Banded killifish

The banded killifish (Fundulus diaphanus) is a North American species of temperate freshwater killifish belonging to the genus Fundulus of the family Fundulidae. Its natural geographic range extends from Newfoundland to South Carolina, and west to Minnesota, including the Great Lakes drainages.[2] This species is the only freshwater killifish found in the northeastern United States. While it is primarily a freshwater species, it can occasionally be found in brackish water.[3]

Banded killifish
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Actinopterygii
Order: Cyprinodontiformes
Family: Fundulidae
Genus: Fundulus
Species:
F. diaphanus
Binomial name
Fundulus diaphanus
(Lesueur, 1817)
Subspecies
Range of the banded killifish
Synonyms[1]
  • Hydrargira diaphana Lesueur, 1817
  • Hydrargira multifaciata Lesueur, 1817
  • Fundulus multifaciatus (Lesueur, 1817)
  • Fundulus menona Jordan & Copeland, 1877
  • Fundulus extensus Jordan & Gilbert, 1882

Etymology

The common name, "banded killifish", commonly refers to the distinct black and white vertical bandings found along their sides. The Latin genus name Fundulus is the diminutive of fundus, which means "bottom," and the specific name diaphanus means "transparent" in Greek.[4]

Description
Closeup of a banded killifish to show anatomical features.

The banded killifish has an olive color on the dorsal surface and white coloring on the ventral. The throat and fins are yellowish. There are vertical blackish and silver-white stripes along the sides.

There are 13–15 rays on a banded killifish's dorsal fin and 10–12 on the anal fin. The homocercal tail is slightly convex or rounded. It also has a small pelvic fin along the abdomen. The body is slender and elongated, with a flattish side and flattened head and small terminal mouth positioned for surface feeding.[5] The banded killifish has a row of small sharp teeth lining their upper and lower jaws. It does not have a lateral line along the side but does have 39 to 43 cycloid scales in the lateral series.[6]

The average banded killifish ranges from 10 to 13 cm in length and weighs a few grams.[4][1] The females tend to grow larger than the males. They also have darker bands on the sides. In contrast, the males have pale gray bands that are close together. The numbers of bands on the dorsal fin of a banded killifish is useful in determining the sex of this species.

Range and distribution

The banded killifish is widely distributed throughout eastern North America, ranging from South Carolina to as far north as the Atlantic Provinces. They are also found in the eastern part of Montana to Minnesota and throughout the suitable habitats of the Great Lakes watershed of southern Ontario to Lake Superior. Due to biogeographical isolation and limitation of potential for range expansion, banded killifish species were assigned a status of special concern by the Committee on the Status of Endangered Wildlife in Canada (COSEWIC).[4]

There are two subspecies, the eastern banded killifish Fundulus diaphanus diaphanus and the western banded killifish Fundulus diaphanus menona. The former is classified as an injurious aquatic invasive species in Ohio. Intergrades occur in the Saint Lawrence and Lake Erie drainages.[7] The eastern subspecies is larger.[8]

Habitat and ecology

Adults range from 2 to 3 years in age. Banded killifish are schooling fish, usually traveling in groups of 3–6 individuals, while the juveniles travel in groups of 8–12. The fish are most often found in the shallow and quiet areas of clear lakes, ponds, rivers, and estuaries with sandy gravel or muddy bottoms and with abundant aquatic vegetation. The sand and gravel provides hatchlings and juveniles with places to hide when threatened by predatory fish such as the largemouth bass Micropterus salmoides, northern pike Esox lucius, bluegill Lepomis macrochirus, and trout. Because the banded killifish is small, it generally does not venture into deeper waters, where it would be vulnerable to predation as well as unable to swim in the fast currents. However, adult banded killifish have been observed to travel into deep bodies of water to feed. Banded killifish often congregate near aquatic vegetation, as it provides protection as well as breeding habitat.

