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Pupfish

Pupfish, members of the genus Cyprinodon within the family Cyprinodontidae, are small-bodied teleost fishes renowned for their exceptional adaptability to extreme aquatic environments, including waters with temperatures ranging from 0°C to 44°C, salinities from fresh to hypersaline, and low oxygen levels.^[1]^[2] Typically measuring 5–8 cm in total length, these egg-laying species exhibit external fertilization and lack a gonopodium in males, distinguishing them from related livebearing fishes.^[1]^[3] The genus Cyprinodon comprises approximately 49 recognized species, primarily distributed across southwestern North America (including the United States and Mexico), the Caribbean Islands, Central America, and parts of northern South America.^[4] These pupfishes inhabit diverse secondary freshwater, brackish, and occasionally coastal marine habitats, such as desert springs, thermal pools, marshes, saline lakes, and ephemeral streams that may dry seasonally.^[1] Their preference for isolated, often harsh environments has driven rapid speciation and endemism, but also vulnerability to habitat alteration.^[2] Biologically, pupfishes are omnivorous, feeding on algae, diatoms, small invertebrates, and detritus, with diet varying by season and habitat availability.^[2] Reproduction occurs year-round in stable habitats but peaks in spring, with females producing multiple clutches of adhesive eggs that hatch within days; breeding success is highly sensitive to temperatures above 32°C, beyond which egg viability drops sharply.^[2] Notable adaptations include broad thermal tolerances with critical thermal maxima of 39–43°C, potential for anaerobic metabolism in oxygen-poor waters, and morphological plasticity that enables survival in fluctuating conditions.^[2] Many Cyprinodon species face severe conservation challenges, with many classified as endangered or critically endangered due to habitat loss from groundwater depletion, invasive species, and climate change; iconic examples include the Devils Hole pupfish (C. diabolis), confined to a single geothermal pool in Nevada whose population dropped to 38 individuals in spring 2025 due to earthquakes but showed signs of recovery by fall 2025, and the desert pupfish (C. macularius), restricted to isolated refugia in the southwestern United States.^[5]^[3]^[6]^[7] Active recovery efforts by agencies like the U.S. Fish and Wildlife Service involve captive breeding, habitat restoration, and legal protections under the Endangered Species Act to preserve these resilient yet imperiled fishes.^[8]

Taxonomy and Classification

Genera and Species Diversity

Pupfish belong to the family Cyprinodontidae, which is classified within the order Cyprinodontiformes. This family encompasses 10 recognized genera, reflecting a moderate level of taxonomic diversity among small killifishes adapted to extreme aquatic environments.^[9] The genus Cyprinodon represents the most species-rich group, containing 49 species that are distributed across North and Central America, often in isolated springs and saline waters. Other notable genera include Aphanius (primarily Old World species), Empetrichthys (endemic to western North America), Floridichthys, Jordanella, Cubanichthys, Cualac, Garmanella, and Megupsilon. These genera collectively highlight the family's evolutionary radiation into varied ecological niches, from freshwater streams to hypersaline lagoons. Recent taxonomic studies suggest potential revisions for some genera, such as the exclusion of Andean Orestias from Cyprinodontidae.^[9]^[10]^[11] According to Eschmeyer's Catalog of Fishes, the family Cyprinodontidae includes 103 valid species. This diversity includes both extant and extinct forms, with several species facing severe conservation challenges due to habitat loss. Notable examples include the Devils Hole pupfish (Cyprinodon diabolis), restricted to a single geothermal pool in Nevada, and the desert pupfish (Cyprinodon macularius), found in fragmented habitats across the southwestern United States and Mexico. Among extinct species, the Catarina pupfish (Megupsilon aporus) represents a significant loss; the last known individual died in captivity in 2014, marking the genus's complete extinction.^[12]^[13] Subspecies distinctions are particularly prominent within certain complexes, such as the Death Valley pupfish group under Cyprinodon nevadensis, which includes forms like C. n. nevadensis (Saratoga Springs pupfish), C. n. amargosae (Amargosa River pupfish), C. n. mionectes (Ash Meadows Amargosa pupfish), and the extinct C. n. calidae (Tecopa pupfish). These subspecies exhibit subtle morphological and genetic variations adapted to localized thermal and salinity gradients in the region.

