Percina macrocephala - (Cope, 1867)
Longhead Darter
Taxonomic Status: Accepted
Related ITIS Name(s): Percina macrocephala (Cope, 1867) (TSN 168486)
Unique Identifier: ELEMENT_GLOBAL.2.814563
Element Code: AFCQC04120
Informal Taxonomy: Animals, Vertebrates - Fishes - Bony Fishes - Perches and Darters
 
Kingdom Phylum Class Order Family Genus
Animalia Craniata Actinopterygii Perciformes Percidae Percina
Genus Size: D - Medium to large genus (21+ species)
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Concept Reference
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Concept Reference: Page, L. M., and T. J. Near. 2007. A new darter from the upper Tennessee River drainage related to Percina macrocephala (Percidae: Etheostomatinae). Copeia 2007:605-613.
Concept Reference Code: A07PAG01NAUS
Name Used in Concept Reference: Percina macrocephala
Taxonomic Comments: Page (1978) found that there are at least three morphologically distinct populations of Percina macrocepahala: one in the upper Tennessee River system, one in the Green River system, and a third in the upper Ohio River system. Further study by Page and Near (2007) determined that the upper Tennessee River population is a distinct species (P. williamsi). The Green River population, although somewhat distinctive from populations in the upper Ohio River drainage, does not appear to be diagnosable morphologically and shares identical mtDNA haplotypes with the upper Ohio River populations, so Page and Near (2007) maintained these populations as P. macrocephala.
Conservation Status
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NatureServe Status

Global Status: G3
Global Status Last Reviewed: 01Aug2012
Global Status Last Changed: 24Sep1996
Ranking Methodology Used: Ranked by inspection
Rounded Global Status: G3 - Vulnerable
Reasons: Spotty distribution in the Ohio River basin; rare and highly localized; extirpated in several areas; narrow habitat preference; threatened by habitat destruction/degradation (pollution, siltation, impoundments).
Nation: United States
National Status: N3 (07Feb2008)

U.S. & Canada State/Province Status
Due to latency between updates made in state, provincial or other NatureServe Network databases and when they appear on NatureServe Explorer, for state or provincial information you may wish to contact the data steward in your jurisdiction to obtain the most current data. Please refer to our Distribution Data Sources to find contact information for your jurisdiction.
United States Kentucky (S1), New York (S2), Ohio (SX), Pennsylvania (S4), Tennessee (S2), West Virginia (S2)

Other Statuses

IUCN Red List Category: DD - Data deficient
American Fisheries Society Status: Vulnerable (01Aug2008)

NatureServe Global Conservation Status Factors

Range Extent: 20,000-200,000 square km (about 8000-80,000 square miles)
Range Extent Comments: Historical range included the Ohio River Basin from southwestern New York (Smith 1985), western Pennsylvania (Cooper 1983), and eastern Ohio (extirpated; Trautman 1981) southward through Kentucky (Burr and Warren 1986), West Virginia (Stauffer et al. 1995), and Tennessee (Etnier and Starnes 1983, Page and Near 2007). Populations in North Carolina (extirpated; Menhinick 1991), western Virginia (Jenkins and Burkhead 1994), and in the upper Tennessee River drainage in eastern Tennessee, formerly included in P. macrocephala, are now regarded as a distinct species (P. williamsi) (Page and Near 2007). Many populations are believed to be extirpated, and the result is a relatively widespread but spotty distribution.

Area of Occupancy:  
Area of Occupancy Comments: Amount of occupied habitat is unknown but probably includes at least a few hundred stream kilometers.

Number of Occurrences: 21 - 80
Number of Occurrences Comments: This species is represented by a couple dozen extant occurrences (subpopulations).

Population Size: Unknown
Population Size Comments: Total adult population size is unknown. On a range-wide basis, this species has been characterized as rare and highly localized (Page and Burr 1991, 2011) and widespread and rare (Etnier and Starnes 1993). In New York, the longhead darter is relatively widespread and fairly common where found (Paul McKewon, New York State Department of Environment and Conservation, pers. comm.). Cooper (1983) reported that this species is common in the Allegheny River, Pennsylvania, whereas other sources characterize the species as sporadically encountered in small numbers in Pennsylvania (Felbaum et al. 1995). In West Virginia, the species is still relatively common in the Elk River system (Dan Cincotta, West Virginia Department of Natural Resources, pers. comm.). The longhead darter is described as sporadic and rare in Kentucky (Burr and Warren 1986), generally rare and locally extirpated in Tennessee (Etnier and Starnes 1993).

