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The Comanche Springs pupfish (''Cyprinodon elegans'') is a species of
pupfish Pupfish are a group of small killifish belonging to ten genera of the family Cyprinodontidae of ray-finned fish. Pupfish are especially noted for being found in extreme and isolated situations. They are primarily found in North America, South ...
in the family Cyprinodontidae. It is endemic to Texas, and is now found only in spring-fed pools near Balmorhea, a small town in West Texas.


Taxonomy and naming

Discovery and formal description of the Comanche Springs pupfish occurred in 1853. The Comanche Springs pupfish was named for the now dry
Comanche Springs The Comanche or Nʉmʉnʉʉ ( com, Nʉmʉnʉʉ, "the people") are a Native American tribe from the Southern Plains of the present-day United States. Comanche people today belong to the federally recognized Comanche Nation, headquartered in Law ...
, an
aquifer An aquifer is an underground layer of water-bearing, permeable rock, rock fractures, or unconsolidated materials ( gravel, sand, or silt). Groundwater from aquifers can be extracted using a water well. Aquifers vary greatly in their characteri ...
of six artesian springs in
Pecos County, Texas Pecos County is a county located in the U.S. state of Texas. As of the 2020 census, its population was 15,193. The county seat is Fort Stockton. The county was created in 1871 and organized in 1875.. By Glenn Justice and John Leffler. Retri ...
.


Description

The Comanche Springs pupfish stands out amongst cogeners due to the striking speckled color pattern of the males, as well as a more streamlined body shape and lack of vertical bars. The back is relatively flat. The species reaches a maximum length of around . Differences also occur between members of the species from different locations in the number of fin rays and patterns of belly scales.


