Etheostomatinae

Last updated

Etheostomatinae
Etheostoma blennioides.jpg
Etheostoma blennioides
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Actinopterygii
Order: Perciformes
Family: Percidae
Subfamily: Etheostomatinae
Agassiz, 1850 [1]
Genera

see text

Etheostomatinae is a species-rich subfamily of freshwater ray-finned fish, the members of which are commonly known as the darters. The subfamily is part of the family Percidae which also includes the perches, ruffes and pikeperches. The family is endemic to North America. It consists of three to five different genera and well over 200 species. A common name for these fish in southern Indiana is pollywog.

Contents

Characteristics

Species within the Etheostomatinae are all small fish, mostly less than 11 centimetres (4.3 in) in length, and their bodies are slightly compressed or fusiform in shape. They have two pterygiophores between the first and second dorsal fins which do not have spines and a reduced swimbladder which may be completely lacking. [2] The common name "darter" owes to the behavior of the fish, which dart around their benthic habitat. They are sexually dimorphic; most species have males with bright colors and patterning, particularly when breeding. These colors and patterns are used to attract females and allow for recognition of species, as the colors and patterns are particular to each species. Many species use typical sites for spawning and they care for their eggs and fry. They have evolved a variety of methods of depositing eggs and these include burying them, which may be the basal habit as it is found in all genera, as well as attaching eggs to a substrate and egg clustering. [3]

Distribution

Etheostomatinae darters are endemic to North America where they are found in the Mississippi River basin and the drainages of the Great Lakes, Hudson Bay, the seaboards of the Atlantic and the Gulf of Mexico and the Pacific coast of Mexico. [3]

Taxonomy

Fishbase recognises five genera as follows: [4] [5]

However, Fishbase places Crystallaria within the subfamily Percinae [6] while the 5th Edition of Fishes of the World regard it as a subgenus of Ammocrypta and Nothonotus as a subgenus of Etheostoma. [2]

Life history and morphology

Phylogeny and diversity

Etheostomatinae, as a species rich and diverse sub-family within the family Percidae of freshwater ray-finned fishes, is composed of multiple genera including Ammocrypta, Crystalluria, Etheostoma, Nothonotus, and Percina, all of which have been observed as monophyletic groupings through molecular phylogenies. [7]

Speciation events can occur to a culmination of factors as both sympatric and allopatric speciation is observed across darter evolutionary history. Factors related to spatiotemporal habitat stability and life history strategies can play into speciation and genetic differentiation of darters, such as dispersal ability and gene flow in headwater reproductive habitats impacting lineage divergence . It was seen that habitat preference and isolation is a major factor contributing to speciation in darters. [8] Speciation events are common across species of this subfamily, one example includes Barcheek darters, an ancient monophyletic grouping residing in the eastern highlands which under time-calibrated molecular phylogenetic analysis of the mitochondrial genome has presented five novel cryptic species previously thought to be associated with Etheostoma basilare. [9]

Hybridization has been accounted for multiple times across Etheostomatinae. It is more common between species with similar egg-laying behavioral mechanisms and within the same phylogenetic clade, with sperm viability being the limiting factor of hybridization. Sperm viability has also been witnessed to decrease as temperatures increase as an extraneous factor. Other factors impacting the viability of hybridization include life history of sympatry as opposed to allopatry, habitat preference, similar spawning periods, and duration of sperm viability. Species known to bury their eggs have been viewed to reproduce under sympatric conditions while egg-guarding species typically reproduce under allopatric conditions. Hybridization is predominantly found within species of large geographic ranges, larger populations, and egg-burying reproductive mechanisms. [10]