Number of Occurrences with Good Viability/Integrity: Unknown
Viability/Integrity Comments: Many historical occurrences are based on old collection data and do not represent currently viable populations.

Overall Threat Impact: High
Overall Threat Impact Comments: Proximate principal threats are most likely increased turbidity and siltation resulting from agricultural, industrial, and municipal development (Page and Near 2007). Populations in the south and in New York (Paul McKeown, New York Department of Environment & Conservation) are probably most affected by agricultural land uses that deposit silt in pools and may smother eggs and larvae. In West Virginia, stream sedimentation resulting from recent coal mining operations may be the biggest threat (Dan Cincotta, West Virginia Department of Natural Resources, pers. comm.). This is primarily because of the need for low sulfur coal, which is available in this area, and new mining technologies ("mountain-topping"). Many populations in the southern portion of the range are isolated by impoundments or other habitat barriers.

Short-term Trend: Decline of <30% to relatively stable
Short-term Trend Comments: In most states where longhead darters are extant, official state status is Threatened. Even before Percina williamsi was split off as a distinct species, Etnier and Starnes (1993) stated that P. macrocephala "probably warrants Threatened status throughout its range." American Fisheries Society (Williams et al. 1989) listed longhead darter as Threatened.

In a review of the conservation status of fish species in the southern United States, Warren et al. (2000) categorized the longhead darter as Threatened (their second-highest category of imperilment; likely to become endangered throughout all or a significant portion of the range). These assessments were made before P. williamsi was split from P. macrocephala (reducing the scope of the latter), so the status of P. macrocephala may be worse than these reports indicate. However, the assessment apparently did not encompass northern populations of P. macrocephala such as those in New York and Pennsylvania. A range-wide categorization by Jelks et al. (2008) assigned this species to the Vulnerable category.

Recent information indicates that some northern populations may be more abundant than formerly believed (Jay Stauffer, Pennsylvania State University, pers. comm.; Dan Cincotta, West Virginia Department of Natural Resources pers. comm; Paul McKeown, New York Department of Environmental Conservation, pers. comm.). Alternatively, traditional collecting methods may have previously underestimated true abundance and distribution. Recent efforts directed specifically at longhead darters have been more successful, but perhaps abundance is cyclic

Long-term Trend: Decline of 30-70%
Long-term Trend Comments: Over the long term, extent of occurrence, area of occupancy, number of subpopulations, and population size clearly have declined, but the degree of decline is uncertain. This darter has been extirpated from much of its range (Page 1978, Trautman 1981, Burr and Warren 1986).

In West Virginia, the species is still relatively common in the Elk River system, but recent land use changes resulting from coal mining have begun to affect aquatic habitats there (Dan Cincotta, West Virginia Department of Natural Resources, pers. comm.).

In Kentucky, the species is probably extirpated from the Cumberland and Kentucky rivers, each of which has one substantiated record (Burr and Warren 1986), but current status is relatively stable in Kentucky (R. Cicerello, pers. comm., 1997).

In Tennessee, the species is rare to extirpated in various streams (Etnier and Starnes 1993), and it has not been found in the Cumberland River since 1891 (Page 1978).

Other NatureServe Conservation Status Information

Protection Needs: Actions are needed to control sediment runoff from mining (primarily West Virgina), row crop agriculture, and degradation of riparian zones and aquatic habitat by livestock (Kentucky, Tennessee, New York, and Pennsylvania). Restoring riparian vegetation, fencing livestock from streams, and providing alternate water sources are recommended. Channelization should be avoided, as should removal of woody debris (snagging and dragging) from stream margins. Where these practices have taken place, natural processes should be allowed to proceed so that the streambed and stream banks become stabilized. Removal of barriers to fish movement may be needed in some watersheds.

Distribution
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Global Range: (20,000-200,000 square km (about 8000-80,000 square miles)) Historical range included the Ohio River Basin from southwestern New York (Smith 1985), western Pennsylvania (Cooper 1983), and eastern Ohio (extirpated; Trautman 1981) southward through Kentucky (Burr and Warren 1986), West Virginia (Stauffer et al. 1995), and Tennessee (Etnier and Starnes 1983, Page and Near 2007). Populations in North Carolina (extirpated; Menhinick 1991), western Virginia (Jenkins and Burkhead 1994), and in the upper Tennessee River drainage in eastern Tennessee, formerly included in P. macrocephala, are now regarded as a distinct species (P. williamsi) (Page and Near 2007). Many populations are believed to be extirpated, and the result is a relatively widespread but spotty distribution.

