev. ea.).
  • Clapp, C. M., 1891. Some points in the development of the toad-fish (Batrachus tau). J. Morph., 5: 494-501.
  • Clapp, C. M., 1898. Relation of the axis of the embryo to the first cleavage plane. Biol. Lectures M. B. L., Wood's Holl, Mass., pp. 139-151.
  • Newman, H. H., 1907. Spawning behavior and sexual dimorphism in Fundulus heteroclitus and allied fish. Biol. Bull., 12: 314-348.
  • Newman, H. H., 1915. Development and heredity in heterogenic teleost hybrids. J. Exp. Zool., 18: 511-576.
  • Newman, H. H., 1918. Hybrids between Fundulus and mackerel. A study of paternal heredity in heterogenic hybrids. J. Exp. Zool., 26: 391-421.
  • Nicholas, J. S., 1927. The application of experimental methods to the study of developing Fundulus embryos. Proc. Nat. acad. Sci., 13: 695-698.
  • Nicholas, J. S., and J. M. Oppenheimer, 1942. Regulation and reconstitution in Fundulus. J. Exp. Zool., 90: 127-157.
  • Oppenheimer, J. M., 1937. The normal stages of Fundulus heteroclitus. Anat. Rec., 68: 1-15.
  • Russell, A., 1939. Pigment inheritance in the Fundulus-Scomber hybrid. Biol. Bull.,., 77: 423-431.
  • Solberg, A. N., 1938. The development of a bony fish. Prog. Fish. Cult., no. 40, pp. 1-19.
  • Sumner, F. B., 1903. A study of early fish development. Experimental and morphological. Arch. f. Entw., 17: 92-149.
  • Trinkaus, J. P., and J. W. Drake, 1956. Exogenous control of morphogenesis in isolated Fundulus blastoderms by nutrient chemical factors. J. Exp. Zool., 132: 311-347.
  • Wilson, H. V., 1889. The embryology of the sea bass (Serranus atrarius). Bull. U. S. Fish Comm., 9: 209-278.
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Preparation of Cultures

To remove the sticky outer jelly layer, roll the eggs on a piece of filter paper or paper towel until the surface of the outer membrane is left smooth and clean. This same procedure should be followed daily for stock cultures, in order to prevent clumping of the eggs.

For experimental work, where it is essential to obtain development as nearly normal as possible, the eggs are usually examined uncovered in shallow depression slides; they may be manipulated with hair loops. For classroom study, when the eggs are to be observed over long periods of time and a specific orientation is desired, either of the following methods is suggested: (1) Place the eggs in sea water in special culture slides having a depression of 1.7 to 1.8 mm. (slightly less than the diameter of the eggs); it is then possible to roll the eggs to the desired position by moving the coverslip. (2) If these special slides are not available, the eggs may be placed in a drop of sea water on an ordinary glass slide and covered with a very thin, flexible sheet of mica; water is then withdrawn (using lens or filter paper) until capillary attraction causes a pressure on the egg, so that it can be rotated as in the previous method.

Recently, Trinkaus and Drake (1956) have described a method for the in vitro culture of Fundulus blastoderms isolated from the subjacent periblast and yolk mass.

  • Agassiz, A., and C. O. Whitman, 1885. The development of osseous fishes. I. The pelagic stages of young fishes. Mem. Mus. Comp. Zool., Harvard, 14: no. 1, part 1, pp. 1-56.
  • Agassiz, A., and C. O. Whitman, 1889. The development of osseous fishes. Ii. The preembryonic stages of development. Part First. The history of the egg from fertilization to cleavage. Mem. Mus. Comp. Zool., Harvard, 14: no. 2, part 2, pp. 1-40.
  • Breder, C. M., Jr., 1948. Field Book of Marine Fishes of the Atlantic Coast from Labrador to Texas. G. P. Putnam's Sons, New York. (Rev. ea.).
  • Clapp, C. M., 1891. Some points in the development of the toad-fish (Batrachus tau). J. Morph., 5: 494-501.
  • Clapp, C. M., 1898. Relation of the axis of the embryo to the first cleavage plane. Biol. Lectures M. B. L., Wood's Holl, Mass., pp. 139-151.
  • Newman, H. H., 1907. Spawning behavior and sexual dimorphism in Fundulus heteroclitus and allied fish. Biol. Bull., 12: 314-348.
  • Newman, H. H., 1915. Development and heredity in heterogenic teleost hybrids. J. Exp. Zool., 18: 511-576.
  • Newman, H. H., 1918. Hybrids between Fundulus and mackerel. A study of paternal heredity in heterogenic hybrids. J. Exp. Zool., 26: 391-421.
  • Nicholas, J. S., 1927. The application of experimental methods to the study of developing Fundulus embryos. Proc. Nat. acad. Sci., 13: 695-698.
  • Nicholas, J. S., and J. M. Oppenheimer, 1942. Regulation and reconstitution in Fundulus. J. Exp. Zool., 90: 127-157.
  • Oppenheimer, J. M., 1937. The normal stages of Fundulus heteroclitus. Anat. Rec., 68: 1-15.
  • Russell, A., 1939. Pigment inheritance in the Fundulus-Scomber hybrid. Biol. Bull.,., 77: 423-431.
  • Solberg, A. N., 1938. The development of a bony fish. Prog. Fish. Cult., no. 40, pp. 1-19.
  • Sumner, F. B., 1903. A study of early fish development. Experimental and morphological. Arch. f. Entw., 17: 92-149.
  • Trinkaus, J. P., and J. W. Drake, 1956. Exogenous control of morphogenesis in isolated Fundulus blastoderms by nutrient chemical factors. J. Exp. Zool., 132: 311-347.
  • Wilson, H. V., 1889. The embryology of the sea bass (Serranus atrarius). Bull. U. S. Fish Comm., 9: 209-278.
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Procuring Gametes

Strip the eggs into a clean four-inch fingerbowl which has been moistened with filtered sea water. Strip the milt into a small amount of sea water, and mix the suspension with the eggs in 1/4 inch of sea water. The eggs should be inseminated as soon as possible after they are obtained from the body of the female. After 30-45 minutes, change the sea water and leave the eggs in about a 1/4 to 1/2-inch depth of sea water. Keep the fingerbowl covered with a glass plate to prevent evaporation; do not allow the eggs to clump or accumulate in one spot. The water should be changed at least twice daily.

