Cyprinodon macularius: Difference between revisions
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#REDIRECT [[Cyprinodon macularius]] |
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{{Taxobox |
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| status = en | status_system = IUCN2.3 |
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| image = Male_female_mecularius.jpg |
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| regnum = [[Animal]]ia |
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| phylum = [[Chordate|Chordata]] |
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| classis = [[Actinopterygii]] |
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| ordo = [[Cyprinodontiformes]] |
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| familia = [[Cyprinodontidae]] |
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| genus = ''[[Cyprinodon]]'' |
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| species = '''''C. macularius''''' |
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| binomial = ''Cyprinodon macularius'' |
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| binomial_authority = [[Spencer Fullerton Baird|S. F. Baird]] & [[Charles Frédéric Girard|Girard]], 1853 |
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}} |
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'''''Cyprinodon macularius''''' is a rare species of [[teleost]] in the family Cyprinodontidae and is known by the common name '''desert pupfish'''. It is a federally listed [[endangered species]] in the United States. |
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Presently, the only remaining natural populations of the desert pupfish are located at a few sites in the [[Salton Sea]] drainage in [[California]], and the [[Colorado River Delta]] in [[Baja California]] and [[Sonora, Mexico]].<ref>Minckley 1973, U.S. Fish and Wildlife Service [USDI, FWS] 1993).</ref> The historic range of the desert pupfish included the lower [[Gila River]] basin in [[Arizona]] and [[Sonora, Mexico]]. It inhabited the Gila, [[Santa Cruz River (Arizona)|Santa Cruz]], [[San Pedro River (Arizona)|San Pedro]], and [[Salt River (Arizona)|Salt River]]s as well as the lower [[Colorado River]] from around [[Needles, California]], to the [[Gulf of California]]. |
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There are natural populations of this fish remaining in California and Mexico, however, populations have been introduced to several areas and some fish are kept in private ponds and aquaria. |
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The desert pupfish was once a common fish, but it is now [[local extinction|extirpated]] from most of its historical natural range. The decrease in populations has been a trend since the early 1900s due to the loss of habitat. The desert pupfish is [[endangered species|endangered]] due to habitat destruction and fragmentation. It has been, and continues to be, preyed upon and displaced by [[introduced species|non-native fishes]]. |
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==Taxonomy and phylogeny== |
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''Cyprinodon macularius'' was first described by [[Spencer Baird]] and [[Charles Girard]] in 1853. <ref name=girard1>{{cite journal|last=Baird|first=Spencer F.|coauthors=Charles Girard|title=Descriptions of New Species of Fishes Collected by Mr. John H. Clark, on the U. S. and Mexican Boundary Survey, under Lt. Col. Jas. D. Graham|journal=Proceedings of the Academy of Natural Sciences of Philadelphia|year=1852-1853|volume=6|page=389}}</ref> Originally, it was considered to be made up of three subspecies: the nominal desert pupfish (''Cyprinodon macularius macularius''), the Quitobaquito pupfish (''Cyprinodon macularius eremus'') and the undescribed Monkey Spring pupfish (''[[Cyprinodon]]'' sp.). <ref name=usfws93>United States fish and Wildlife Service (1993). Desert Pupfish (Cyprinodon macularius) Recovery Plan. (Unpublished). Prepared for the USFWS, Region 2.</ref> Recently, the three subspecies have been classified as three distinct species: the desert pupfish (''C. macularius''), the Sonoyta (Quitobaquito) pupfish (''[[Cyprinodon eremus|C. eremus]]''), and the Santa Cruz (Money Spring) pupfish (''[[Cyprinodon arcuatus|C. arcuatus]]'').<ref name=usfws10>Unites States Fish and Wildlife Service (2010). Desert Pupfish (Cyprinodon macularius); 5-Year Review: Summary and Evaluation. Phoenix, Arizona: USFWS.</ref> |