U.S. States and Canadian Provinces

Due to latency between updates made in state, provincial or other NatureServe Network databases and when they appear on NatureServe Explorer, for state or provincial information you may wish to contact the data steward in your jurisdiction to obtain the most current data. Please refer to our Distribution Data Sources to find contact information for your jurisdiction.
Color legend for Distribution Map
Endemism: endemic to a single nation

U.S. & Canada State/Province Distribution
United States KY, NY, OHextirpated, PA, TN, WV

Range Map
No map available.


U.S. Distribution by County Help
State County Name (FIPS Code)
KY Adair (21001), Allen (21003), Barren (21009)*, Butler (21031)*, Casey (21045)*, Clay (21051)*, Green (21087), Hart (21099), Lewis (21135), Monroe (21171)*, Pike (21195)*, Taylor (21217), Warren (21227), Wayne (21231)*
NC Buncombe (37021), Madison (37115)
NY Cattaraugus (36009), Chautauqua (36013)*
PA Allegheny (42003)*, Armstrong (42005), Clarion (42031), Crawford (42039), Erie (42049), Fayette (42051)*, Forest (42053), McKean (42083), Mercer (42085), Potter (42105), Somerset (42111)*, Venango (42121), Warren (42123), Washington (42125), Westmoreland (42129)*
TN Bedford (47003)*, Coffee (47031)*, Lewis (47101), Macon (47111)*, Pickett (47137)*
WV Braxton (54007), Clay (54015), Kanawha (54039), Webster (54101)
* Extirpated/possibly extirpated
U.S. Distribution by Watershed Help
Watershed Region Help Watershed Name (Watershed Code)
05 Upper Allegheny (05010001)+, Middle Allegheny-Tionesta (05010003)+, French (05010004)+, Middle Allegheny-Redbank (05010006)+, Lower Monongahela (05020005)+, Youghiogheny (05020006)+*, Walhonding (05040003)*, Elk (05050007)+, Lower Levisa (05070203)+, Ohio Brush-Whiteoak (05090201)+, South Fork Kentucky (05100203)+*, Upper Green (05110001)+, Barren (05110002)+, Middle Green (05110003)+*, South Fork Cumberland (05130104)+*, Obey (05130105)+
06 Upper French Broad (06010105)+, Powell (06010206), Upper Duck (06040002)+, Buffalo (06040004)+
+ Natural heritage record(s) exist for this watershed
* Extirpated/possibly extirpated
U.S. Distribution by Watershed (based on multiple information sources) Help
Ecology & Life History
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Basic Description: A small fish (darter).
General Description: This fish is a relatively large darter (reaching nearly five inches total length), with a slender body and long, pointed head and snout and a broad and irregular dark lateral stripe with depigmented area where the lateral line passes through its center. This stripe begins in front of a dark caudal spot, and continues across opercle and beyond eye to tip of snout. The area above the lateral stripe is also dark. Below the lateral stripe, the belly is mostly immaculate, with some scattered dark pigment. Dark bands are present along the edge and middle of the spinous dorsal fin, and all other fins are speckled with dark pigment. No bright colors are present. Dorsal spines 11-16; dorsal soft rays 11-14; and lateral-line scales 69-90.
Diagnostic Characteristics: Longhead darters are similar to the blackside, dusky, slenderhead and frecklebelly darters (P. maculata, P. sciera, P. phoxocephala, and P. stictogaster), all of which are potentially sympatric in various parts of the longhead darter's range. Longhead darters are separated from all in having naked, as opposed to well scaled opercles. They are separated from blackside and dusky darters in having a longer snout, and a different lateral pattern including blotches that are fused into the stripe described above. Frecklebelly darters, which may look, superficially like longhead darters, also have a fully scaled head (naked in longhead darters) and lack the modified midventral scales present in longhead darters.
Reproduction Comments: The scant life history information on Percina macrocephala and its close relatives (including P. williamsi), based on Kentucky and Tennessee populations, was summarized as follows by Page (1978), Etnier and Starnes (1993), and Jenkins and Burkhead (1994). Spawning takes place in spring (March to May). There is no parental care given to eggs or larvae, but details of spawning are not published. Eggs hatch after 27 days at water temperature of 10 C; hatchlings were 10 mm. Individuals grow to 50-60 mm by age one, 70-80 mm by age two, and 90-100 mm by age three or four. Sexual reproduction is apparently not reached until individuals are two years of age. Lifespan is three to four years.
Habitat Type: Freshwater
Non-Migrant: Y
Locally Migrant: N
Long Distance Migrant: N
Riverine Habitat(s): CREEK, MEDIUM RIVER, Moderate gradient, Pool, Riffle
Special Habitat Factors: Benthic
Habitat Comments: Habitat includes boulder- and cobble-strewn flowing pools and the areas above and below deep, fast riffles underlain with cobble, in larger upland creeks and small to medium rivers (Burr and Warren 1986). Spawning presumably occurs in gravel shoals.
Adult Food Habits: Invertivore
Immature Food Habits: Invertivore
Food Comments: In Kentucky, eats mainly crayfish and insect larvae (e.g., mayflies) (Page 1978, Page 1983).
Length: 9 centimeters
Economic Attributes Not yet assessed
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Management Summary
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Stewardship Overview: Much basic information required as a basis for population management is unknown. Basic range-wide information on the status of the species is lacking, as is the best method to survey for this difficult-to-collect fish. Traditional fish sampling methods may seriously underestimate distribution and abundance of longhead darters. Some details of life history are lacking, as is adequate information on population and metapopulation dynamics, including movement patterns. This information is necessary for proper management. Regular monitoring of all populations is lacking, but needed. This is especially important in the southern portion of the range. Some southern populations may require population augmentation; other extirpated populations may be restored by reintroduction. Captive propagation techniques should be developed for this potential need.
Restoration Potential: Some extant populations are extremely vulnerable because of small size and extent. If events occur that result in further fragmentation (e.g., establishment of habitat barriers, even if temporary), recolonization may be unlikely because of small source populations. Recovery may be aided by reintroduction or augmentation, and should be considered a reasonable management activity in areas where habitat conditions have improved in areas where the species is currently believed to have been extirpated. Captive propagation might be necessary to obtain enough individuals for these efforts.
Preserve Selection & Design Considerations: Mostly unknown at present.
Management Requirements: Some obvious factors that can be managed are habitat barriers present in a watershed containing the fish. Channelization should be avoided, as should removal of woody debris (snagging and dragging) from stream margins. Where these practices have taken place, natural processes should be allowed to proceed so that the streambed and stream banks become stabilized.
Monitoring Requirements: Apparent variations in abundance in a particular locality may be the result of cyclic population fluctuations, or they could reflect differences in the effectiveness of various survey methods. Regular monitoring using standardized methods is needed to clarify this situation.