  • Agassiz, A., and C. O. Whitman, 1885. The development of osseous fishes. I. The pelagic stages of young fishes. Mem. Mus. Comp. Zool., Harvard, 14: no. 1, part 1, pp. 1-56.
  • Agassiz, A., and C. O. Whitman, 1889. The development of osseous fishes. Ii. The preembryonic stages of development. Part First. The history of the egg from fertilization to cleavage. Mem. Mus. Comp. Zool., Harvard, 14: no. 2, part 2, pp. 1-40.
  • Breder, C. M., Jr., 1948. Field Book of Marine Fishes of the Atlantic Coast from Labrador to Texas. G. P. Putnam's Sons, New York. (Rev. ea.).
  • Clapp, C. M., 1891. Some points in the development of the toad-fish (Batrachus tau). J. Morph., 5: 494-501.
  • Clapp, C. M., 1898. Relation of the axis of the embryo to the first cleavage plane. Biol. Lectures M. B. L., Wood's Holl, Mass., pp. 139-151.
  • Newman, H. H., 1907. Spawning behavior and sexual dimorphism in Fundulus heteroclitus and allied fish. Biol. Bull., 12: 314-348.
  • Newman, H. H., 1915. Development and heredity in heterogenic teleost hybrids. J. Exp. Zool., 18: 511-576.
  • Newman, H. H., 1918. Hybrids between Fundulus and mackerel. A study of paternal heredity in heterogenic hybrids. J. Exp. Zool., 26: 391-421.
  • Nicholas, J. S., 1927. The application of experimental methods to the study of developing Fundulus embryos. Proc. Nat. acad. Sci., 13: 695-698.
  • Nicholas, J. S., and J. M. Oppenheimer, 1942. Regulation and reconstitution in Fundulus. J. Exp. Zool., 90: 127-157.
  • Oppenheimer, J. M., 1937. The normal stages of Fundulus heteroclitus. Anat. Rec., 68: 1-15.
  • Russell, A., 1939. Pigment inheritance in the Fundulus-Scomber hybrid. Biol. Bull.,., 77: 423-431.
  • Solberg, A. N., 1938. The development of a bony fish. Prog. Fish. Cult., no. 40, pp. 1-19.
  • Sumner, F. B., 1903. A study of early fish development. Experimental and morphological. Arch. f. Entw., 17: 92-149.
  • Trinkaus, J. P., and J. W. Drake, 1956. Exogenous control of morphogenesis in isolated Fundulus blastoderms by nutrient chemical factors. J. Exp. Zool., 132: 311-347.
  • Wilson, H. V., 1889. The embryology of the sea bass (Serranus atrarius). Bull. U. S. Fish Comm., 9: 209-278.
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Care of Adults

Both eggs and sperm are obtained by "stripping": the fish is held firmly in one hand while gentle pressure is applied to its abdomen with the thumb and forefinger of the other hand. As these fingers are drawn towards the anus of the fish, the pressure forces out the gametes. If the fish is held in front of a strong light source during the stripping process, the eggs may be seen passing through the oviduct which runs along the anal fin.

  • Agassiz, A., and C. O. Whitman, 1885. The development of osseous fishes. I. The pelagic stages of young fishes. Mem. Mus. Comp. Zool., Harvard, 14: no. 1, part 1, pp. 1-56.
  • Agassiz, A., and C. O. Whitman, 1889. The development of osseous fishes. Ii. The preembryonic stages of development. Part First. The history of the egg from fertilization to cleavage. Mem. Mus. Comp. Zool., Harvard, 14: no. 2, part 2, pp. 1-40.
  • Breder, C. M., Jr., 1948. Field Book of Marine Fishes of the Atlantic Coast from Labrador to Texas. G. P. Putnam's Sons, New York. (Rev. ea.).
  • Clapp, C. M., 1891. Some points in the development of the toad-fish (Batrachus tau). J. Morph., 5: 494-501.
  • Clapp, C. M., 1898. Relation of the axis of the embryo to the first cleavage plane. Biol. Lectures M. B. L., Wood's Holl, Mass., pp. 139-151.
  • Newman, H. H., 1907. Spawning behavior and sexual dimorphism in Fundulus heteroclitus and allied fish. Biol. Bull., 12: 314-348.
  • Newman, H. H., 1915. Development and heredity in heterogenic teleost hybrids. J. Exp. Zool., 18: 511-576.
  • Newman, H. H., 1918. Hybrids between Fundulus and mackerel. A study of paternal heredity in heterogenic hybrids. J. Exp. Zool., 26: 391-421.
  • Nicholas, J. S., 1927. The application of experimental methods to the study of developing Fundulus embryos. Proc. Nat. acad. Sci., 13: 695-698.
  • Nicholas, J. S., and J. M. Oppenheimer, 1942. Regulation and reconstitution in Fundulus. J. Exp. Zool., 90: 127-157.
  • Oppenheimer, J. M., 1937. The normal stages of Fundulus heteroclitus. Anat. Rec., 68: 1-15.
  • Russell, A., 1939. Pigment inheritance in the Fundulus-Scomber hybrid. Biol. Bull.,., 77: 423-431.
  • Solberg, A. N., 1938. The development of a bony fish. Prog. Fish. Cult., no. 40, pp. 1-19.
  • Sumner, F. B., 1903. A study of early fish development. Experimental and morphological. Arch. f. Entw., 17: 92-149.
  • Trinkaus, J. P., and J. W. Drake, 1956. Exogenous control of morphogenesis in isolated Fundulus blastoderms by nutrient chemical factors. J. Exp. Zool., 132: 311-347.
  • Wilson, H. V., 1889. The embryology of the sea bass (Serranus atrarius). Bull. U. S. Fish Comm., 9: 209-278.
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Not available.

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This fish preys on mosquito larvae and is therefore occasionally used instead of harmful pesticides to control mosquito population. They are an extremely important food source for many larger fish, which are valuable commercially, and for wading birds and seabirds (National Oceanic and Atmospheric Administration Coastal Services Center, 2001). For some of these birds mummichogs compose up to 95% of their entire diet. One final economic value of the mummichog is its use as bait for recreational fishing (Abraham, 1985).

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Economic Uses

Comments: Suitable for use in sophisticated genetic analyses; has been used to study endocrine mechanisms in osmoregulation; has played a central role in embryology in U.S.; being used to address questions about the evolutionary significance of protein polymorphism (Powers 1989). Used extensively and increasingly as a bioassay organism in toxicological investigations and as an indicator of marine water quality (Eisler 1986).

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Importance

aquarium: commercial
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Preparation of Eggs for Sectioning

Eggs stripped from a female fish into diluted sea water (70% fresh water, 30% sea water) retain the morphological characteristics of freshly-extruded eggs, including the yolk platelets, oil drops, membranes, etc. A micropyle is present, but it must be observed before removal of the chorionic jelly.