Banded killifish are euryhaline, but they usually inhabit freshwater streams and lakes. The largest adult recorded, observed in Indian Bay, Canada, was 12.8 cm. People have used banded killifish as fish bait. Most people do not favor them as pets because they require a high level of maintenance and therefore do not survive well in an aquarium setting. They are important to aquatic ecosystems because they are a food source for larger fish such as largemouth bass, northern pike, and trout. They are also a food source for birds such as belted kingfisher (Megaceryle alcyon), common merganser (Mergus merganser), and herons.

In January 2005 the banded killifish was listed as a vulnerable species under the Newfoundland Labrador Endangered Species Act and the Canada Species at Risk Act (SARA). This species is facing habitat degradation due to industrial development, motorized watercraft activities, and removal of aquatic vegetations.[9]

Diet

Banded killifish have been observed to feed at all levels of the water column. The adults feed on a variety of items such as insects, nymphs, mollusks, turbellarians, and other small crustaceans. Mosquito larvae are also a popular food source. In contrast, the smaller individuals are limited to fewer items such as chironomid larvae, cladocerans, copepods, and midge larvae. Both young and adult banded killifish have been observed to feed mostly in the afternoon.[9]

Reproduction and life cycle

Banded killifish are commonly observed to spawn in dense aquatic vegetation because they practice external fertilization where the female lays her eggs that are equipped with adhesive threads that adhere to plants. Spawning occurs from June to mid-August in shallow waters. During the spawning season, the males go through a color change phase. They develop a bright blue patch near the anal fin. In addition, the lower portion of the body changes to a bright blue color.

Spawning occurs at water temperatures of 21˚ C to 23˚ C. The male chooses a site in the shallow part of the water and protects it from other males.[4] When a female appears, the male will court the female and fight with the other prospecting males. The female will emit one egg while the male pursues her. Once together, the female emits 10 eggs that falls onto the bottom or gets attached to aquatic plants in the chosen spawn area. The male will continue to pursue the female until the female have laid 50 to 100 eggs.

A single female may lay several clutches of eggs during one summer. After the eggs have been fertilized, both the parents will leave and go their separate ways; the eggs do not receive parental care. 6-7 millimeter fries emerge within 10 to 12 days depending on the temperature of the water. They reach maturity at approximately 1 year with an average length of 6 cm. Banded killifish can live for a little over 2 years.[9] However, there have been some that have been observed to live up to 3 years.

Behavior

Shoaling behavior

Banded killifish form group shoals.[2][1] The conditions in which shoaling occurs, the individuals who comprise the shoal, and the size of the shoal are all highly variable. Shoaling can be understood through a cost and benefit analysis. The costs and benefits of group membership are influenced by food availability and predatory risk. Bigger groups allow for better predatory protection, but are not as conducive to foraging because the food will be distributed amongst all of the group members.[10] For smaller groups, there is not as effective predatory protection, but in regards to foraging each individual will have access to more food.[10]

The nutritional state of banded killifish influences individual's decisions to shoal or not. Food-deprived individuals spent more time by themselves and not in shoals. When food-deprived individuals were found in shoals their shoals were not any smaller than that of a well-fed banded killifish. Hungry banded killifish are more likely to leave a shoal than a well-fed individual.[11] In the presence of a food stimulus, group size decreases, so that each individual does not have to compete with others for access to these resources.[12] When presented with food odour, individuals were less attracted to neighbors, but when presented with a predatory stimulus, they were more attracted to neighbors and formed tighter shoals. When in the presence of both stimuli, the group size is intermediate.[13]

In the presence of a predatory stimulus, banded killifish decrease the amount of food attempts and duration of feeding posture. This is done to allocate more energy to predatory vigilance and avoidance. In both the presence and absence of predatory stimuli, banded killifish feeding rate for individuals is independent of shoal size.[14] Lone individuals incur additional cost because they do not gain the improved predatory protection granted by shoal formation, and thus will attempt to join larger shoals rather than smaller shoals. When deciding whether to join a shoal or not, banded killifish value predatory protection over foraging opportunities.[14]