Global Distribution

Pupfish of the genus Cyprinodon are predominantly distributed across North America, with the core range concentrated in the arid southwestern United States—including states such as California, Arizona, Nevada, New Mexico, and Texas—and northern Mexico, where over 30 of the 49 recognized species occur.^[14]^[14] This distribution extends eastward into the southeastern U.S. along coastal drainages and southward into the Yucatán Peninsula of Mexico and Belize, while also reaching the Caribbean islands, such as Cuba, the Bahamas, and Hispaniola (shared by Haiti and the Dominican Republic).^[15]^[16] Minor occurrences are noted in northern South America, including coastal regions of Colombia and Venezuela.^[17] The West Indies harbor several insular populations, contributing to the genus's fragmented Neotropical presence.^[16] Key endemic hotspots underscore the genus's concentration in isolated aquatic systems. In the United States, Death Valley National Park in California hosts the Devils Hole pupfish (C. diabolis), confined to a single geothermal spring within a limestone cavern.^[18] In Mexico, the Cuatro Ciénegas Basin in Coahuila represents a biodiversity hotspot with a species flock adapted to desert oases, including several Cyprinodon taxa restricted to spring-fed pools and saline wetlands.^[19] Similarly, San Salvador Island in the Bahamas supports a remarkable assemblage of endemic pupfishes, such as C. sator and C. desquamator, limited to hypersaline lakes formed by karst topography.^[20] These sites highlight regional endemism driven by geographic barriers. Historically, pupfish ranges were more extensive, but many populations have been extirpated due to habitat alterations and invasive species introductions. For instance, the desert pupfish (C. macularius) once inhabited broader stretches of the Colorado River Delta and associated basins in the southwestern U.S. and Mexico, but natural populations have been eliminated from Arizona and severely reduced in California and Sonora.^[21] In Mexico's Lake Chichancanab on the Yucatán Peninsula, a endemic radiation including the boxer pupfish (C. simus) and largefin pupfish (C. verecundus) has been driven to functional extinction in the wild, with wild populations lost primarily to competition from introduced cichlids; as of 2025, captive breeding efforts continue to prevent total extinction.^[22]^[23]^[24] Endemism in Cyprinodon arises from patterns of habitat isolation, particularly in spring-fed aquifers and closed desert basins that fragment populations during aridification events. These refugia, such as desert springs and endorheic lakes, promote speciation through vicariance, as seen in the southwestern North American clade where ancestral lineages diverged in disconnected water bodies following Pleistocene climate shifts.^[20]^[25] Such isolation has resulted in high levels of local adaptation and narrow ranges for many species.

Physical Description and Adaptations

Morphology and Size

Pupfish in the genus Cyprinodon are small, ray-finned teleosts exhibiting an elongate body form with a robust, often deep profile that is accentuated in breeding males. They feature a terminal or slightly inferior mouth, large eyes positioned high on the head, and rounded fins, including a dorsal fin originating at or near the midbody and a caudal fin with 14–22 rays. Scales are typically large and ctenoid, covering the body without preorbital scales in some species.^[26]^[27]^[28] Adults generally range from 5 to 8 cm in total length, with species-specific variations; for instance, the Amargosa River pupfish (C. nevadensis amargosae) rarely exceeds 5 cm, while the desert pupfish (C. macularius) can reach up to 7.5 cm. The smallest species, such as the Devils Hole pupfish (C. diabolis), average around 2.3 cm and maximize at 4.3 cm.^[27]^[29]^[5] Sexual dimorphism is evident in size and coloration, with males typically larger than females and displaying brighter hues during the breeding period, including iridescent blue on the head and sides, yellow on the caudal peduncle, and black margins on the fins. Females are smaller, with subdued olive-brown or grayish tones and faint vertical bars along the sides. In certain Death Valley endemics like C. diabolis, pelvic fins are absent or greatly reduced, contributing to a more streamlined appearance.^[27]^[28]^[30]