  • Agassiz, A., and C. O. Whitman, 1885. The development of osseous fishes. I. The pelagic stages of young fishes. Mem. Mus. Comp. Zool., Harvard, 14: no. 1, part 1, pp. 1-56.
  • Agassiz, A., and C. O. Whitman, 1889. The development of osseous fishes. Ii. The preembryonic stages of development. Part First. The history of the egg from fertilization to cleavage. Mem. Mus. Comp. Zool., Harvard, 14: no. 2, part 2, pp. 1-40.
  • Breder, C. M., Jr., 1948. Field Book of Marine Fishes of the Atlantic Coast from Labrador to Texas. G. P. Putnam's Sons, New York. (Rev. ea.).
  • Clapp, C. M., 1891. Some points in the development of the toad-fish (Batrachus tau). J. Morph., 5: 494-501.
  • Clapp, C. M., 1898. Relation of the axis of the embryo to the first cleavage plane. Biol. Lectures M. B. L., Wood's Holl, Mass., pp. 139-151.
  • Newman, H. H., 1907. Spawning behavior and sexual dimorphism in Fundulus heteroclitus and allied fish. Biol. Bull., 12: 314-348.
  • Newman, H. H., 1915. Development and heredity in heterogenic teleost hybrids. J. Exp. Zool., 18: 511-576.
  • Newman, H. H., 1918. Hybrids between Fundulus and mackerel. A study of paternal heredity in heterogenic hybrids. J. Exp. Zool., 26: 391-421.
  • Nicholas, J. S., 1927. The application of experimental methods to the study of developing Fundulus embryos. Proc. Nat. acad. Sci., 13: 695-698.
  • Nicholas, J. S., and J. M. Oppenheimer, 1942. Regulation and reconstitution in Fundulus. J. Exp. Zool., 90: 127-157.
  • Oppenheimer, J. M., 1937. The normal stages of Fundulus heteroclitus. Anat. Rec., 68: 1-15.
  • Russell, A., 1939. Pigment inheritance in the Fundulus-Scomber hybrid. Biol. Bull.,., 77: 423-431.
  • Solberg, A. N., 1938. The development of a bony fish. Prog. Fish. Cult., no. 40, pp. 1-19.
  • Sumner, F. B., 1903. A study of early fish development. Experimental and morphological. Arch. f. Entw., 17: 92-149.
  • Trinkaus, J. P., and J. W. Drake, 1956. Exogenous control of morphogenesis in isolated Fundulus blastoderms by nutrient chemical factors. J. Exp. Zool., 132: 311-347.
  • Wilson, H. V., 1889. The embryology of the sea bass (Serranus atrarius). Bull. U. S. Fish Comm., 9: 209-278.
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Source: Egg Characteristics and Breeding Season for Woods Hole Species

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Permanent Total Preparations

Eggs to be sectioned must be dechorionated before fixation, so that fluids can penetrate to the interior. (For details of this process, see the paper by Nicholas, 1927.) The following schedule for dehydration and embedding is useful.

1. Fix in Bouin's or Zenker's solution, 12-24 hours.

2. Dehydrate as usual through the alcohol series (up to and including 95% alcohol), leaving the eggs in each for one hour.

3. Absolute alcohol, two hours&emdash;use several changes.

4. Equal parts absolute alcohol and amyl acetate, two hours. 5. Amyl acetate, 24-48 hours.

6. Equal parts amyl acetate and paraffin, 12 hours (incubate at 30°).

7. Three changes of infiltrating paraffin (15 minutes in each); embed in 56-58° paraffin.

  • Agassiz, A., and C. O. Whitman, 1885. The development of osseous fishes. I. The pelagic stages of young fishes. Mem. Mus. Comp. Zool., Harvard, 14: no. 1, part 1, pp. 1-56.
  • Agassiz, A., and C. O. Whitman, 1889. The development of osseous fishes. Ii. The preembryonic stages of development. Part First. The history of the egg from fertilization to cleavage. Mem. Mus. Comp. Zool., Harvard, 14: no. 2, part 2, pp. 1-40.
  • Breder, C. M., Jr., 1948. Field Book of Marine Fishes of the Atlantic Coast from Labrador to Texas. G. P. Putnam's Sons, New York. (Rev. ea.).
  • Clapp, C. M., 1891. Some points in the development of the toad-fish (Batrachus tau). J. Morph., 5: 494-501.
  • Clapp, C. M., 1898. Relation of the axis of the embryo to the first cleavage plane. Biol. Lectures M. B. L., Wood's Holl, Mass., pp. 139-151.
  • Newman, H. H., 1907. Spawning behavior and sexual dimorphism in Fundulus heteroclitus and allied fish. Biol. Bull., 12: 314-348.
  • Newman, H. H., 1915. Development and heredity in heterogenic teleost hybrids. J. Exp. Zool., 18: 511-576.
  • Newman, H. H., 1918. Hybrids between Fundulus and mackerel. A study of paternal heredity in heterogenic hybrids. J. Exp. Zool., 26: 391-421.
  • Nicholas, J. S., 1927. The application of experimental methods to the study of developing Fundulus embryos. Proc. Nat. acad. Sci., 13: 695-698.
  • Nicholas, J. S., and J. M. Oppenheimer, 1942. Regulation and reconstitution in Fundulus. J. Exp. Zool., 90: 127-157.
  • Oppenheimer, J. M., 1937. The normal stages of Fundulus heteroclitus. Anat. Rec., 68: 1-15.
  • Russell, A., 1939. Pigment inheritance in the Fundulus-Scomber hybrid. Biol. Bull.,., 77: 423-431.
  • Solberg, A. N., 1938. The development of a bony fish. Prog. Fish. Cult., no. 40, pp. 1-19.
  • Sumner, F. B., 1903. A study of early fish development. Experimental and morphological. Arch. f. Entw., 17: 92-149.
  • Trinkaus, J. P., and J. W. Drake, 1956. Exogenous control of morphogenesis in isolated Fundulus blastoderms by nutrient chemical factors. J. Exp. Zool., 132: 311-347.
  • Wilson, H. V., 1889. The embryology of the sea bass (Serranus atrarius). Bull. U. S. Fish Comm., 9: 209-278.
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Source: Egg Characteristics and Breeding Season for Woods Hole Species

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Methods of Observation

Fix the eggs in Stockard's solution (formalin, 5 parts; glacial acetic acid, 4 parts; glycerine, 6 parts; distilled water, 85 parts). This turns the protoplasm white but leaves the yolk transparent. The fixative may be used as a preservative, or the material can be transferred to 10% formalin after two days.