Morphological features

Darters are composed of fusiform shaped bodies that are typically less than 11 centimeters in length, and characteristically host a reduced or lack an air bladder, allowing them to reside in benthic regions of lakes, streams, and rivers. A prominent feature across many species of darters is their diverse coloration. Color preferences have been found in common darter predators such as the largemouth bass, imposing selective pressures on certain species of darters limiting nuptial coloration. Chromatic species of darters are more prominent in fast flowing habitats with coarse substrate present within the water column and low flow accumulation, more light penetration, and fewer predators present. Species with blue fins are present in habitats supporting coarse substrate along with low flow accumulation, while blue body and red fins are commonly present in rifle ecosystems, and species with red coloration on the body are present in small streams. [11]

The size of darters can be linked to increased fitness as larger darters tend to grow faster, produce more offspring, mature earlier at a larger size, and have longer lifespans along with reproductive lifespans. Diversity of life species traits across darters have been linked to influences such as reproductive behavior, geographic range, and population size of a species. [12]

Coloration has been viewed to be heightened through ingestion of carotenoids, which could explain how coloration is phenotypically correlated to fitness across chromatic species of Etheostomatinae. [11]

Another prominent feature held by females across darter lineage is the presence of gential papillae, fleshy protuberances between the anus and anal fin and are associated with egg laying abilities. Phylogenetic analyses expressed that the common ancestor of Etheostomatinae buried eggs with the help of tube-like appendage with a genital pore, a pore that releases the females eggs, held at the dorsal-posterior end known as tube papillae, leading researchers to believe this is the ancestral state of darters. Tube papillae are present in majority egg laying species across Etheostomatinae, with the exception of the genus Nothonotus, which instead have a mound-tube papillae allowing females to completely bury themselves as they lay eggs under the substrate. Egg clustering species tend to express complexes of pleated rosette papillae with ventral genital pores allowing the females to attach their eggs to the underhangs of rocks and other objects. [13]

Behavior

Egg laying and reproductive behavior

Selective traits may vary across species and genera, female darters have been observed to lay 500 to 1000 eggs sequentially, alluding to a promiscuous mating system with limited female mate preference and no parental care. [14] Observations have been made that the majority of species across the diverse genus Etheostoma and that they attach eggs to their surroundings for protection. These egg attaching species mate when a female selects a location above the substrate such as a plant or rock where she will deposit her eggs, the male then mounts the female and as they vibrate the females attach eggs to her location of choosing while the male deposits sperm fertilizing the eggs. Other egg-laying behavioral characteristics include egg-guarding in which a female are courted by a male guarding a cavity, typically under flat stones, the couple then invert as the female lays eggs and the cavity ceiling as the male fertilizes them. When burying eggs, the male initiates with a courtship dance, in response the female wiggles until she is partially buried in the substrate. The male then mounts the female and similar to other species of darter, they vibrate as the females release eggs into her desired location while the male deposits sperm atop the eggs. Typically no parental care is associated with egg-burying species across the genus. [13]

While parental care is not common within the darter subfamily, is has developed across Etheostomatinae's evolutionary history multiple times, once in the clade Goneaperca, and a more novel evolutionarily derived clade Nothonotus. Male parental care across these clades have expressed evolutionary reversals back to the lack of care for their offspring multiple times. There are at least 250 species of darters with known reproductive methodologies across the Etheostomatinae sub-family with roughly 40 species experiencing paternal care and the rest showing no parental care at all. [15]

Sexual selection

Males, while predominantly giving no parental care, do invest in courtship along with egg guarding among certain species. Intra-sexual and intersexual interactions affect speciation via sexual selection, predominantly via behavioral isolation through male aggression more so than female biases, likely due to the idea that males expending more energy into courtship reproduction than females. Male darters have been found to hold female preferences and mate choice and aggression correlated to their coloration differences expressing biases regarding intra and intersexual selection. Coloration has been viewed to be correlated more with male aggression as opposed to previously suggested female selective biases expressing that the color divergence is due to intra-sexual mechanisms in certain species lacking parental care, and that male coloration and aggressiveness may have coevolved. [14] It has been viewed across species exhibiting parental care that achromatic coloration can be a trade off between paternal care and energy allocation for reproduction. [16] Nuptial color preference has been observed across females of the darter species E. barrenses supporting the idea that female choice can drive the evolution of male coloration; while this may not apply to all species of darters, it does help support the general idea behind coloration and how it develops under the realm of sexual selection. [17]