Distribution and habitat

Comanche Springs pupfish are currently found in a system of interconnected springs near Balmorhea, Texas: Phantom Lake Spring in
Jeff Davis County, Texas Jeff Davis County is a county in the U.S. state of Texas. As of the 2020 census, its population was 1,996. Its county seat is Fort Davis. The county is named for Jefferson Davis, who served as the 23rd United States Secretary of War in the 1 ...
, and
San Solomon Springs San Solomon Springs is a collection of artesian springs located near the small towns of Toyahvale and Balmorhea in Reeves County, Texas. Between 20 million and 28 million US gallons (90,850 cubic meters) of water a day flow from the springs, s ...
, and Giffin Spring, as well as
Toyah Creek Toyah Creek is a stream in western Texas that flows from Balmorhea to the Pecos River The Pecos River ( es, Río Pecos) originates in north-central New Mexico and flows into Texas, emptying into the Rio Grande. Its headwaters are on the eastern ...
, in
Reeves County, Texas Reeves County is a county located in the U.S. state of Texas. As of the 2020 census, its population was 14,748. Its county seat and most populous city is Pecos. The county was created in 1883 and organized the next year. It is named for Georg ...
. Base water flow for these springs comes from the Edwards-Trinity aquifer system. Additionally, runoff from the
Davis Mountains The Davis Mountains, originally known as Limpia Mountains, are a range of mountains in West Texas, located near Fort Davis, after which they are named. The fort was named for then United States Secretary of War and later Confederate President ...
recharges the overlying,
alluvial Alluvium (from Latin ''alluvius'', from ''alluere'' 'to wash against') is loose clay, silt, sand, or gravel that has been deposited by running water in a stream bed, on a floodplain, in an alluvial fan or beach, or in similar settings. Al ...
aquifers. Quantity and quality of habitat respond rapidly to these recharge events. Historically, these springs formed extensive
marsh A marsh is a wetland that is dominated by herbaceous rather than woody plant species.Keddy, P.A. 2010. Wetland Ecology: Principles and Conservation (2nd edition). Cambridge University Press, Cambridge, UK. 497 p Marshes can often be found a ...
es that likely supported large numbers of pupfish. Toyah Creek is intermittent, only flowing after heavy rainfall, and as a result only occasionally provides pupfish habitat. A population formerly existed in Comanche Springs in
Fort Stockton, Texas Fort Stockton is a city in and the county seat of Pecos County, Texas, United States. It is located on Interstate 10, future Interstate 14, U.S. Highways 67, 285, and 385, and the Santa Fe Railroad, northwest of San Antonio and southeas ...
, for which the species was named. These springs went dry in 1955, resulting in the
extirpation Local extinction, also known as extirpation, refers to a species (or other taxon) of plant or animal that ceases to exist in a chosen geographic area of study, though it still exists elsewhere. Local extinctions are contrasted with global extinct ...
of the local pupfish population. Small numbers of pupfish have also been found immediately downstream of East Sandia Spring during surveys in 1993 and 2001. West Sandia Spring and Saragosa Spring, though now mostly dry, likely supported the species before human modification. Most of the surviving habitat has been converted to a network of interconnected concrete or earthen
irrigation canal An acequia () or séquia () is a community-operated watercourse used in Spain and former Spanish colonies in the Americas for irrigation. Particularly in Spain, the Andes, northern Mexico, and the modern-day American Southwest particul ...
s. Water flow in these canals is swift and often
ephemeral Ephemerality (from the Greek word , meaning 'lasting only one day') is the concept of things being transitory, existing only briefly. Academically, the term ephemeral constitutionally describes a diverse assortment of things and experiences, fr ...
, and much of the network is unsuitable for use by pupfish. Because the water in the canals is sometimes diverted, resulting in mortalities, the canals are not considered permanent habitat, but rather serve as connections between larger populations in the springs. Low water levels in the aquifer have greatly reduced flow from Phantom Lake Spring, which led to the drying of the canals and reduction of pupfish habitat to a single pool at the cave from which the spring flows. As a result, the
U.S. Fish and Wildlife Service The United States Fish and Wildlife Service (USFWS or FWS) is an agency within the United States Department of the Interior dedicated to the management of fish, wildlife, and natural habitats. The mission of the agency is "working with othe ...
filled in the old canals and constructed a more natural
ciénega A ciénega (also spelled ciénaga) is a wetland system unique to the American Southwest. Ciénagas are alkaline, freshwater, spongy, wet meadows with shallow-gradient, permanently saturated soils in otherwise arid landscapes that often occupy n ...
at the cave mouth. A pumping system was put in place to maintain water levels, and an alarm alerts authorities to failures in the pump system. Artificial refugia for the species have also been constructed at
Balmorhea State Park Balmorhea State Park is a state park located on the San Solomon Springs in Reeves County, Texas, opened in 1968. The closest city is Balmorhea, Texas. Texas Parks and Wildlife Department manages the park. The park is open year-round, and visit ...
, which contains San Solomon spring. The entire head of this spring was converted into a concrete-lined swimming pool by the
Civilian Conservation Corps The Civilian Conservation Corps (CCC) was a voluntary government work relief program that ran from 1933 to 1942 in the United States for unemployed, unmarried men ages 18–25 and eventually expanded to ages 17–28. The CCC was a major part o ...
in the 1930s, destroying the natural ciénega. The first refugium, a long concrete-lined canal curving around the park motel, was completed in 1975. 1996 saw the addition of a ciénega within the grounds of the park, designed to replicate the appearance and functionality of the original. This wetland now contains the largest known concentrations of Comanche Springs pupfish. Finally, from 2009 to 2010, a second small ciénega was built adjacent to the 1975 canal in order to replace the older, deteriorating structure. This fish prefers water temperatures between 20-30 °C, and cannot survive above 40 °C. It occurs in water with lower salinity than that occupied by other pupfish species within the
Pecos River The Pecos River ( es, Río Pecos) originates in north-central New Mexico and flows into Texas, emptying into the Rio Grande. Its headwaters are on the eastern slope of the Sangre de Cristo mountain range in Mora County north of Pecos, New Mexico ...
system, suggesting a long interval of isolation.


Diet and feeding

The diet of adult Comanche Springs pupfish consists mostly of
filamentous algae Algae ( , ; : alga ) are any of a large and diverse group of photosynthetic, eukaryotic organisms. The name is an informal term for a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from un ...
as well as some snails, while juveniles subsist solely on
infusoria Infusoria are minute freshwater life forms including ciliates, euglenoids, protozoa, unicellular algae and small invertebrates. Some authors (e.g., Bütschli) used the term as a synonym for Ciliophora. In modern formal classifications, the te ...
until they grow larger.


Reproduction

Comanche Springs pupfish spawn year-round in spring outflows and small pools. Females reach sexual maturity at 5 months of age. Males use visual cues to identify and establish a territory. They select sites over algal mats in swift-moving water. Males take a position upstream of their territory thus allowing them to use the current to make high-speed attacks on intruders. Females will enter the territory from downstream and select a breeding site on the algal mat. The female lays her eggs singly, up to 30 a day, onto the algal mat or rock. The male aggressively guards the eggs until hatching. Less dominant, average-sized males occupy the periphery of the larger male territories, while the smallest males, known as sneaker males, mimic the coloration of females in order to access eggs in the territories of dominant males without provoking an attack. Breeding behavior is similar in slow-moving water except that the territories are centered around a convenient landmark such as a rock or patch of plants. Eggs average around in diameter, and hatch within 5 days at 20 °C. Newly hatched fry retain a relatively large yolk sack. Growth is rapid, and sexual maturity is reached at 5 months. Most Comanche Springs pupfish live for roughly one year.