  • Agassiz, A., and C. O. Whitman, 1885. The development of osseous fishes. I. The pelagic stages of young fishes. Mem. Mus. Comp. Zool., Harvard, 14: no. 1, part 1, pp. 1-56.
  • Agassiz, A., and C. O. Whitman, 1889. The development of osseous fishes. Ii. The preembryonic stages of development. Part First. The history of the egg from fertilization to cleavage. Mem. Mus. Comp. Zool., Harvard, 14: no. 2, part 2, pp. 1-40.
  • Breder, C. M., Jr., 1948. Field Book of Marine Fishes of the Atlantic Coast from Labrador to Texas. G. P. Putnam's Sons, New York. (Rev. ea.).
  • Clapp, C. M., 1891. Some points in the development of the toad-fish (Batrachus tau). J. Morph., 5: 494-501.
  • Clapp, C. M., 1898. Relation of the axis of the embryo to the first cleavage plane. Biol. Lectures M. B. L., Wood's Holl, Mass., pp. 139-151.
  • Newman, H. H., 1907. Spawning behavior and sexual dimorphism in Fundulus heteroclitus and allied fish. Biol. Bull., 12: 314-348.
  • Newman, H. H., 1915. Development and heredity in heterogenic teleost hybrids. J. Exp. Zool., 18: 511-576.
  • Newman, H. H., 1918. Hybrids between Fundulus and mackerel. A study of paternal heredity in heterogenic hybrids. J. Exp. Zool., 26: 391-421.
  • Nicholas, J. S., 1927. The application of experimental methods to the study of developing Fundulus embryos. Proc. Nat. acad. Sci., 13: 695-698.
  • Nicholas, J. S., and J. M. Oppenheimer, 1942. Regulation and reconstitution in Fundulus. J. Exp. Zool., 90: 127-157.
  • Oppenheimer, J. M., 1937. The normal stages of Fundulus heteroclitus. Anat. Rec., 68: 1-15.
  • Russell, A., 1939. Pigment inheritance in the Fundulus-Scomber hybrid. Biol. Bull.,., 77: 423-431.
  • Solberg, A. N., 1938. The development of a bony fish. Prog. Fish. Cult., no. 40, pp. 1-19.
  • Sumner, F. B., 1903. A study of early fish development. Experimental and morphological. Arch. f. Entw., 17: 92-149.
  • Trinkaus, J. P., and J. W. Drake, 1956. Exogenous control of morphogenesis in isolated Fundulus blastoderms by nutrient chemical factors. J. Exp. Zool., 132: 311-347.
  • Wilson, H. V., 1889. The embryology of the sea bass (Serranus atrarius). Bull. U. S. Fish Comm., 9: 209-278.
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Source: Egg Characteristics and Breeding Season for Woods Hole Species

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Preparation of Cultures

To remove the sticky outer jelly layer, roll the eggs on a piece of filter paper or paper towel until the surface of the outer membrane is left smooth and clean. This same procedure should be followed daily for stock cultures, in order to prevent clumping of the eggs.

For experimental work, where it is essential to obtain development as nearly normal as possible, the eggs are usually examined uncovered in shallow depression slides; they may be manipulated with hair loops. For classroom study, when the eggs are to be observed over long periods of time and a specific orientation is desired, either of the following methods is suggested: (1) Place the eggs in sea water in special culture slides having a depression of 1.7 to 1.8 mm. (slightly less than the diameter of the eggs); it is then possible to roll the eggs to the desired position by moving the coverslip. (2) If these special slides are not available, the eggs may be placed in a drop of sea water on an ordinary glass slide and covered with a very thin, flexible sheet of mica; water is then withdrawn (using lens or filter paper) until capillary attraction causes a pressure on the egg, so that it can be rotated as in the previous method.

Recently, Trinkaus and Drake (1956) have described a method for the in vitro culture of Fundulus blastoderms isolated from the subjacent periblast and yolk mass.

  • Agassiz, A., and C. O. Whitman, 1885. The development of osseous fishes. I. The pelagic stages of young fishes. Mem. Mus. Comp. Zool., Harvard, 14: no. 1, part 1, pp. 1-56.
  • Agassiz, A., and C. O. Whitman, 1889. The development of osseous fishes. Ii. The preembryonic stages of development. Part First. The history of the egg from fertilization to cleavage. Mem. Mus. Comp. Zool., Harvard, 14: no. 2, part 2, pp. 1-40.
  • Breder, C. M., Jr., 1948. Field Book of Marine Fishes of the Atlantic Coast from Labrador to Texas. G. P. Putnam's Sons, New York. (Rev. ea.).
  • Clapp, C. M., 1891. Some points in the development of the toad-fish (Batrachus tau). J. Morph., 5: 494-501.
  • Clapp, C. M., 1898. Relation of the axis of the embryo to the first cleavage plane. Biol. Lectures M. B. L., Wood's Holl, Mass., pp. 139-151.
  • Newman, H. H., 1907. Spawning behavior and sexual dimorphism in Fundulus heteroclitus and allied fish. Biol. Bull., 12: 314-348.
  • Newman, H. H., 1915. Development and heredity in heterogenic teleost hybrids. J. Exp. Zool., 18: 511-576.
  • Newman, H. H., 1918. Hybrids between Fundulus and mackerel. A study of paternal heredity in heterogenic hybrids. J. Exp. Zool., 26: 391-421.
  • Nicholas, J. S., 1927. The application of experimental methods to the study of developing Fundulus embryos. Proc. Nat. acad. Sci., 13: 695-698.
  • Nicholas, J. S., and J. M. Oppenheimer, 1942. Regulation and reconstitution in Fundulus. J. Exp. Zool., 90: 127-157.
  • Oppenheimer, J. M., 1937. The normal stages of Fundulus heteroclitus. Anat. Rec., 68: 1-15.
  • Russell, A., 1939. Pigment inheritance in the Fundulus-Scomber hybrid. Biol. Bull.,., 77: 423-431.
  • Solberg, A. N., 1938. The development of a bony fish. Prog. Fish. Cult., no. 40, pp. 1-19.
  • Sumner, F. B., 1903. A study of early fish development. Experimental and morphological. Arch. f. Entw., 17: 92-149.
  • Trinkaus, J. P., and J. W. Drake, 1956. Exogenous control of morphogenesis in isolated Fundulus blastoderms by nutrient chemical factors. J. Exp. Zool., 132: 311-347.
  • Wilson, H. V., 1889. The embryology of the sea bass (Serranus atrarius). Bull. U. S. Fish Comm., 9: 209-278.
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Procuring Gametes

Strip the eggs into a clean four-inch fingerbowl which has been moistened with filtered sea water. Strip the milt into a small amount of sea water, and mix the suspension with the eggs in 1/4 inch of sea water. The eggs should be inseminated as soon as possible after they are obtained from the body of the female. After 30-45 minutes, change the sea water and leave the eggs in about a 1/4 to 1/2-inch depth of sea water. Keep the fingerbowl covered with a glass plate to prevent evaporation; do not allow the eggs to clump or accumulate in one spot. The water should be changed at least twice daily.