Related Research Articles

<span class="mw-page-title-main">Rainbow darter</span> Species of fish

The rainbow darter is a small species of freshwater ray-finned fish, a darter from the subfamily Etheostomatinae, part of the family Percidae, which also contains the perches, ruffes and pikeperches. It is native to North America where it is found in small, fast-moving streams and small to medium-sized rivers. It grows to 2 to 3 inches in length. The species is very sensitive to pollution and silt, staying in clean, pollution-free water. The rainbow darter is easily identified by three dark spots on the back, and blue and orange in the dorsal and anal fins.

The Cumberland snubnose darter is a species of freshwater ray-finned fish, a darter from the subfamily Etheostomatinae, part of the family Percidae, which also contains the perches, ruffes and pikeperches. This species is found in the middle Cumberland River drainage in Tennessee, Kentucky, Virginia, North Carolina, Georgia, and Alabama. It is absent in reaches above the Big South Fork, rare in North Carolina, and absent in western tributaries of the Tennessee River. While research on the ecology of E. atripinne is not extensive, what is known is they are usually found in small to medium freshwater streams in gravel riffle areas where their eggs can attach to the substrate and be left unguarded. E. atripinne can be found within a wide range of depths in its environment, leading its being classified as benthopelagic. While its global status is secure, the American Fisheries Society labels it with a status of “Special Concern”.

The emerald darter, is a species of freshwater ray-finned fish, a darter from the subfamily Etheostomatinae, part of the family Percidae, which also contains the perches, ruffes and pikeperches. It is found only in northern and eastern Kentucky and northeastern Tennessee.

The corrugated darter is a species of freshwater ray-finned fish, a darter from the subfamily Etheostomatinae, part of the family Percidae, which also contains the perches, ruffes and pikeperches. It is endemic to the eastern United States only known from Caney Fork, Tennessee, where it has been shown to have as many as five subgroups. They are active hunters which prey on insects, fry, and possibly young snails. Multiple species and subspecies of these darters can coexist where substrate is varied, so this must be carefully preserved to accommodate many similar species. With the warming of spring, E. basilare enters its breeding cycle and males develop their dimorphic coloration. Like most darters, it hugs the stream bed, using its body shape and fin posture to maintain its position.

The splendid darter is a species of freshwater ray-finned fish, a darter from the subfamily Etheostomatinae, part of the family Percidae, which also contains the perches, ruffes and pikeperches. It is endemic to the Barren River system in south-central Kentucky and north-central Tennessee. This species is usually found in small rocky pools on the sides of creeks and rivers.

<span class="mw-page-title-main">Orangefin darter</span> Species of fish

The orangefin darter is a species of freshwater ray-finned fish, a darter from the subfamily Etheostomatinae, part of the family Percidae, which also contains the perches, ruffes and pikeperches. It is found in Barren River and Green River systems in Tennessee and Kentucky.

<span class="mw-page-title-main">Buffalo darter</span> Species of fish

The buffalo darter is a species of freshwater ray-finned fish, a darter from the subfamily Etheostomatinae, part of the family Percidae, which also contains the perches, ruffes and pikeperches. It is found in the tributaries of the lower Duck and lower Tennessee Rivers. It is distinguished from other darter species by the presence of eight anal rays, as well breeding males having a unique hump behind the head reminiscent of a buffalo.

<span class="mw-page-title-main">Bluebreast darter</span> Species of fish

The bluebreast darter is a small species of freshwater ray-finned fish, a darter from the subfamily Etheostomatinae, part of the family Percidae, which also contains the perches, ruffes and pikeperches. It is endemic to the eastern United States from New York and Illinois to Tennessee and North Carolina.