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The study of [[mitochondrial DNA]] variation between different populations of pupfish traditionally considered as ''C. macularius'' provide the basis for the species distinction between ''C. macularius'' and ''C. eremus.''<ref name=usfws10 /> The mtDNA [[haplotypes]] of the pupfish endemic to the Rio Sonoyta Basin/Quitobaquito Springs and the pupfish endemic to the Salton Sea/Lower Colorado River Basin suggest sustained, mutually exclusive evolutionary histories.<ref name=neigel1>{{cite book|last=Neigel|first=J.E.|title=Evolutionary Processes and Theory|year=1986|publisher=Academic Press|pages=515–534|coauthors=J.C. Avise|editor=S. Karlin|chapter=Phylogenetic relationships of mitochondrial DNA under various demographic models of speciation}}</ref> Furthermore, the inconsistencies between haplotype frequency are highly significant and suggest species-divergence of about 100,000 years.<ref name=eschelle1>{{cite journal|last=Echelle|first=Anthony A.|coauthors=Van Den Bussche, Ronald A.; Malloy, Terrence P.; Haynie, Michelle L.; Minckley, C. O.; McEachran, J. D.|title=Mitochondrial DNA Variation in Pupfishes Assigned to the Species (Atherinomorpha: Cyprinodontidae): Taxonomic Implications and Conservation Genetics|journal=Copeia|volume=2000|issue=2|pages=353–364|doi=10.1643/0045-8511(2000)000[0353:MDVIPA]2.0.CO;2}}</ref><ref name=ives1>{{cite book|last=Ives|first=Ronald|title=The Pinacate region, Sonora, Mexico, (Occasional Papers of the California Academy of Sciences)|year=1964|publisher=The California Academy of Sciences|pages=1-43}}</ref> In 2002, the Monkey Spring pupfish, renamed the Santa Cruz pupfish, was described as a third distinct species of ''Cyprinodon'', ''C. arcuatus.''<ref name=minckley1>{{cite journal|last=Minckley|first=W. L.|coauthors=Miller, Robert Rush; Norris, Steven Mark; Schaefer, S. A.|title=Three New Pupfish Species, Cyprinodon (Teleostei, Cyprinodontidae), from Chihuahua, México, and Arizona, USA|journal=Copeia|date=1 August 2002|volume=2002|issue=3|pages=687–705|doi=10.1643/0045-8511(2002)002[0687:TNPSCT]2.0.CO;2}}</ref> |
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==Description== |
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[[File:Male macularius.jpg|thumb|left|Bright blue coloration of the male ''C. macularius'' can be seen.]] |
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The desert pupfish is a small fish that is typically less than three inches long; males are larger than females and generally have more vivid markings, specifically during breeding seasons. Females and juveniles typically have tan or olive backs and silvery sides with narrow, dark vertical bars situated laterally. These bars are often interrupted to give the impression of a disjunct, lateral band. During mating season, males become bright blue on the dorsal portion of the head and sides, and yellow or orange on the [[fish fin|caudal fin]] and posterior caudal peduncle.<ref name=fr1>51 FR 10843 (1986-03-31).</ref> |
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The profile of the desert pupfish is smoothly rounded, while its body is thickened and laterally compressed. The desert pupfish has a protruding mouth equipped with tricuspid teeth.<ref name=usfws10 /> |
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==Habitat and distribution== |
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[[File:Macularius distribution.png|thumb|Distribution of ''[[Cyprinodon macularius|C. macularius]]''.]] |
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The desert pupfish is found in shallow waters of desert springs, small streams, and marshes below 5,000 feet in elevation.<ref name=usfws93 /> The species can tolerate high salinities, water temperatures, and lower oxygen content than most fish, and will occupy habitats inhospitable to invasive fish. Pupfish typically prefer clear waters with either rooted or unattached aquatic plants, restricted surface flow, or sand-silt substrates, and will do well if their habitats have little vegetation apart from mats of benthic algae.<ref name=drecp1>Dudek and ICF International (2012). Desert Renewable Energy Conservation Plan (DRECP) Baseline Biology Report. (Draft). Prepared for the California Energy Commission.</ref> |
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The distribution of the desert pupfish historically ranged from the lower [[Colorado River]] in [[Arizona]] and [[California]] to the [[Gulf of Mexico]] and onto its delta in [[Sonora]] and [[Baja California]]. Specifically, the desert pupfish occurred the [[Salton Sink]] basin in California; the [[Gila River]] basin in Arizona and Sonora, including the [[Gila River|Gila]], [[Santa Cruz River (Arizona)|Santa Cruz]], [[San Pedro River (Arizona)|San Pedro]], and [[Salt River (Arizona)|Salt Rivers]]; the Rio Sonoyta in Arizona and Sonora; [[Puerto Peñasco]], Sonora; and the [[Laguna Salada (Mexico)|Laguna Salada]] basin in [[Baja California]].<ref name=drecp1 /> Due to both habitat destruction and species reclassification, however, the desert pupfish has a smaller distribution than previously considered. |