Sampling with traditional collecting methods (seines, backpack electroshockers, gill nets, boat shocking) may not accurately document their presence or abundance. These fish may be able to avoid collection by these methods. However, recent surveys indicate that electric seines may work well in smaller streams (Paul McKewon, New York State Department of Environmental Conservation, pers. comm.), and direct observation with snorkel or SCUBA may be the best method in larger streams (Jay Stauffer, Pennsylvania State University, pers. comm.; and P. W. Shute, pers. obs.). Because longhead darters apparently move to deeper areas in winter, surveys during the colder months may be difficult without appropriate cold-water gear, and are probably not necessary to document trends.

Biological Research Needs: The most immediate research need is to determine the actual current range-wide distribution and abundance, movement/dispersal patterns, and metapopulation dynamics. This information will be necessary before we can determine the watershed area appropriate for viable populations. Also, a more complete understanding of life history (more details on spawning sites, and larval juvenile habitat requirements for example) will help ensure management activities are appropriate to protect habitats and other factors necessary to complete all life history stages. In the event reintroduction or population augmentation is believed to be necessary or beneficial, techniques should be developed to propagate longhead darters in captivity (appropriate natural populations may not be large enough to serve as sources for reintroductions).
Population/Occurrence Delineation
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Group Name: Darters

Use Class: Not applicable
Minimum Criteria for an Occurrence: Occurrences are based on evidence of historical presence, or current and likely recurring presence, at a given location. Such evidence minimally includes collection or reliable observation and documentation of one or more individuals (including eggs and larvae) in appropriate habitat.
Separation Barriers: Dam lacking a suitable fishway; high waterfall; upland habitat.
Separation Distance for Unsuitable Habitat: 10 km
Separation Distance for Suitable Habitat: 10 km
Separation Justification: Data on dispersal and other movements generally are not available. Though larvae of some species may drift with the current, Turner (2001) found no significant relationship between a larval transport index and gene flow among several different darter species.