<span class="mw-page-title-main">Greenfin darter</span> Species of fish

The greenfin darter is a species of freshwater ray-finned fish, a darter from the subfamily Etheostomatinae, part of the family Percidae, which also contains the perches, ruffes and pikeperches. It is endemic to the eastern United States.

The stripetail darter is a species of freshwater ray-finned fish, a darter from the subfamily Etheostomatinae, part of the family Percidae, which also contains the perches, ruffes and pikeperches. It is endemic to the eastern United States. It is found in small rivers and streams in the states of Tennessee, Ohio, Kentucky, Illinois, Alabama, Georgia, and Mississippi. Males are a golden-orange color with black barring on the fins, and grow to a length of about 2.8 in (7 cm). This fish feeds on midge larvae and other small invertebrates. It breeds in the spring; eggs are attached to the substrate, often under slab rocks, where they are guarded by the male. The population trend of this fish seems to be stable and it is a common species with numerous sub-populations over a wide range, and the International Union for Conservation of Nature has assessed its conservation status as being of "least concern".

Nothonotus microlepidum, the smallscale darter, is a species of freshwater ray-finned fish, a darter from the subfamily Etheostomatinae, part of the family Percidae, which also contains the perches, ruffes and pikeperches. It is endemic to the southeastern United States. It occurs in the lower Cumberland River drainage in the states of Kentucky and Tennessee. It inhabits shallow riffles with gravel substrates in small rivers. Breeding habits of the smallscale darter are typical of the E. maculatum group in that females deposit large masses of eggs on the undersides of rocks to be protected by the males. This species is a fairly large, deep-bodied fish with dark background coloration mixed with bright red spots scattered alongside the body. Nuptial males of this species also have deep green fins with orange margins on the spinous dorsal and caudal fins, with the soft dorsal fins having a dark coloration moving marginally to yellow-orangish center with black margins. This species can reach a length of 7.2 centimetres (2.8 in) TL though most only reach about 4.8 centimetres (1.9 in).

<span class="mw-page-title-main">Snubnose darter</span> Species of fish

The snubnose darter is a species of freshwater ray-finned fish, a darter from the subfamily Etheostomatinae, part of the family Percidae, which also contains the perches, ruffes and pikeperches. It is endemic to the southeastern United States.

<span class="mw-page-title-main">Tippecanoe darter</span> Species of fish

The Tippecanoe darter is a species of freshwater ray-finned fish, a darter from the subfamily Etheostomatinae, part of the family Percidae, which also contains the perches, ruffes and pikeperches. It is endemic to the eastern United States.

The wounded darter is a species of freshwater ray-finned fish, a darter from the subfamily Etheostomatinae, part of the family Percidae, which also contains the perches, ruffes and pikeperches. It is endemic to the eastern United States. Its range includes the upper Tennessee River drainage, western Virginia, western North Carolina, and eastern Tennessee. Its typical habitat is among boulders or coarse rubble and cobble, often with overhanging ledges, in medium to large slow-moving rivers. It feeds on small insect larvae, especially midge larvae. Spawning occurs when the water warms up in late spring. Females deposit clutches of adhesive eggs on the underside of rock ledges or slabs, and the male guards the nest. The population trend of this fish seems to be decreasing slowly but it is a relatively common species with numerous sub-populations, and the International Union for Conservation of Nature has assessed its conservation status as being of "least concern". The greatest threat comes from impoundment, and a management plan is in place, including captive breeding.