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As of 2010, naturally occurring populations of the desert pupfish are restricted in the United States to two tributaries and various shoreline pools and irrigation drains of the Salton Sea in California.<ref name=usfws10 /><ref name=lau1>Lau, S.; Boehm, C. (1991). A distribution survey of desert pupfish (Cyprinodon macularius) around the Salton Sea, California. Final Report for Section 6, Project No. EF9oXII-1. Prepared for the California Department of Fish and Game, Inland Fisheries Division.</ref> In Mexico, the desert pupfish is scattered along the [[Colorado River Delta]] and the Laguna Salada basin.<ref name=hendrickson1>{{cite journal|last=Hendrickson|first=D.|coauthors=A. Valera-Romera|title=Conservation Status of Desert Pupfish, Cyprinodon macularius, in México and Arizona|journal=Copoeia|year=1989|volume=1989|pages=478-483}}</ref> Collectively, there are 11 known populations of extant wild desert pupfish within the [[United States]] and [[Mexico]]. Attempted reintroductions have yielded 16 transplanted populations in Arizona, as well as 46 captive or refuge populations in Arizona, California, and Mexico.<ref name=usfws10 /> |
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==Diet== |
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Pupfish are omnivores that forage in shallow zones during early morning and late evening; they will move out of the shallows during the day when water temperatures may rise to 36°C or higher.<ref name=thompson1>{{cite web|last=Thompson|first=Lisa C.|title=California Fish Species|url=http://calfish.ucdavis.edu/species/?uid=28&ds=241}}</ref> Rate of surface foraging drops when water temperatures fall below 21ºC.<ref name=legner1>{{cite journal|last=Legner|first=E. F.|coauthors=Medved, R. A.; Hauser, W. J.|title=Predation by the desert pupfish, Cyprinodon macularius onCulex mosquitoes and benthic chironomid midges|journal=Entomophaga|date=1 March 1975|year=1975|month=March|volume=20|issue=1|pages=23–30|doi=10.1007/BF02373447}}</ref> The diet of the desert pupfish may include algae, small invertebrates, aquatic crustaceans, aquatic insect larvae, snails, detritus, and occasionally intraspecies eggs and young.<ref name=thompson1 /><ref name=legner1 /><ref name=keeney1>{{cite web|last=Keeney|first=Sharon|title=Desert pupfish (Cyprinodon macularis)|url=http://www.dfg.ca.gov/regions/6/Conservation/Desert_Fish/Desert_Pupfish.html}}</ref><ref name=naiman1>{{cite journal|last=Naiman|first=Robert J.|title=Preliminary Food Studies of Cyprinodon macularius and Cyprinodon nevadensis (Cyprinodontidae)|journal=The Southwestern Naturalist|date=15 September 1979|year=1979|month=September|volume=24|issue=3|pages=538-541}}</ref><ref name=cox1>{{cite dissertation|last=Cox|first=Thomas J.|title=A Behavioral and Ecological Study of the Desert Pupfish (Cyprinodon macularis) in Quitobaquito Springs, Organ Pipe Cactus National Monument, Arizona|date=1966|pages=91|publisher=The University of Arizona|location=Arizona}}</ref> Pupfish will feed somewhat heavily on [[nektonic]] organisms, going so far as to suppress or even eliminate local populations of mosquitoes. In the absence of mosquito prey, pupfish will consume [[benthic]] [[chironomid]] [[midges]] both from substrate and from midwater when midges attempt to come to the surface. <ref name=legner1 /> |
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Exposed eggs are typically and readily eaten by other pupfish if not concealed by the bearer, while mobile fry are ignored unless adults are very hungry.<ref name=legner1 /><ref name=loiselle1>{{cite journal|last=Loiselle|first=Paul V.|title=Filial cannibalism and egg recognition by males of the primitively custodial teleost Cyprinodon macularius californiensis girard (Atherinomorpha: Cyprinodontidae)|journal=Ethology and Sociobiology|date=1 January 1983|volume=4|issue=1|pages=1–9|doi=10.1016/0162-3095(83)90002-X}}</ref> Furthermore, though pupfish may engage in [[filial cannibalism]], males have been observed to distinguish between eggs fertilized by themselves and by other males using olfactory cues. This behavior has been suggested as an analogue to the rejection of alien eggs by avian victims of [[nest parasitism]].<ref name=loiselle2 /><ref name=loiselle1 /> Similarly, pupfish eggs are typically only consumed by females that do not spawn them; these two factors, coupled with the fact that males show sexual preference towards larger (and therefore more [[fecund]]) females, are consistent with a model for the maximization of filial fitness.<ref name=loiselle2 /> |
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==Biology and ecology== |
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===Motor patterns=== |
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The desert pupfish exhibits characteristic movements that can be grouped into various units as follows: |