Predator protection

In the presence of a predatory stimulus, shoal size increases to allow for greater protection via the dilution effect. From the dilution effect, each individual will have a lesser chance of being attacked by a predator, since there are many other individuals in the group. In a larger group, there is a larger chance that a predator will attack a different shoalmate than it will attack the individual of interest.[10] Individuals who stray from the shoal have an increased mortality risk. This leads to selection for grouping.[15]

Besides the dilution effect, shoaling also benefits the individual because of group vigilance.[10] To best protect against predation, the banded killifish must be on the lookout to detect predators. In a group, each individual can spend less time looking out for predators because others in the shoal can share in this responsibility. With increasing shoal size, each individual spends less time being vigilant, but with more individuals the group increases overall vigilance and therefore benefit each individual in the shoal. When one individual detects a predator, an alarm call is made to alert the shoal, so that the individuals can act to protect themselves from predatory attack.[16]

To improve predator avoidance, banded killifish will often take refuge. Frightened fish in the presence of a predatory stimulus will hide in a foodless refuge. This preference is modulated by individual body size. Larger individuals spend more time in the refuge than their smaller conspecifics. Smaller banded killifish individuals are more prone to risk than their larger counterparts.[17]

Shoaling preferences

When choosing which shoals to join, banded killifish often choose to join ones that consist of others with similar phenotypes. In order to achieve this, banded killifish have shown the ability to discriminate between individuals within the species. Banded killifish distinguish individuals by body coloration phenotype. Banded killifish are likely to enter shoals with those of similar body color in order to maximize predator avoidance.[18] An individual of different color might present as a contrast. This contrast is likely to make the shoal stand out and gain the predator's attention. Banded killifishes' level of preference for similarly colored shoalmates differs in the presence and absence of a predatory stimulus.[18]

Banded killifish use body coloration to determine if an individual has been parasitized or not. Black spots indicate the presence of a parasite, whereas the absence of these spots indicates an unparasitized individual. Both parasitized and unparasitized individuals preferred to join unparasitized shoals. The preference for unparasitized shoals increases with an increase in parasite load of a parasitized individual.[19]

Shoaling preferences in banded killifish differ in shoal species composition, shoal size, and individual body size within the shoal. The shift in preferences is triggered by predatory stimuli.[20] In the presence of predatory stimuli, a banded killifish individual will prefer the bigger conspecific shoal, but only as long as the shoal is homogenous in individual fish size. If the size of individual fish were different, they would join a shoal of similarly sized fish regardless of shoal size. This preference is maintained by predatory threat. Body size is a more significant determinant of shoaling preference than shoal size and species composition.[20]

Banded killifish have been found to engage in heterospecific shoaling with other species of killifish as well as other fish. There are many situational factors that determine if the banded killifish individual would want to engage in heterospecific shoaling. If body size is similar, then banded killifish prefer conspecific shoals as opposed to heterospecific ones. Size is a sorting mechanism for conspecific as opposed to heterospecific shoaling.[21]