  • Agassiz, A., and C. O. Whitman, 1885. The development of osseous fishes. I. The pelagic stages of young fishes. Mem. Mus. Comp. Zool., Harvard, 14: no. 1, part 1, pp. 1-56.
  • Agassiz, A., and C. O. Whitman, 1889. The development of osseous fishes. Ii. The preembryonic stages of development. Part First. The history of the egg from fertilization to cleavage. Mem. Mus. Comp. Zool., Harvard, 14: no. 2, part 2, pp. 1-40.
  • Breder, C. M., Jr., 1948. Field Book of Marine Fishes of the Atlantic Coast from Labrador to Texas. G. P. Putnam's Sons, New York. (Rev. ea.).
  • Clapp, C. M., 1891. Some points in the development of the toad-fish (Batrachus tau). J. Morph., 5: 494-501.
  • Clapp, C. M., 1898. Relation of the axis of the embryo to the first cleavage plane. Biol. Lectures M. B. L., Wood's Holl, Mass., pp. 139-151.
  • Newman, H. H., 1907. Spawning behavior and sexual dimorphism in Fundulus heteroclitus and allied fish. Biol. Bull., 12: 314-348.
  • Newman, H. H., 1915. Development and heredity in heterogenic teleost hybrids. J. Exp. Zool., 18: 511-576.
  • Newman, H. H., 1918. Hybrids between Fundulus and mackerel. A study of paternal heredity in heterogenic hybrids. J. Exp. Zool., 26: 391-421.
  • Nicholas, J. S., 1927. The application of experimental methods to the study of developing Fundulus embryos. Proc. Nat. acad. Sci., 13: 695-698.
  • Nicholas, J. S., and J. M. Oppenheimer, 1942. Regulation and reconstitution in Fundulus. J. Exp. Zool., 90: 127-157.
  • Oppenheimer, J. M., 1937. The normal stages of Fundulus heteroclitus. Anat. Rec., 68: 1-15.
  • Russell, A., 1939. Pigment inheritance in the Fundulus-Scomber hybrid. Biol. Bull.,., 77: 423-431.
  • Solberg, A. N., 1938. The development of a bony fish. Prog. Fish. Cult., no. 40, pp. 1-19.
  • Sumner, F. B., 1903. A study of early fish development. Experimental and morphological. Arch. f. Entw., 17: 92-149.
  • Trinkaus, J. P., and J. W. Drake, 1956. Exogenous control of morphogenesis in isolated Fundulus blastoderms by nutrient chemical factors. J. Exp. Zool., 132: 311-347.
  • Wilson, H. V., 1889. The embryology of the sea bass (Serranus atrarius). Bull. U. S. Fish Comm., 9: 209-278.
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Care of Adults

Both eggs and sperm are obtained by "stripping": the fish is held firmly in one hand while gentle pressure is applied to its abdomen with the thumb and forefinger of the other hand. As these fingers are drawn towards the anus of the fish, the pressure forces out the gametes. If the fish is held in front of a strong light source during the stripping process, the eggs may be seen passing through the oviduct which runs along the anal fin.

  • Agassiz, A., and C. O. Whitman, 1885. The development of osseous fishes. I. The pelagic stages of young fishes. Mem. Mus. Comp. Zool., Harvard, 14: no. 1, part 1, pp. 1-56.
  • Agassiz, A., and C. O. Whitman, 1889. The development of osseous fishes. Ii. The preembryonic stages of development. Part First. The history of the egg from fertilization to cleavage. Mem. Mus. Comp. Zool., Harvard, 14: no. 2, part 2, pp. 1-40.
  • Breder, C. M., Jr., 1948. Field Book of Marine Fishes of the Atlantic Coast from Labrador to Texas. G. P. Putnam's Sons, New York. (Rev. ea.).
  • Clapp, C. M., 1891. Some points in the development of the toad-fish (Batrachus tau). J. Morph., 5: 494-501.
  • Clapp, C. M., 1898. Relation of the axis of the embryo to the first cleavage plane. Biol. Lectures M. B. L., Wood's Holl, Mass., pp. 139-151.
  • Newman, H. H., 1907. Spawning behavior and sexual dimorphism in Fundulus heteroclitus and allied fish. Biol. Bull., 12: 314-348.
  • Newman, H. H., 1915. Development and heredity in heterogenic teleost hybrids. J. Exp. Zool., 18: 511-576.
  • Newman, H. H., 1918. Hybrids between Fundulus and mackerel. A study of paternal heredity in heterogenic hybrids. J. Exp. Zool., 26: 391-421.
  • Nicholas, J. S., 1927. The application of experimental methods to the study of developing Fundulus embryos. Proc. Nat. acad. Sci., 13: 695-698.
  • Nicholas, J. S., and J. M. Oppenheimer, 1942. Regulation and reconstitution in Fundulus. J. Exp. Zool., 90: 127-157.
  • Oppenheimer, J. M., 1937. The normal stages of Fundulus heteroclitus. Anat. Rec., 68: 1-15.
  • Russell, A., 1939. Pigment inheritance in the Fundulus-Scomber hybrid. Biol. Bull.,., 77: 423-431.
  • Solberg, A. N., 1938. The development of a bony fish. Prog. Fish. Cult., no. 40, pp. 1-19.
  • Sumner, F. B., 1903. A study of early fish development. Experimental and morphological. Arch. f. Entw., 17: 92-149.
  • Trinkaus, J. P., and J. W. Drake, 1956. Exogenous control of morphogenesis in isolated Fundulus blastoderms by nutrient chemical factors. J. Exp. Zool., 132: 311-347.
  • Wilson, H. V., 1889. The embryology of the sea bass (Serranus atrarius). Bull. U. S. Fish Comm., 9: 209-278.
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Wikipedia

Mummichog

The mummichog, Fundulus heteroclitus, is a small killifish found along the Atlantic coast of the United States and Canada. Also known as mummies, gudgeons, and mud minnows, these fish inhabit brackish and coastal waters including estuaries and salt marshes. The species is noted for its hardiness and ability to tolerate highly variable salinity, temperature fluctuations from 6 °C to 35 °C (43 °F to 95 °F), very low oxygen levels (down to 1 mg/L), and heavily polluted ecosystems. As a result, the mummichog is a popular research subject in embryological, physiological, and toxicological studies. It is also the first fish ever sent to space, aboard Skylab in 1973.

Etymology and taxonomy[edit]

Fundulus comes from fundus, meaning bottom, from the fish's habit of swimming near muddy bottoms. Heteroclitus means irregular or unusual. The type specimen was first described by Carl Linnaeus in 1766, from near Charleston, South Carolina. Past scientific names for this species include Cobitis heteroclita, Fundulus fasciatus, Fundulus pisculentus, and Fundulus nigrofasciatus.[4]

The mummichog belongs to the order Cyprinodontiformes, and the family Fundulidae. There are two subspecies: F. h. heteroclitus (Linnaeus, 1766), in the south and F. h. macrolepidotus (Walbaum, 1792) in the north.