Physiological Adaptations

Pupfish exhibit extraordinary physiological tolerances that allow them to inhabit environments lethal to most other fish species, including extreme temperatures, salinities, and oxygen levels. These adaptations involve specialized osmoregulatory, respiratory, and metabolic processes that maintain homeostasis under chronic stress. For instance, species within the genus Cyprinodon can endure water temperatures ranging from near freezing to over 42°C, with some populations demonstrating critical thermal maxima exceeding 45.5°C (114°F).^[31] Similarly, the Devils Hole pupfish (Cyprinodon diabolis) thrives in a geothermal pool with stable temperatures around 34.4°C (94°F) and dissolved oxygen concentrations often below 2.5 mg/L, conditions that approach the lethal limits for many aquatic vertebrates.^[32]^[33] Osmoregulation in pupfish is highly efficient, enabling survival across a salinity gradient from freshwater to hypersaline conditions exceeding 200 ppt (g/L), far beyond typical seawater levels of 35 ppt. This is facilitated by active ion transport mechanisms in the gills and kidneys, where Na⁺-K⁺ ATPase activity in gill epithelia increases dramatically in low-salinity environments to promote ion uptake and decreases in high-salinity settings to minimize ion loss.^[29]^[34] Pupfish acclimate rapidly to salinity shifts, adjusting osmoregulatory function within hours through hormonal regulation, such as arginine vasotocin, which modulates ion and water flux across gill and kidney tissues.^[35]^[36] Low oxygen tolerance is supported by respiratory adaptations, including enhanced oxygen delivery to tissues via hemoglobin with population-specific oxygen affinities that optimize unloading in hypoxic waters below 2 mg/L.^[37]^[38] Thermal stress triggers protective metabolic responses in pupfish, including the upregulation of heat-shock proteins that stabilize cellular proteins and mitigate oxidative damage at high temperatures.^[39] These fish also demonstrate metabolic flexibility, with enzymatic adjustments allowing sustained activity and reproduction in hot springs where temperatures fluctuate widely, such as up to 47.5°C in some habitats.^[40] In fluctuating arid environments, pupfish exhibit resistance to starvation, enduring prolonged periods without food through efficient energy conservation and lipid storage, which supports survival during seasonal resource scarcity.^[41] These integrated adaptations underscore the pupfish's resilience, with their compact body form further aiding heat dissipation in confined, warm waters.

Habitat and Ecology

Environmental Preferences

Pupfish primarily inhabit shallow, spring-fed pools, desert streams, saline lakes, and geothermal waters across arid and semi-arid regions. These environments are often characterized by clear waters in vegetated shallows supported by rocky or sand-silt substrates, which provide stable microhabitats amid surrounding harsh conditions.^[42]^[43]^[44] Key abiotic factors influencing pupfish distribution include a wide pH range from slightly acidic to highly alkaline conditions (approximately 6 to 9), low water flow rates in lentic or pool-dominated systems, and seasonal drying cycles typical of desert aquatic habitats. These species thrive in low-oxygen environments and exhibit remarkable tolerance to fluctuations in these parameters, allowing persistence in otherwise marginal waters.^[45] Recent drying events, exacerbated by climate change and groundwater depletion, have led to habitat loss in some regions, such as the Leon Springs area as of 2024.^[46] Within these habitats, pupfish often utilize specific microhabitats as refugia, such as the shallow underwater limestone shelves in sites like Devils Hole, which buffer against temperature extremes and maintain consistent conditions for daily activities.^[5]^[18] Regional variations highlight the adaptability of pupfish to diverse abiotic regimes; for instance, populations in Mexico's Cuatro Ciénegas Basin endure hot, hypersaline conditions exceeding 95 g/L salinity in shallow, thermally variable lakes, while those in Bahamian blue holes occupy cooler, often freshwater-influenced karst sinkholes with overlying oxic layers.^[47]^[20]^[48] Such tolerances align with their physiological adaptations to high-heat geothermal springs in various locales.^[49]