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*''Meandering''—the female swims somewhat aimlessly in midwater or near the surface in slightly exaggerated movements. |
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*''Nuzzling''—the male swims directly under the meandering female, tilting his body upwards while keeping his head against or near the abdominal region of the female. |
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*''Contacting''—as the female moves slowly, the male stays beside and slightly behind her, typically maintaining contact either side-to-side or snout-to-side. |
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*''Tilting and nipping''—while swimming slowly, the female tilts her body downwards towards the substrate; from the tilted position, the female takes up a mouthful of substrate and drops her body to the bottom of the habitat. The female then expels the substrate and may nip two or three more times from the horizontal position. This action has been interpreted as food-seeking, and has been observed in other species as a signal for the moment of spawning. |
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*''Halting and sidling''—after nipping, the female will halt, whereupon the male will swim forward and laterally against the female, thrusting the area at the base of his anal fin against the posterior line of the female’s abdomen. |
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*''S-shaping''—seen from above, the pupfish forms an s-shape. Typically, both male and female will couple in parallel s-shapes on the bottom of their habitat during mating. |
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*''Wrapping''—while both fish maintain an s-shape, the male will wrap his anal fin around the posterior area of the female’s belly, forming a crude cup beneath her vent. |
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*''Jerking''—while still in an s-shape, the pupfish will jerk its head in the opposite direction it is situated in, causing a contraction that reverses the direction of the s-shape. During jerking, the female will extrude an egg, while the male presumably releases his sperm. |
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*''Patrolling''—the territorial male will swim forwards in small, fast spurts, then stop abruptly before swimming in another direction. The male will patrol his entire territory in this way. |
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*''Facing and eyeing''—two territorial males will approach each other head-on and face each other eye-to-eye with one to one-half body length between them. |
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*''Arching''—the pupfish will form a c-shape with the concave side facing the opponent fish. |
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*''Tail beating''—a competing male will beat his caudal towards the opponent fish, causing the beating fish to swing away from its opponent. |
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*''Circling''—two fish will circle each other at high speeds in a head-to-tail orientation; this is likely due to each fish simultaneously attempting to charge into the flank of its respective opponent, while dodging the charge of the other by turning. |
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*''Fleeing''—a fleeing pupfish will swim rapidly towards cover, or, in its absence, towards the water's surface. |
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*''Escaping''—a pupfish will dive into the substrate of its habitat and remain motionless.<ref name=barlow1>{{cite journal|last=Barlow|first=George W.|title=Social Behavior of the Desert Pupfish, Cyprinodon macularius, in the Field and in the Aquarium|journal=American Midland Naturalist|date=April 1961|year=1961|month=April|volume=65|issue=2|pages=339-359}}</ref> |
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===Reproduction=== |
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The reproductive behavior of the desert pupfish occurs in characteristic fashion. Typically, males remain in their territory, while females, either alone or in groups, visit them; this is characteristic of [[lek mating|arena-breeding]].<ref name=loiselle2>{{cite journal|last=Loiselle|first=Paul V.|title=Male Spawning-Partner Preference in an Arena-Breeding Teleost Cyprinodon macularius californiensis Girard (Atherinomorpha: Cyprinodontidae)|journal=The American Naturalist|date=December 1982|year=1982|month=December|volume=120|issue=6|pages=721-732}}</ref><ref name=barlow1 /><ref name=cowles1>{{cite journal|last=Cowles|first=Raymond B.|title=Notes on the Ecology and Breeding Habits of the Desert Minnow, Cyprinodon macularius Baird and Girard|journal=Copeia|date=24 April 1934|year=1934|month=April|volume=1934|issue=1|pages=40-42}}</ref> If a single female enters his territory, spawning occurs immediately; however, if multiple females are present, the male will swim from one female to the next until the proper response is given, whereupon spawning will occur. Females out of [[oestrus]] will continuously forage. Rarely, a male from a neighboring territory will rush to the side of the female opposite the resident male and spawn with the pair.<ref name=barlow1 /> |