Common name[edit]

The name mummichog is derived from a Narragansett term which means "going in crowds", which reflects the mummichog's strong shoaling tendency.[5] Colloquial names include mummy, killie, salt water minnow, mud minnow, mud dabbler, marsh minnow, brackish water chub, gudgeon, and common killifish. Some of these terms may lead to confusion: the term minnow should be reserved for species of the Cyprinidae family, the mudminnows are members of the Umbridae family, and the name gudgeon is used for various bottom-dwelling species of cyprinid, eleotrid, and ptereleotrid fishes, none of which belongs to the same family as the fundulid mummichog.

Description[edit]

The body of the mummichog is elongate but thick, with a deep caudal peduncle. Usual length is 75 to 90 mm (3 to 3.5 inches) but maximum lengths of 130 to 150 mm (5 to 6 inches) are possible. The mouth is upturned and the lower jaw protrudes when the mouth is closed. Pectoral and tail fins are round. Mummichogs have 10-13 dorsal fin rays, 9-12 anal fin rays, 16-20 pectoral fin rays. Males have larger dorsal and anal fins than females. There is no lateral line on the body, but lateral line pores are present on the head. The colour is variable (and may even change in shade within the same individual when placed near different backgrounds)[6][7] but is generally olive-brown or olive-green. There can be vertical bars on the sides that are thin, wavy, and silvery. Colors are more intense in males during the reproductive season, as they become dark olive-green on the back, steel-blue on the sides with about 15 silvery bars, and yellow or orange-yellow on the underside; the dorsal fin is mottled and a small eyespot may be present near the rear edge. Females tend to be paler, without bars or the intense yellow on the belly, and their dorsal fin is uniformly coloured.

Adults of the two subspecies can be distinguished based on slight morphological[8] and genomic[9] differences. Further, eggs of the northern subspecies have filaments (adhesive chorionic fibrils) that eggs of the southern subspecies lack. While the northern subspecies deposits eggs in the sand, the southern subspecies often deposits eggs inside empty mussel shells.[10][11]

The mummichog is very similar to the banded killifish, Fundulus diaphanus, and indeed the two species have been known to interbreed.[12] The two species may overlap in their choice of habitat, but in general the banded killifish is more commonly found in freshwater, which is not the case for the mummichog. The banded killifish tends to have thin dark bars on a light side, whereas in the mummichog the bars are thin and light on a dark side. Internally, the banded killifish has 4-7 gill rakers, as opposed to 8-12 in the mummichog.

Distribution[edit]

This species ranges along the Atlantic coast of North America, from Gaspé Peninsula, Anticosti Island and Port au Port Bay in the north to northeastern Florida in the south. It is present on Sable Island, 175 km (109 mi) southeast of the closest point of mainland Nova Scotia in the Atlantic Ocean.[13] The approximate geographical division between the two subspecies lies in New Jersey, Delaware and Virginia.

Introduced populations have established themselves on the Atlantic coast of Portugal and southwestern Spain, starting in the 1970s[14][15][16] and some have now reached the western Mediterranean basin.[17] There may also be introduced populations in Hawaii and the Philippines.[18] As bait fish, mummichogs are sometimes released in freshwater habitats, where they can survive, and there have been reports of individuals in New Hampshire ponds, as well as the upper Ohio River and Beaver River (Pennsylvania).[19]

Habitat[edit]

The mummichog is a common fish in coastal habitats such as salt marshes, muddy creeks, tidal channels, brackish estuaries, eelgrass or cordgrass beds, and sheltered shorelines. It can be found within coastal rivers but seldom beyond the head of tide. A few landlocked populations may exist in freshwater lakes close to shore, for example on Digby Neck, Nova Scotia.[20]

Diet[edit]

Mummichogs are omnivorous. Analyses of their stomach contents have found diatoms, amphipods and other crustaceans, molluscs, fish eggs (including their own species), very small fish, insect larvae, and bits of eelgrass.

Physiology[edit]

This fish is well known for its ability to withstand a variety of environmental conditions.[21] They can survive temperatures between 6 °C and 35 °C; even within the same tidal cycle they can tolerate rapid temperature changes from 15 °C to 30 °C.[22] They are among fish species most tolerant of salinity changes (euryhaline).[23] Mummichog larvae can grow in salinities ranging from 0.4 to 100 parts per thousand, the latter being about three times the normal salinity of seawater. Adult mummichogs tolerate low oxygen levels down to 1 mg/L, at which they resort to aquatic surface respiration (breathing in the surface layer of water, richer in oxygen because of contact with air) to survive.[24][25] They can even survive for a few hours in moist air outside of water, breathing air directly.

Populations have developed resistance to methylmercury, kepone, dioxins, polychlorinated biphenyl, and polyaromatic hydrocarbons.[26] One study[27] has looked at the genomic variation exhibited by mummichogs populations living in Newark Bay, New Bedford Harbor, and the Elizabeth River (Virginia) (in some areas heavily polluted with polychlorinated biphenyls and creosote, a complex mixture containing dioxin-like chemicals) and has found that about 20% of their genes were modified as compared to populations living in clean sites.

Behavior[edit]

Mummichogs live in dense shoals that can include several hundred individuals.

During cold winter months in the northern parts of their range, mummichogs move to upstream tidal pools, where they burrow into the mud at depths up to 20 cm (8 inches) to overwinter.[28][29] They can also bury themselves in mud if they are caught in a drying tidal pool between spring tides. Alternatively, they can travel short distances on land to get back to the sea.[30]

Reproduction[edit]

Spawning takes place from spring through fall. In the southern most populations, up to eight spawnings are possible in a season. Spawning takes place most often at high tide and when the moon is new or full. Maximal spawning occurs when high spring tides coincide with night,[31] though spawning during the day remains possible.

During courtship, males may pursue females, and females may attract males by turning on their sides near the bottom and flicking their tails. A male and female may swim together for a while, after which the male crowds the female against a rock or a plant and clasps her: the male's larger dorsal and anal fins curve around the female's body. Fingerlike projections that develop on the male's scales behind and below the dorsal fin may help the male maintain contact with the female. The pair quivers vigorously and eggs and sperm are released.[32]

The eggs are pale yellow, about 2 mm in diameter, and strongly adhesive. During a spawning event, a female can deposit up to 740 eggs in separate clutches of 10 to 300 eggs at a time.[10] The eggs adhere to plants, algal mats, empty mussel shells, sand, or mud at sites that are reached by water only at high spring tides.[10] Eggs therefore develop while exposed to moist air, and they hatch when the next high spring tides reach them.[33][34][35]

As opposed to their northern counterparts, the southern subspecies have eggs that lack filaments (adhesive chorionic fibrils)[36] and they often deposit those eggs inside empty mussel shells.[10][11] The two subspecies are also distinguished based on slight morphological[8] and genomic[9] differences.