Diet and Foraging Behavior

Pupfish exhibit an omnivorous diet, primarily consisting of algae, diatoms, detritus, insects, and small crustaceans such as ostracods and copepods.^[50]^[8] In resource-poor environments, such as isolated desert springs, they shift toward greater reliance on algae and detritus to sustain energy needs, reflecting their opportunistic feeding habits.^[8] This dietary flexibility allows pupfish to exploit limited food resources in extreme habitats, where invertebrate availability may be low.^[51] Foraging strategies among pupfish species include surface picking for insects at the air-water interface, bottom scraping to remove epilithic algae from substrates, and mid-water schooling to capture drifting detritus or small prey.^[52] These behaviors are facilitated by morphological adaptations, such as upturned mouths suited for surface feeding in many species.^[52] In nutrient-scarce habitats, pupfish engage in opportunistic foraging, actively scraping limited algal growth or picking sparse invertebrates during early morning and late evening when activity peaks.^[43] Social dynamics influence foraging, with non-breeding individuals often forming loose schools for mutual protection while foraging in open water.^[43] In denser populations, aggression over food resources increases, leading to defense of small feeding territories through displays like charging and tail-beating, particularly when schools disperse to feed. For example, the Devils Hole pupfish (Cyprinodon diabolis) relies almost exclusively on epilithic algae and diatoms scraped from a single 20 m² limestone shelf, with diet shifting seasonally from diatom dominance in winter-spring to cyanobacteria and green algae in summer.^[50] Similarly, desert pupfish (Cyprinodon macularius) in intermittent streams show seasonal diet variations, consuming more insect larvae and snails in wetter periods and detritus during dry seasons.^[43]

Reproduction and Life History

Mating Systems

Pupfish typically exhibit a polygynous mating system, in which males establish and vigorously defend small territories in shallow, warm waters to attract multiple females for spawning.^[53] During courtship, territorial males display vibrant nuptial coloration—often intensifying to blues, oranges, or blacks on the body and fins—while performing zig-zag swims, looping maneuvers, and fin flaring to signal readiness and dominance.^[54] These displays, combined with herding behaviors to isolate receptive females, facilitate mate attraction, though some species like the Devils Hole pupfish (Cyprinodon diabolis) show reduced territoriality outside of brief mating interactions.^[28] Breeding occurs year-round in stable, warm-spring habitats but is largely seasonal in fluctuating desert environments, typically from spring through autumn when water temperatures exceed 20°C and photoperiod increases.^[8] Optimal reproductive temperatures range from 28–29°C, with spawning triggered by rising temperatures and longer daylight hours that stimulate gonadal development in both sexes.^[55] Females, often gravid with multiple egg batches, exercise mate choice by preferring males demonstrating vigorous displays and intense coloration, which correlate with higher reproductive success.^[53] A single female may spawn repeatedly, releasing a few eggs (typically 1–5) per spawning act in a broadcast manner directly onto the substrate or vegetation without nest-building, with females often spawning multiple times per day and laying up to 50–200 eggs per week, allowing fertilization by the territory holder or occasionally intruding males.^[56]^[57] Variations in mating strategies exist across species; for instance, in the Comanche Springs pupfish (Cyprinodon elegans), male-male combat is particularly intense, involving aggressive chases, bites, and displays to secure prime spawning sites, with larger males dominating territories while smaller "satellite" males opportunistically intercept spawnings.^[58] In contrast, some populations shift to less territorial, school-based systems under high-density or low-resource conditions, reducing aggression but maintaining polygynous pairings through persistent courtship.^[59] These adaptations highlight the flexibility of pupfish mating systems in response to environmental pressures.^[60]