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When a female is ready to spawn, she will typically nip and tilt every 3 to 5 seconds. At this point, the male will immediately contact and attempt to sidle the female. After the third to fifth nip, the female will halt, allowing the male to fully sidle. Sidling by the male causes the female to s-shape, which causes to male to s-shape and concurrently wrap his anal fin around the female’s vent. After the female is wrapped, both fish jerk; jerking typically causes the female to release one egg, while ostensibly causing the male to fertilize said egg. Though the pair usually separates to leave one fertilized egg in the substrate, the female will very occasionally immediately s-shape and both fish will again perform the sequence through jerking.<ref name=barlow1 /> |
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Spawning lasts from 30 minutes to 2 hours, with larger females spawning more than smaller females. Typically, the last coupling of the pupfish is the most prolonged, and several eggs are laid in succession. After [[copulation]], both fish lie in the substrate, and after about 2 seconds, the male will contact the female again. If the female is finished spawning, she will flee in a “panicky” style with the male in pursuit. This chase lasts for about 5 minutes before the male swims to the bottom and both fish begin foraging.<ref name=barlow1 /> |
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===Breeding=== |
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The breeding season of the desert pupfish typically occurs during early spring and into winter when water temperatures exceed about 20ºC. Under conditions of abundant food and suitable temperature, the desert pupfish may become sexually mature as early as six weeks of age. Though they may breed during their first summer, most pupfish do not reproduce until their second summer.<ref name=moyle1>{{cite book|last=Moyle|first=Peter|title=Inland Fishes of California|year=1976|publisher=University of California Press}}</ref> Breeding behavior includes both territorial arena-breeding (high aggression) and consort-pair breeding (low aggression). The former occurs in large habitats with high primary productivity, limited breeding substrates, and high population density and is most common; the latter occurs in habitats with low primary productivity, abundant breeding substrates, and low population density.<ref name=barlow1 /><ref name=kodric1>{{cite book|last=Kodric-Brown|first=A.|title=Fishes in North American Deserts|year=1981|publisher=John Wiley & sons, Incorporated|pages=205-235|editor=R. Naiman, D. Soltz|chapter=Variable breeding systems in pupfishes (genus Cyprinodon): adaptations to changing environments.}}</ref> |
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During the breeding season, male pupfish establish and actively patrol and defend individual territories that are typically less than 1 meter deep and 1-2 meters in area; these territories vary in size based on individual fish size, density of other male pupfish, and water temperature, and are associated with small structures or incongruities of substrate.<ref name=barlow1 /><ref name=moyle1 /> Adult females will swim in loose schools and forage inconspicuously. When a female is ready to spawn, she will leave the school to enter the territory of an attractive male, and the mating process described above will commence.<ref name=usfws93 /><ref name=barlow1 /><ref name=cowles1 /> |
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==Extreme conditions== |
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''C. macularius'' are [[euryhaline]] and [[eurythermal]], tolerating temperatures between approximately 4º and 45ºC and [[salinity|salinities]] ranging from 0 to 70 parts per thousand, exceeding the tolerances of virtually all other [[freshwater]] fish. The desert pupfish can also survive dissolved-oxygen concentrations as low as 0.13 ppm.<ref name=thompson1 /><ref name=legner1 /><ref name=lowe1>{{cite journal|last=Lowe|first=Charles H.|coauthors=Wallace G. Heath|title=Behavioral and Physiological Responses to Temperature in the Desert Pupfish Cyprinodon macularius|journal=Physiological Zoology|date=January 1969|year=1969|month=January|volume=42|issue=1|pages=53-59}}</ref><ref name=usfws93 /><ref name=drecp1 /> It has been noted that the desert pupfish are more abundant in environments with high vegetative cover, [[pH]], and salinity, and low dissolved oxygen and sediment factors, suggesting that living in extreme conditions allows them to survive in environments that are hostile to invasive or nonnative fishes.<ref name=drecp1 /> |