Most mummichogs become sexually mature when two years old, around 38 mm in length. Normal lifespan is four years.[10]

Parasites[edit]

Mummichogs are hosts to a parasitic fluke, Homalometron pallidum, which has a complex lifecycle involving the aquatic snail, Ecrobia truncata.[37] Other parasite species reported in mummichogs include 10 protozoans, eight trematodes, one nematode, two acanthocephalans, and two crustaceans.[38] A study in New Jersey found that mummichogs heavily infested with the digenean gill parasite Ascocotyle phagicola, spent more time near the surface and exhibited conspicuous behaviors such as jerking, an example of a parasite affecting the behavior of its host in a way beneficial to the parasite, as conspicuous behaviors near the surface make the fish more likely to be noticed by predatory wading birds, the next host in the parasite's life cycle.[39]

Interest to humans[edit]

Mummichogs are sold as bait in sport fisheries for marine species such as summer flounder and bluefish, or even sometimes for freshwater species.[22]

Mummichogs readily eat mosquito larvae and attempts have been made to use them as biocontrol agents of mosquito populations.[22]

Scientific Utility[edit]

Mummichogs are considered an important environmental model organism because of their ability to tolerate various extremes of chemical (pollution, etc.) and physical (temperature, salinity, oxygen, etc.) conditions. They are relatively abundant in nature and can be easily captured, transported and reared in laboratory facilities. They are commonly used in scientific studies of stress biology,[40] thermal physiology and toxicology, and have also been studied in the contexts of evolutionary biology, developmental biology, endocrinology, cancer biology, and chronobiology (study of circadian rhythms).[41][42] With the successful sequencing and assembly of the full killifish genome,[43] they serve as a premier scientific model for studying biochemical and physiological responses to varying environmental conditions.[44]

Their remarkable ability to tolerate various extremes of temperature and salinity has made them popular subjects in scientific studies of toxicology. For decades the killifish has been a useful laboratory model for toxicological studies that include exposures to single chemicals, chemical mixtures, and complex contaminated media. It is sometimes the only fish species found in severely polluted and oxygen-deprived waterways, such as the Elizabeth River in Virginia and, in New Jersey, the Hackensack River and the Arthur Kill.

Killifish eggs are used in developmental studies and when teaching embryology because the eyes, the beating heart, and the different stages of ontogenesis can be easily examined. Embryos are also extremely durable and easy to manipulate in the laboratory.

Mummichogs were the first fish sent to space.[45] In 1973 a couple of them were flown in a plastic bag aquarium aboard Skylab, during the Skylab 3 mission. In the absence of gravity the fish at first exhibited an unusual swimming behavior: they constantly pitched forward and therefore described tight circles. However, by day 22 of the mission they swam normally. Fifty eggs at an advanced stage of development had also been taken on board, and 48 of them hatched during the flight. The hatchlings swam normally.[46] More experiments with mummichogs in space followed as part of the Apollo-Soyuz Test Project[47] and as part of a biological package aboard the Bion 3 satellite.

References[edit]