Conservation status

The Comanche Springs pupfish is included on the
IUCN Red List The International Union for Conservation of Nature (IUCN) Red List of Threatened Species, also known as the IUCN Red List or Red Data Book, founded in 1964, is the world's most comprehensive inventory of the global conservation status of biol ...
of endangered species and was federally listed as endangered by the U.S. Government in 1967. Threats to this species include loss of habitat due to human alteration, reduced spring flows due to high levels of water mining, and competition from introduced species. Modification of the Balmorhea area spring system starting in 1875 destroyed most of the natural marsh and stream habitat. Construction of irrigation canals diverted water into agricultural fields, leading to the drying out of marshy areas and increasing the speed at which water left the spring area. In addition to Comanche Springs, the type locale for this species, a number of other springs in the area have gone dry as a result of the pumping of irrigation water from the aquifer, demonstrating the vulnerability of the remaining springs to desiccation. It is possible that continued pumping from deep-lying aquifers could eventually cause the entire Balmorhea spring system to dry up. Historic stocking of
black bass Black is a color which results from the absence or complete absorption of visible light. It is an achromatic color, without hue, like white and grey. It is often used symbolically or figuratively to represent darkness. Black and white have ...
species into the irrigation canals of the Phantom Cave Spring system likely severely impacted that population of pupfish. The introduction of another ''Cyprinodon'' species, the
sheepshead minnow The sheepshead minnow or sheepshead pupfish (''Cyprinodon variegatus'') is a species of ray-finned fish in the family Cyprinodontidae, the pupfishes. It is found in salt marsh and estuary environments and is native to the eastern coasts of North ...
, into Lake Balmorhea has resulted in a
hybrid Hybrid may refer to: Science * Hybrid (biology), an offspring resulting from cross-breeding ** Hybrid grape, grape varieties produced by cross-breeding two ''Vitis'' species ** Hybridity, the property of a hybrid plant which is a union of two diff ...
zone where the two species interbreed in the lower reaches of the Phantom Cave irrigation canals. One study found that when these hybrid offspring mated with Comanche Springs pupfish the resulting backcrossed offspring had lower fitness than those resulting from matings with sheepshead minnows, which could potentially result in
extinction Extinction is the termination of a kind of organism or of a group of kinds ( taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed ...
of the Comanche Springs pupfish through hybridization. As of 2013, physical barriers have prevented the sheepshead minnow from spreading further upstream in the spring system. Captive breeding stocks of the Comanche Springs pupfish are maintained at the Southwestern Native Aquatic Resources and Recovery Center (SNARRC) in
Dexter, New Mexico Dexter is a town in Chaves County, New Mexico. The population was 1,266 at the time of the 2010 census. Geography Dexter is located in south-central Chaves County at (33.196062, -104.370222), approximately west of the Pecos River. New Mexico ...
, and the Uvalde National Fish Hatchery in
Uvalde County, Texas Uvalde County ( ) is a county located in the U.S. state of Texas. As of the 2020 census, its population was 24,564. Its county seat is Uvalde. The county was created in 1850 and organized in 1856. It is named for Juan de Ugalde, the Spanish gov ...
. The Uvalde population is derived from individuals taken from the morphologically distinct Phantom Lake Spring subpopulation. In 2013, the total stock at the two facilities was 17,500 individuals. These stocks provide fish that can be used to replenish natural populations in the event of a catastrophic loss. Additionally, researchers can draw from the captive stocks instead of collecting wild fish. The Comanche Springs pupfish is further at risk simply due to the susceptibility of the small population to random catastrophic loss, such as a natural disaster. The risk of extinction for any one species is known to be strongly indirectly
correlated In statistics, correlation or dependence is any statistical relationship, whether causal or not, between two random variables or bivariate data. Although in the broadest sense, "correlation" may indicate any type of association, in statisti ...
to its population size.


References

{{Authority control Cyprinodon Endemic fauna of Texas Fish of the Western United States Freshwater fish of the United States Fauna of the Chihuahuan Desert Endangered fish Endangered fauna of the United States Taxa named by Spencer Fullerton Baird Taxa named by Charles Frédéric Girard Fish described in 1853 Taxonomy articles created by Polbot ESA endangered species