Separation distances are arbitrary but reflect the likely low probability that two occupied locations separated by less than several kilometers of aquatic habitat would represent truly independent populations.

Because of the difficulty in defining suitable versus unsuitable habitat, especially with respect to dispersal, and to simplify the delineation of occurrences, a single separation distance is used regardless of habitat quality.

Occupied locations that are separated by a gap of 10 km or more of any aquatic habitat that is not known to be occupied generally represent different occurrences. However, it is important to evaluate seasonal changes in habitat to ensure that an occupied habitat occurrence for a particular population does not artificially separate spawning areas and nonspawning areas as different occurrences simply because there have been no collections/observations in an intervening area that may exceed the separation distance.

Date: 21Sep2004
Author: Hammerson, G.
Population/Occurrence Viability
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U.S. Invasive Species Impact Rank (I-Rank) Not yet assessed
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Authors/Contributors
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NatureServe Conservation Status Factors Edition Date: 01Aug2012
NatureServe Conservation Status Factors Author: Hammerson, G., P. Shute, R. Jennings, and S. Roble
Management Information Edition Date: 03Nov1999
Management Information Edition Author: SHUTE, PEGGY W.
Management Information Acknowledgments: Thanks to Dan Cincotta, West Virginia Department of Natural Resources; Jay Stauffer, Pennsylvania State University; Paul McKewon, New York State Department of Environment and Conservation.
Element Ecology & Life History Edition Date: 08Feb2008
Element Ecology & Life History Author(s): Shute, P. W., and G. Hammerson

Zoological data developed by NatureServe and its network of natural heritage programs (see Local Programs) and other contributors and cooperators (see Sources).

References
Help
  • Burr, B. M., and M. L. Warren, Jr. 1986a. Distributional atlas of Kentucky fishes. Kentucky Nature Preserves Commission, Scientific and Technical Series No. 4, Frankfort, Kentucky. 398 pp.

  • Carlson, Douglas M. 1998. Species Accounts for the rare fishes of New York. New York State Department of Environmental Conservation, Division of Fish, Wildlife and Marine Resources. Bureau of Fisheries, Endangered Fish Project. 95pp.

  • Carlson, Douglas. 1998. Summary of activities relating to management of ETs Fishes (as listed in 1983) from 1995 to present. 5pp.

  • Cooper, E. L. 1983. Fishes of Pennsylvania and the northeastern United States. Pennsylvania State University Press, University Park. 243 pp.

  • Cooper, E.L. 1983. Fishes of Pennsylvania. Penn State Univ. Press, University Park, PA.

  • Eaton, S.W., R.J. Nemecek and M.M. Kozubowski. 1982. Fishes of the Allegheny River above Kinzua Dam. New York Fish and Game J. 29(2):189-198.

  • Etnier, D. A. 1997. Longhead darter, Percina macrocephala. Page 78 in E.F. Menhinick and A.L. Braswell. Endangered, Threatened, and rare fauna of North Carolina. Part IV. A reevaluation of the freshwater fishes. Occasional papers of the North Carolina State Museum of Natural Sciences and the North Carolina Biological Survey 11.

  • Etnier, D. A., and W. C. Starnes. 1993. The fishes of Tennessee. University of Tennessee Press, Knoxville, Tennessee. xiv + 681 pp.

  • Etnier, David A. and Wayne C. Starnes. 1993. The Fishes of Tennessee. University of Tennessee Press, Knoxville. 681 pp.

  • Felbaum, F., B. Mitchell, K. McKenna, J. Hassinger, A. Shiels, J. Hart and D. Brauning. 1995. Endangered and threatened species of Pennsylvania. Wildlife Conservation Resource Fund, Harrisburg, PA, 80 pp

  • Greenberg, L. A. 1991. Habitat use and feeding behavior of thirteen species of benthic stream fishes. Environmental Biology of Fishes 31:389-401.

  • Jelks, H. L., S. J. Walsh, N. M. Burkhead, S. Contreras-Balderas, E. Díaz-Pardo, D. A. Hendrickson, J. Lyons, N. E. Mandrak, F. McCormick, J. S. Nelson, S. P. Platania, B. A. Porter, C. B. Renaud, J. Jacobo Schmitter-Soto, E. B. Taylor, and M.L. Warren, Jr. 2008. Conservation status of imperiled North American freshwater and diadromous fishes. Fisheries 33(8):372-407.