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==Management factors== |
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Listed as endangered in 1986, the desert pupfish has been the focus of many conservation studies and attempts.<ref name=fr1 /> The threats to desert pupfish survival include the destruction of habitat, including loss and/or degradation of habitat through groundwater pumping and water diversion; contamination from agricultural return flows and other contaminants; and physical changes in water quality. Disease and predation due to invasive and nonnative species, as well as environmental competition and behavioral interference of these species are cited as possible threats, though the direct effects of these factors are unknown.<ref name=drecp1 /> |
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Various management factors have been suggested as beneficial towards saving the desert pupfish, including the control and management of exotic and invasive species in pupfish habitats; the maintenance of water levels, quality, and the proper functioning condition of ponds, springs, and drains; the restoration and enhancement of degraded habitat; and the establishment of large primary refuge populations for each group of wild ''C. macularius''.<ref name=usfws10 /><ref name=drecp1 /> The reintroduction of the desert pupfish to wild habitats has been moderately successful, with the establishment of captive or refuge desert pupfish populations has being more fruitful. Water management has been largely cited as having the greatest impact on the survival of ''C. macularius''.<ref name=usfws10 /> |
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==See also== |
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* [[Pupfish]] |
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*''[[Cyprinodon]]'' - Genus |
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* [[Death Valley pupfish]], Salt Creek pupfish, ''Cyprinodon salinus'' |
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* [[Shoshone Pupfish]], ''Cyprinodon nevadensis shoshone'' |
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* [[Tecopa Pupfish]], ''[[Cyprinodon nevadensis calidae]]'' (extinct) |
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*[[Owens pupfish]] ‘’Cyprinodon radiosus’’ |
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*[[Devil's Hole pupfish]] ‘’Cyprinodon diabolis’’ |
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*[[:Category:Cyprinodon|Category: ''Cyprinodon'']] (Pupfish) |
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* [[Julimes pupfish]], ''Cyprinodon julimes'' |
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==References== |
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{{reflist|colwidth=30em}} |
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==External links== |
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* [http://www.desertmuseum.org/books/nhsd_fish.php#pupfish Arizona Desert Museum: - Pupfish] |
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* [http://www.azgfd.gov/w_c/edits/documents/Cyprmacu.fo_000.pdf List of Arizona Native Fishes] |
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*{{FishBase_species|genus=Cyprinodon|species=macularius|year=2010|month=January}} |
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[[Category:Cyprinodon|macularius]] |
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[[Category:Cave fish]] |
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[[Category:Fauna of Northern Mexico]] |
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[[Category:Fauna of the Colorado Desert]] |
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[[Category:Fauna of the Mojave Desert]] |
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[[Category:Fauna of the Sonoran Desert]] |
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[[Category:Fish of the Western United States]] |
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[[Category:Endemic fauna of the United States]] |
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[[Category:Colorado River]] |
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[[Category:Gila River]] |
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[[Category:Endangered fauna of California]] |
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[[Category:Endangered fauna of the United States]] |
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[[Category:Animals described in 1853]] |
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Revision as of 22:57, 29 October 2013
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