<span class="mw-page-title-main">Gilt darter</span> Species of fish

The gilt darter is a species of freshwater ray-finned fish, a darter from the subfamily Etheostomatinae, part of the family Percidae, which also contains the perches, ruffes and pikeperches. It can be found in a number of states in the Mississippi River drainage of the United States although it has been extirpated from some river systems in which it was at one time present, mostly due to siltation and pollution problems. Males are more colorful than females and can grow to a length of about 9 cm (3.5 in). It is a benthic fish that feeds primarily on small aquatic insect larvae. Males form territories during the breeding season in late spring and early summer. Spawning typically takes place at the upper ends of riffles with sandy and gravelly bottoms interspersed with larger cobbles. Some organisations are endeavouring to conserve populations of the gilt darter and re-introduce it to states where the fish has been extirpated but suitable habitat still exists.

<span class="mw-page-title-main">Blackbanded darter</span> Species of fish

The blackbanded darter is a species of freshwater ray-finned fish, a darter from the subfamily Etheostomatinae, part of the family Percidae, which also contains the perches, ruffes and pikeperches. It is native to the river systems of the southeastern United States where it is found in the states of Tennessee, Georgia, Florida, Mississippi, Alabama, and Louisiana. It lives over sandy or gravelly bottoms in smaller rivers and streams and its color varies depending on the different habitats in which it lives. It feeds on small insect larvae and is itself preyed on by larger fish. It spawns between February and June depending on locality. It is generally a common fish throughout most of its range.

The bronze darter is a species of freshwater ray-finned fish, a darter from the subfamily Etheostomatinae, part of the family Percidae, which also contains the perches, ruffes and pikeperches. It is restricted to the Coosa and Tallapoosa River systems in Georgia, Alabama, and southeastern Tennessee in the United States. It feeds on insect larvae and other small invertebrates, and spawns in April and May. It is a common fish within its range and the International Union for Conservation of Nature has classified its conservation status as being of "least concern". However, it is susceptible to environmental disturbance and an ongoing management plan has been put in place to protect its habitat.

The sickle darter is a recently identified freshwater ray-finned fish, a darter from the subfamily Etheostomatinae, part of the family Percidae, which also contains the perches, ruffes and pikeperches. It is closely related to a well-known darter, Percina macrocephala. Discovered in 2007 in the upper Tennessee River drainage, the sickle darter is around 90.5 mm (3.56 in) in length and is a slender bodied, freshwater and benthopelagic fish that most likely feeds on small crayfish and mayflies, since it shares a large mouth and long snout with its sister species. The known distribution of the sickle darter is the upper drainage of the Tennessee River of Tennessee, North Carolina and Virginia. It is distinguished from all other Percina darters except its sister species P. macrocephala by a dark suborbital bar and a black bar subtending a medial black spot on the base of the caudal fin. It stays in mostly rocky, sandy, or silty substrates in clear creeks or small rivers in the Appalachian Mountains. The sickle darter spawns in late winter in gravel shoals. It also has large scales which make it different from macrocephala. It is known to be extirpated from most of its home habitat mainly because of agricultural practices that cause siltation and turbidity which causes most populations of the sickle darter to be widely scattered. But where it is found, it can be observed with regularity. Taking all this into consideration, the sickle darter does not have a formal conservation status under any federal or state law although its closely related species is considered a species of concern by the TWRA. Future management plans should include finding more locations of the sickle darter and decreasing further habitat destruction in known distributions by stating it as a fish of concern by state law.

<span class="mw-page-title-main">Christmas darter</span> Species of ray-finned fish, a darter from the subfamily Etheostomatinae

The Christmas darter i is a species of ray-finned fish, a darter from the subfamily Etheostomatinae, part of the family Percidae which includes the perches, ruffes and pike-perches. It is found in the Savannah, Ogeechee and Altamaha catchment areas in South Carolina and Georgia. It is a small but colorful fish, typically less than 5 cm (2 in) in total length, with blackish, red and green streaks on its flanks. Green and red are associated with Christmas, hence the common name Christmas darter. Like other darters, it lives on or close to the stream-bed, in riffles and vegetation-laden stretches of small rivers, creeks and spring-fed streams. It feeds on aquatic insects and probably breeds in the spring, but its biology and behavior are poorly known.