  • Jenkins, R. E., and N. M. Burkhead. 1994. Freshwater fishes of Virginia. American Fisheries Society, Bethesda, Maryland. xxiii + 1079 pp.

  • Kuehne, R. A., and R. W. Barbour. 1983. The American Darters. University Press of Kentucky, Lexington, Kentucky. 177 pp.

  • Lee, D. S., C. R. Gilbert, C. H. Hocutt, R. E. Jenkins, D. E. McAllister, and J. R. Stauffer, Jr. 1980. Atlas of North American freshwater fishes. North Carolina State Museum of Natural History, Raleigh, North Carolina. i-x + 854 pp.

  • Nelson, J. S., E. J. Crossman, H. Espinosa-Perez, L. T. Findley, C. R. Gilbert, R. N. Lea, and J. D. Williams. 2004. Common and scientific names of fishes from the United States, Canada, and Mexico. American Fisheries Society, Special Publication 29, Bethesda, Maryland. 386 pp.

  • Page, L. M. 1978. Redescription, distribution, variation and life history notes on Percina macrocephala (Percidae). Copeia 1978:655-664.

  • Page, L. M. 1983a. Handbook of Darters. T. F. H. Publications, Inc., Neptune City, New Jersey. 271 pp.

  • Page, L. M., H. Espinosa-Pérez, L. T. Findley, C. R. Gilbert, R. N. Lea, N. E. Mandrak, R. L. Mayden, and J. S. Nelson. 2013. Common and scientific names of fishes from the United States, Canada, and Mexico. Seventh edition. American Fisheries Society, Special Publication 34, Bethesda, Maryland.

  • Page, L. M., and B. M. Burr. 1991. A field guide to freshwater fishes: North America north of Mexico. Houghton Mifflin Company, Boston, Massachusetts. 432 pp.

  • Page, L. M., and B. M. Burr. 2011. Peterson field guide to freshwater fishes of North America north of Mexico. Second edition. Houghton Mifflin Harcourt, Boston. xix + 663 pp.

  • Page, L. M., and T. J. Near. 2007. A new darter from the upper Tennessee River drainage related to Percina macrocephala (Percidae: Etheostomatinae). Copeia 2007:605-613.

  • Robins, C.R., R.M. Bailey, C.E. Bond, J.R. Brooker, E.A. Lachner, R.N. Lea, and W.B. Scott. 1991. Common and scientific names of fishes from the United States and Canada. American Fisheries Society, Special Publication 20. 183 pp.

  • Smith, C. L. 1985. The inland fishes of New York State. New York State Department of Environmental Conservation. Albany, New York, xi + 522 pp.

  • Smith, C.L. 1985. The Inland Fishes of New York State. New York State Department of Environmental Conservation. Albany, NY. 522pp.

  • Starnes, W. C. 1995. Taxonomic validation for fish species on the U.S. Fish and Wildlife Service Category 2 species list. 28 pp.

  • Warren, M. L., Jr., B. M. Burr, S. J. Walsh, H. L. Bart, Jr., R. C. Cashner, D. A. Etnier, B. J. Freeman, B. R. Kuhajda, R. L. Mayden, H. W. Robison, S. T. Ross, and W. C. Starnes. 2000. Diversity, distribution, and conservation status of the native freshwater fishes of the southern United States. Fisheries 25(10):7-31.

  • Werner, R.G. 1980. Freshwater fishes of New York State. N.Y.: Syracuse University Press. 186 pp.

  • Williams, J.E, J.E. Johnson, D.A. Hendrickson, S. Contreras-Balderas, J.D. Williams, M. Navarro-Mendoza, D.E. McAllister, and J.E. Deacon. 1989b. Fishes of North America endangered, threatened or of special concern: 1989. Fisheries 14(6):2-20.

References for Watershed Distribution Map
  • State Natural Heritage Data Centers. 1996a. Aggregated element occurrence data from all U.S. state natural heritage programs, including the Tennessee Valley Authority, Navajo Nation and the District of Columbia. Science Division, The Nature Conservancy.

  • Stauffer, J. R., Jr., J. M. Boltz, and L. R. White. 1995. The fishes of West Virginia. Proceedings of the Academy of Natural Sciences of Philadelphia 146:1-389.

  • Trautman, M. B. 1981. The fishes of Ohio. Second edition. Ohio State University Press, Columbus, Ohio. 782 pp.

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