Larval Development and Growth

Pupfish reproduction is oviparous, involving external fertilization during spawning events where males release milt over eggs laid by females on suitable substrates like algae-covered rocks or limestone ledges. Egg development duration is temperature-dependent, typically ranging from 4 to 10 days, with hatching producing larvae measuring 4–6.5 mm in total length. For instance, in the desert pupfish (Cyprinodon macularius), eggs of about 1 mm in diameter hatch after approximately 10 days at 20°C.^[29]^[28] Upon hatching, pupfish larvae exhibit rapid initial growth and behavioral adaptations for survival, including active foraging on microalgae, detritus, and small invertebrates. Larvae often form schools shortly after emergence, which helps mitigate predation risks in their shallow, open habitats. Growth rates are notably fast, with many species reaching about 25 mm (1 inch) within the first 1–3 months; for example, Devils Hole pupfish (Cyprinodon diabolis) larvae grow at an average of 0.65 mm per week under natural conditions. However, early larval stages experience high mortality, often exceeding 90% in the first few weeks, primarily due to predation by invertebrates and conspecifics, as well as fluctuating water levels and temperatures.^[28]^[18] Pupfish reach sexual maturity relatively quickly, typically at 2–3 months of age and sizes of 15–30 mm standard length, enabling multiple reproductive cycles within their short lifespan. In the wild, individuals generally live 1–2 years, though maximum recorded lifespans reach up to 2.5 years in some populations; captive conditions can extend this to 3 years by reducing stressors.^[29]^[61]^[43] Larval growth and development are strongly influenced by environmental conditions, with warmer temperatures (e.g., 28–34°C) accelerating hatching times and increasing growth rates compared to cooler regimes. Abundant food availability in nutrient-rich springs further enhances somatic growth and survival. Captive breeding programs, such as those for the endangered desert pupfish, demonstrate these effects by maintaining optimal temperatures and feeding regimes, resulting in faster maturation (as little as 30–45 days to adulthood in some hybrids) and higher juvenile survival rates than observed in the wild.^[62]^[63]

Evolutionary History

Fossil Record and Origins

The family Cyprinodontidae, to which pupfish belong, originated through divergence within the order Cyprinodontiformes during the Miocene epoch, with the earliest known fossils dating to the Oligocene in Europe.^[64] These include species of the genus Prolebias, such as P. aymardi and P. cephalotes, preserved in southern French deposits and representing early cyprinodontid forms adapted to freshwater and brackish environments. The family's radiation is estimated to have occurred around 23 million years ago, coinciding with the late Oligocene expansion of diverse killifish lineages across Eurasian paleolakes and coastal systems.^[65] Ancestral cyprinodontids likely evolved from marine teleosts, transitioning to euryhaline and freshwater habitats as part of broader adaptations in the Tethyan region.^[66] Migration patterns involved dispersal from the Old World to the Americas, facilitated by ancient land bridges or vicariance events across the proto-Atlantic and Tethys Sea connections during the late Eocene to early Miocene.^[14] This colonization enabled pupfish ancestors to exploit inland riverine and lacustrine systems, marking a shift from oceanic origins to continental endemism. In the New World, pupfish-like forms of the genus Cyprinodon appear in the fossil record by the late Miocene, reflecting early adaptations to arid and hypersaline conditions in western North American basins.^[14] Key examples include C. breviradius from Neogene deposits in Death Valley, California, which exhibits morphological traits suited to fluctuating desert aquatic environments.^[67] Similar pupfish fossils from Miocene basins in northern Mexico indicate contemporaneous diversification along coastal and interior drainages, such as ancestral extensions of the Rio Grande and Gulf of Mexico systems. By the late Miocene, approximately 7–9 million years ago, Cyprinodon had established a widespread presence from the Atlantic slope of Mexico to California, setting the stage for further inland radiations.^[14]