<i>Nothonotus</i> Genus of fishes

Nothonotus is a genus or subgenus of freshwater ray-finned fish, a darter from the subfamily Etheostomatinae, part of the family Percidae, which also contains the perches, ruffes and pikeperches. It is endemic to the southeastern United States. First proposed as a sub-genus of Etheostoma in 1988, there is still debate regarding the appropriate taxonomic rank of Nothonotus in the literature. Darter species comprise more than 180 of the Percidae taxa.

References

  1. Richard van der Laan; William N. Eschmeyer & Ronald Fricke (2014). "Family-group names of Recent fishes". Zootaxa. 3882 (2): 001–230.
  2. 1 2 J. S. Nelson; T. C. Grande; M. V. H. Wilson (2016). Fishes of the World (5th ed.). Wiley. pp. 448–450. ISBN   978-1-118-34233-6. Archived from the original on 2019-04-08. Retrieved 2020-09-19.
  3. 1 2 Carol A. Stepien & Amanda Haponski (2015). "Taxonomy, Distribution, and Evolution of the Percidae". In Patrick Kestemont; Konrad Dabrowski & Robert C. Summerfelt (eds.). Biology and Culture of Percid Fishes. Springer, Dordrecht. pp. 3–60. doi:10.1007/978-94-017-7227-3_1. ISBN   978-94-017-7227-3.
  4. Froese, Rainer, and Daniel Pauly, eds. (2019). "Etheostomatinae" in FishBase . December 2019 version.
  5. Eschmeyer, William N.; Fricke, Ron & van der Laan, Richard (eds.). "Genera in the family Etheostomatinae". Catalog of Fishes . California Academy of Sciences . Retrieved 19 September 2020.
  6. Froese, Rainer and Pauly, Daniel, eds. (2019). Species of Crystallaria in FishBase . December 2019 version.
  7. Near, Thomas J. (February 2002). "Phylogenetic Relationships of Percina (Percidae: Etheostomatinae)". Copeia. 2002 (1): 1–14. doi:10.1643/0045-8511(2002)002[0001:proppe]2.0.co;2. ISSN   0045-8511.
  8. Fluker, Brook L.; Kuhajda, Bernard R.; Harris, Phillip M. (2014-09-19). "The influence of life‐history strategy on genetic differentiation and lineage divergence in darters (Percidae: Etheostomatinae)". Evolution. 68 (11): 3199–3216. doi:10.1111/evo.12505. ISSN   0014-3820.
  9. Hollingsworth Jr, Phillip R.; Near, Thomas J. (January 2009). "Temporal patterns of diversification and microendemism in Eastern Highland endemic barcheek darters (Percidae: Etheostomatinae)". Evolution. 63 (1): 228–243. doi:10.1111/j.1558-5646.2008.00531.x. ISSN   0014-3820.
  10. Keck, Benjamin P.; Near, Thomas J. (2009-12-29). "Patterns of Natural Hybridization in Darters (Percidae: Etheostomatinae)". Copeia. 2009 (4): 758–773. doi:10.1643/ci-09-008. ISSN   0045-8511.
  11. 1 2 Ciccotto, Patrick J.; Mendelson, Tamra C. (2016-03-22). "The ecological drivers of nuptial color evolution in darters (Percidae: Etheostomatinae)". Evolution. 70 (4): 745–756. doi:10.1111/evo.12901. ISSN   0014-3820.
  12. Paine, M. D. (September 1990). "Life history tactics of darters (Percidae: Etheostomatiini) and their relationship with body size, reproductive behaviour, latitude and rarity". Journal of Fish Biology. 37 (3): 473–488. doi:10.1111/j.1095-8649.1990.tb05877.x. ISSN   0022-1112.
  13. 1 2 Martin, Zachary P.; Page, Lawrence M. (March 2015). "Comparative Morphology and Evolution of Genital Papillae in a Genus of Darters (Percidae: Etheostoma)". Copeia. 103 (1): 99–124. doi:10.1643/ci-14-081. ISSN   0045-8511.
  14. 1 2 Martin, Michael D.; Mendelson, Tamra C. (February 2016). "Male behaviour predicts trait divergence and the evolution of reproductive isolation in darters (Percidae: Etheostoma)". Animal Behaviour. 112: 179–186. doi: 10.1016/j.anbehav.2015.11.027 . ISSN   0003-3472.
  15. Kelly, N. B.; Near, T. J.; Alonzo, S. H. (2012-02-22). "Diversification of egg‐deposition behaviours and the evolution of male parental care in darters (Teleostei: Percidae: Etheostomatinae)". Journal of Evolutionary Biology. 25 (5): 836–846. doi:10.1111/j.1420-9101.2012.02473.x. ISSN   1010-061X.
  16. Héjja-Brichard, Yseult; Renoult, Julien; Mendelson, Tamra (2022-12-07). "Mate preference in sympatric and allopatric darters (genus Etheostoma): comparative evidence for a sex effect in reinforcement". dx.doi.org. Retrieved 2024-04-27.
  17. Williams, T. H.; Gumm, J. M.; Mendelson, T. C. (2013-09-18). "Sexual selection acting on a speciation trait in darters (Percidae: Etheostoma)". Behavioral Ecology. 24 (6): 1407–1414. doi:10.1093/beheco/art080. ISSN   1045-2249.