  1. ^ NatureServe (2013). "Fundulus heteroclitus". IUCN Red List of Threatened Species. Version 2014.3. International Union for Conservation of Nature. Retrieved March 12, 2015. 
  2. ^ Nicolas Bailly (2014). Nicolas Bailly, ed. "Fundulus heteroclitus heteroclitus (Linnaeus, 1766)". FishBase. World Register of Marine Species. Retrieved March 12, 2015. 
  3. ^ Nicolas Bailly (2014). Nicolas Bailly, ed. "Fundulus heteroclitus macrolepidotus (Walbaum, 1792)". FishBase. World Register of Marine Species. Retrieved March 12, 2015. 
  4. ^ Scott, W.B., and Crossman, E.J. 1973. Freshwater fishes of Canada. Bulletin 184 of the Fisheries Research Board of Canada, Ottawa.
  5. ^ "Mummichog." Merriam-Webster.com. Merriam-Webster, n.d. Web. 6 Feb. 2014. <http://www.merriam-webster.com/dictionary/mummichog>
  6. ^ Connolly, C.J. 1925. Adaptive changes in shades and color of Fundulus. Biological Bulletin (Woods Hole) 48: 56-77.
  7. ^ Bagnara, J.T., and Hadley, M.E. 1973. Chromatophores and color change. Prentice-Hall, New Jersey.
  8. ^ a b Able, K.W. and Felley, J.D. 1986. Geographical variation in Fundulus heteroclitus: tests for concordance between egg and adult morphologies. American Zoologist 26: 145-157.
  9. ^ a b Brown, B.L. and Chapman, R.W. 1991. Gene flow and mitochondrial DNA variation in the killifish, Fundulus heteroclitus. Evolution 45: 1147-1161.
  10. ^ a b c d e Coad, B.W. 1995. Encyclopedia of Canadian Fishes. Canadian Museum of Nature, Ottawa, 928p.
  11. ^ a b Taylor, M.H. 1986. Environmental and endocrine influences on reproduction of Fundulus heteroclitus. American Zoologist 26: 159-171.
  12. ^ Hubbs, C.L., Walker, B.W., and Johnson, R.E. 1943. Hybridization in nature between species of American cyprinodont fishes. Contributions to the Laboratory of Vertebrate Biology of the University of Michigan 23: 21 p.
  13. ^ Garside, E.T. 1969. Distribution of insular fishes of Sable Island, Nova Scotia. Journal of the Fisheries Research Board of Canada 26: 1390-1392.
  14. ^ Hernando, J.A., 1975. Nuevas localidades de Valencia hispanica (Pisces: Ciprinodontidae) en el Suroeste de España. Doñana Acta Vertebrata 2: 265-267.
  15. ^ Coelho, M., Gomes, J., and Ré, P.B. 1976. Valencia hispanica, a new fish to Portugal. Arquivos do Museu Bocage 6: 1-3.
  16. ^ Gutiérrez-Estrada, J.C., Prenda, J., Oliva, F, and Fernandez-Delgado, C. 1998. Distribution and habitat preferences of the introduced mummichog Fundulus heteroclitus (Linneaus) in South-western Spain. Estuarine Coastal and Shelf Science 46: 827-835.
  17. ^ Gisbert, E., and Lopez, M.A. 2007. First record of a population of the exotic mummichog, Fundulus heteroclitus (L., 1766) in the Mediterranean Sea basin (Ebro River delta). Journal of Fish Biology 71: 1220-1224.
  18. ^ Fish Base Mummichog distribution
  19. ^ USGS Nonindigenous aquatic species database Mummichog occurrences
  20. ^ Klawe, W.L. 1957. Common mummichog and newt in a lake on Digby Neck, Nova Scotia. Canadian Field-Naturalist 71: 154-155.
  21. ^ Burnett, K.G., Bain, L.J., Baldwin, D.S., et al. 2007. Fundulus as the premier teleost model in environmental biology: Opportunities for new insights using genomics. Comparative Biochemistry and Physiology (D) 2: 257-266.
  22. ^ a b c 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 and Wildlife Service Biological Reports 82 (11.40): 23 p. http://www.nwrc.usgs.gov/wdb/pub/species_profiles/82_11-040.pdf
  23. ^ Whitehead, A. 2010. The evolutionary radiation of diverse osmotolerant physiologies in killifish (Fundulus sp.). Evolution 64: 2070-2085.
  24. ^ Wannamaker, C.M., and Rice, J.A. 2000. Effects of hypoxia on movements and behavior of selected estuarine organisms from the southeastern United States. Journal of Experimental Marine Biology and Ecology 249: 145-163.
  25. ^ Stierhoff, K.L., Targett, T.E., and Grecay, P.A. 2003. Hypoxia tolerance of the mummichog: the role of access to the water surface. Journal of Fish Biology 63: 580-592.
  26. ^ Weis, J,S. 2002. Tolerance to environmental contaminants in the mummichog, Fundulus heteroclitus. Human and Ecological Risk Assessment 8: 933-953.
  27. ^ Whitehead, A., Galvez, F., Zhang, S., Williams, L.M., and Oleksiak, M.F. 2011. Functional genomics of physiological plasticity and local adaptation in killifish. Journal of Heredity 102: 499-511. doi: 10.1093/jhered/esq077
  28. ^ Chidester, F.E. 1920. The behavior of Fundulus heteroclitus in the salt marshes of New Jersey. American Naturalist 54: 244-245.
  29. ^ Raposa, K. 2003. Overwintering habitat selection by the mummichog, Fundulus heteroclitus, in a Cape Cod (USA) salt marsh. Wetlands Ecology and Management 11: 175-182.
  30. ^ Mast, S. O. 1915. The behavior of Fundulus, with especial reference to overland escape from tide-pools and locomotion on land. Journal of Animal Behavior 5: 341-350. http://dx.doi.org/10.1037/h0075747
  31. ^ Taylor, M.H., Leach, G.J., DiMichele, L., Levithan, W.H., and Jacob, W.F. 1979. Lunar spawning cycle in the mummichog, Fundulus Heteroclitus (Pisces: Cyprinodontidae). Copeia 1979: 291-297.
  32. ^ Newman, H.H. 1907. Spawning behavior and sexual dimorphism in Fundulus heteroclitus and allied fish. Biological Bulletin (Woods Hole) 12: 314-345.
  33. ^ Taylor, M.H., DiMichele, L., and Leach, G.J. 1977. Egg stranding in the life cycle of the mummichog Fundulus heteroclitus. Copeia: 1977: 397-399.
  34. ^ Taylor, M.H., and DiMichele, L. 1983. Spawning site utilization in a Delaware population of Fundulus heteroclitus (Pisces: Cyprinodontidae). Copeia 1983: 719-725.
  35. ^ Taylor, M.H. 1999. A suite of adaptations for intertidal spawning. American Zoologist 39: 313-320.
  36. ^ Morin, R.P. and Able, K.W. 1983. Patterns of geographic variation in the egg morphology of the fundulid fish, Fundulus heteroclitus. Copeia 1983: 726-740.
  37. ^ Stunkard, Horace W. (1964). "The morphology, life history and systematics of the digenetic trematode Homalometron pallidum Stafford 1904". The Biological Bulletin 126 (1): 163–173. doi:10.2307/1539426. 
  38. ^ Hoffman, G.L. 1967. Parasites of North American freshwater fishes. University of California Press, Berkeley, 486 pp.
  39. ^ Santiago Bass, C. and Weis, J.S. 2009. Conspicuous behaviour of Fundulus heteroclitus associated with high digenean metacercariae gill abundances. Journal of Fish Biology 74: 763-772.
  40. ^ Schulte, Patricia M. 2014. What is environmental stress? Insights from fish living in a variable environment. Journal of Experimental Biology 217: 23-34.
  41. ^ Kavaliers, M. 1980. Social groupings and circadian activity of the killifish, Fundulus heteroclitus. Biological Bulletin 158: 69-76.
  42. ^ Kavaliers, M., and Abbott, F.S. 1977. Rhythmic colour change of the killifish, Fundulus heteroclitus. Canadian Journal of Zoology 55: 553-561.
  43. ^ http://www.ncbi.nlm.nih.gov/genome/
  44. ^ Lister, AL, Van Der Kraak, GJ, Rutherford, R, MacLatchy, D. 2011. Fundulus heteroclitus: ovarian reproductive physiology and the impact of environmental contaminants. Comparative Biochemistry and Physiology, Part C: Toxicology & Pharmacology 154:4 278-287.
  45. ^ Reebs, S.G. (2009) Fish in space Retrieved 12 December 2014.
  46. ^ von Baumgarten, R.J., Simmonds, R.C., Boyd, J.F., and Garriott, O.K. 1975. Effects of prolonged weightlessness on the swimming pattern of fish aboard Skylab 3. Aviation Space and Environmental Medicine 46: 902-906.
  47. ^ Hoffman, R.B., Salinas, G.A., and Baky, A.A. 1977. Behavioral analyses of killifish exposed to weightlessness in the Apollo-Soyuz test project. Aviation Space and Environmental Medicine 48: 712-717.
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Names and Taxonomy

Taxonomy

Comments: A number of subspecies have been described, but only two (heteroclitus and macrolepidotus) have been recognized in most recent studies. The genus Fundulus was removed from Atheriniformes:Cyprinodontidae and placed in Cyprinodontiformes:Fundulidae by Parenti (1981); pending confirmation based on other character suites, this change was not accepted in the 1991 AFS checklist (Robins et al. 1991). Studies of mitochondrial DNA of subspecies heteroclitus and macrolepidotus of the U.S. Atlantic coast indicate that these two taxa exhibit secondary intergradation in northern New Jersey, Long Island, and in Chesapeake and Delaware bays (Gonzalez-Villasenor and Powers 1990), Able and Felley 1986). See Brown and Chapman (1991) and Powers (1989) for additional genetic studies of these populations. See Wiley (1986) for a study of the evolutionary relationships of Fundulus topminnows based on morphological characters. See Cashner et al. (1992) for an allozyme-based phylogenetic analysis of the genus Fundulus.

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