References
  1. Froese, Rainer and Pauly, Daniel, eds. (2019). "Fundulus diaphina" in FishBase. September 2019 version.
  2. Page, Lawrence M. and Brooks M. Burr (1991), Freshwater Fishes, p. 216, Houghton Mifflin, New York. ISBN 0-395-91091-9
  3. Werner, Robert G. (2004), Freshwater Fishes of the Northeastern United States, p. 206, Syracuse University Press, Syracuse, New York. ISBN 0-8156-3020-4.
  4. Chippett, Jamie D. 2003. "Update COSEWIC status report on the banded killifish Fundulus diaphanus, Newfoundland population in Canada", in COSEWIC assessment and update status report on the banded killifish Fundulus diaphanous in Canada. Committee on the Status of Endangered Wildlife in Canada. Ottawa. 1-21 pp
  5. "The Virtual Aquarium". Archived from the original on 2011-02-11. Retrieved 2011-05-06.
  6. "Iowa DNR Fish and Fishing". Archived from the original on 2012-03-14. Retrieved 2011-05-06.
  7. "Banded Killifish (Fundulus diaphanus) - FactSheet". USGS. Retrieved 23 August 2014.
  8. "Banded killifish (Fundulus diaphanus)". Arkive. Archived from the original on 26 September 2011. Retrieved 23 August 2014.
  9. Paulson, N. and Jay T. Hatch
  10. Nicholas B. Davies; John R. Krebs; Stuart A. West (2012-04-02). An introduction to behavioural ecology (4th ed.). Oxford: Wiley-Blackwell. pp. 148–163. ISBN 978-1405114165.
  11. Hensor, E.M.A.; J.-G. J Godin; D.J. Hoare; J. Krause (April 2003). "Effects of nutritional state on the shoaling tendency of banded killifish, Fundulus diaphanus, in the field". Animal Behaviour. 65 (4): 663–669. doi:10.1006/anbe.2003.2075. S2CID 53177142.
  12. Hoare, D.J.; I.D. Couzin; J-G J. Godin; J. Krause (January 2004). "Context-dependent group size choice in fish". Animal Behaviour. 67 (1): 155–164. doi:10.1016/j.anbehav.2003.04.004. S2CID 2244463.
  13. Nicholas B. Davies; John R. Krebs; Stuart A. West (2012-04-02). An introduction to behavioural ecology (4th ed.). Oxford: Wiley-Blackwell. pp. 171–173. ISBN 978-1405114165.
  14. Goudin, Jean-Guy J. (1986). "Risk of predation and foraging behaviour in shoaling banded killifish (Fundulus diaphanus)". Canadian Journal of Zoology. 64 (8): 1675–1678. doi:10.1139/z86-251.
  15. Morgan, M. Joanne; Jean-Guy J. Godin (January–December 1985). "Antipredator Benefits of Schooling Behaviour in a Cyprinodontid Fish, the Banded Killifish (Fundulus diaphanus)". Zeitschrift für Tierpsychologie. 70 (3): 246. doi:10.1111/j.1439-0310.1985.tb00515.x.
  16. Abrahams, M.V.; Godin, J.-G.J.; Classon, L.J. (1 January 1988). "Group Vigilance and Shoal Size in a Small Characin Fish". Behaviour. 104 (1): 29–40. doi:10.1163/156853988x00584.
  17. Dowling, Lisa M.; Jean-Guy J. Godin (April 2002). "Refuge use in a killifish: influence of body size and nutritional state". Canadian Journal of Zoology. 80 (4): 782–788. doi:10.1139/z02-036.
  18. McROBERT, SCOTT P; BRADNER, JOSHUA (1 September 1998). "The influence of body coloration on shoaling preferences in fish". Animal Behaviour. 56 (3): 611–615. doi:10.1006/anbe.1998.0846. PMID 9784209. S2CID 24479889.
  19. Krause, Jens; Godin, Jean-Guy J. (26 April 2010). "Influence of Parasitism on Shoal Choice in the Banded Killifish (Fundulus diaphanus, Teleostei, Cyprinodontidae)". Ethology. 102 (1): 40–49. doi:10.1111/j.1439-0310.1996.tb01102.x.
  20. Krause, Jens; Jean-Guy J. Godin (26 April 2010). "Shoal Choice in the banded Killifish (Fundulus diaphanus, Teleostei, Cyprinodontidae): Effects of Predation Risk, Fish Size, Species Composition and Size of Shoals". Ethology. 98 (2): 128–136. doi:10.1111/j.1439-0310.1994.tb01063.x.
  21. Blakeslee, Carrie; Ruhl, Nathan; Currie, Warren; McRobert, Scott (1 May 2009). "Shoaling preferences of two common killifish (Fundulus heteroclitus and F. diaphanus) in the laboratory and in the field: A new analysis of heterospecific shoaling". Behavioural Processes. 81 (1): 119–125. doi:10.1016/j.beproc.2009.02.012. PMID 19429206. S2CID 23225234.
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