Fluker, B. L., Kuhajda, B. R., & Harris, P. M. (2014). The influence of life-history strategy on genetic differentiation and lineage divergence in darters (Percidae: Etheostomatinae). Evolution, 68(11), 3199-3216.

Héjja-Brichard, Y., Renoult, J. P., & Mendelson, T. C. (2022). Preference for conspecific mates in sympatric and allopatric darters (genus Etheostoma): comparative evidence for geographical and sex effects.

Hollingsworth Jr, P. R., & Near, T. J. (2009). Temporal patterns of diversification and microendemism in Eastern Highland endemic barcheek darters (Percidae: Etheostomatinae). Evolution, 63(1), 228-243.

Hopper, G. W. (2015). Ecological and morphological variation of darters among assemblages in Oklahoma streams (Doctoral dissertation, Kansas State University).

Keck, B. P., & Near, T. J. (2009). Patterns of natural hybridization in darters (Percidae: Etheostomatinae). Copeia, 2009(4), 758-773.

Kelly, N. B., Near, T. J., & Alonzo, S. H. (2012). Diversification of egg‐deposition behaviours and the evolution of male parental care in darters (Teleostei: Percidae: Etheostomatinae). Journal of Evolutionary Biology, 25(5), 836-846.

Martin, M. D., & Mendelson, T. C. (2016). Male behaviour predicts trait divergence and the evolution of reproductive isolation in darters (Percidae: Etheostoma). Animal Behaviour, 112, 179-186.

Martin, Z. P., & Page, L. M. (2015). Comparative morphology and evolution of genital papillae in a genus of darters (Percidae: Etheostoma). Copeia, 103(1), 99-124.

Near, T. J. (2002). Phylogenetic relationships of Percina (Percidae: Etheostomatinae). Copeia, 2002(1), 1-14.

Paine, M. D. (1990). Life history tactics of darters (Percidae: Etheostomatiini) and their relationship with body size, reproductive behaviour, latitude and rarity. Journal of Fish Biology, 37(3), 473-488.m

Sherman, A. N. (2022). Investigating Differences in the Gene Expression Profiles of Rainbow Darter (Percidae: Etheostoma caeruleum) Populations Across Space and Time (Doctoral dissertation, Southeastern Louisiana University). Williams, T. H., Gumm, J. M., & Mendelson, T. C. (2013). Sexual selection acting on a speciation trait in darters (Percidae: Etheostoma). Behavioral Ecology, 24(6), 1407-1414.