Rapid Speciation and Diversification

Pupfish exemplify accelerated evolutionary processes, with speciation and diversification rates far exceeding those typical for teleost fishes. In the Bahamian lineage on San Salvador Island, morphological evolution proceeds at rates 50–130 times faster than in other Cyprinodon species, enabling the rapid assembly of a species flock through the occupation of novel ecological roles.^[68] Similarly, Death Valley pupfish lineages diversify reflecting post-Pleistocene isolation and adaptation to extreme desert aquifers.^[23] These patterns highlight pupfish as a model for understanding how small, fragmented populations can generate substantial biodiversity in geologically brief intervals. The primary mechanisms driving this rapidity involve allopatric isolation in disconnected habitats, such as geothermal springs and hypersaline lakes, which impose bottlenecks and founder effects that amplify genetic drift.^[69] In tandem, ecological niche partitioning fosters divergence by favoring adaptations to specific environmental axes, including salinity gradients, water depth, and temperature regimes. For example, on San Salvador Island, pupfish populations exploit varying conditions within shallow, hypersaline lakes (40–50 ppt salinity, depths up to 12 m), where species differentiate along thermal clines and foraging strata to reduce competition.^[20] In Death Valley, isolation in springs with distinct chemistries—ranging from freshwater outflows to highly saline pools—similarly promotes speciation via localized selection on osmoregulatory and thermal tolerance traits.^[70] Illustrative cases underscore the tempo of these events. The Death Valley pupfish complex arose less than 10,000 years ago, following the drying of Pleistocene lakes that stranded ancestors in isolated refugia, leading to the emergence of multiple endemic forms like the Devils Hole pupfish (Cyprinodon diabolis).^[71] On San Salvador Island, a radiation of at least three Cyprinodon species has unfolded within 6,000–10,000 years, filling niches across a mosaic of blue holes and lakes with divergent temperature profiles, from near-boiling geothermal inflows to cooler margins.^[20] These examples demonstrate how vicariance and opportunity in marginal habitats can catalyze flock formation from Miocene progenitors.^[72] Genomic analyses reveal the underpinnings of this swift divergence, including elevated allelic differentiation (e.g., F_ST ≈ 0.40 in related systems) despite minimal temporal separation, often exceeding expectations under neutral models.^[69] Founder effects during colonization reduce initial genetic diversity, while bottlenecks intensify drift, as seen in the Devils Hole population's recent establishment (105–830 years ago) and subsequent fixation of traits like reduced pelvic fins through introgression and selection.^[70] In Bahamian clades, directional selection on craniofacial morphology drives adaptive peaks, with hybridization occasionally introducing variation that accelerates niche invasion.^[73] Collectively, these factors—drift, isolation, and selection—enable pupfish to achieve reproductive isolation and ecological specialization in under millennia.

Conservation Status

Major Threats

Pupfish populations face severe threats from anthropogenic activities that have drastically reduced their already limited habitats in desert springs, streams, and isolated aquatic systems. Primary among these is habitat loss driven by water diversion and groundwater pumping for agriculture and urban development, which has caused the drying of essential desert water sources across the southwestern United States and northern Mexico. For instance, extensive pumping in the Amargosa Valley has lowered water tables, threatening the sole habitat of the Devils Hole pupfish by reducing flow in its geothermal spring.^[42] Similarly, channelization and impoundment projects have dewatered streams like those feeding the Salton Sea, leading to the extirpation of pupfish from numerous historic sites since the late 19th century.^[74] Climate change exacerbates these pressures through rising temperatures and prolonged droughts, which alter hydrology and increase thermal stress in pupfish habitats. Higher air temperatures have been linked to reduced reproduction in species like the Devils Hole pupfish, as warmer waters exceed optimal ranges and diminish dissolved oxygen levels critical for survival.^[75] Droughts, intensified by climate shifts, further diminish spring flows and concentrate salinity, pushing pupfish beyond their physiological tolerances in isolated refugia.^[76] Pollution from agricultural runoff introduces toxins and elevates salinity, while predation by invasive species compounds mortality, particularly among larvae. Runoff carrying pesticides and nutrients has contaminated sites like Quitobaquito Springs and San Felipe Creek, causing direct toxicity and indirect effects through algal overgrowth that disrupts foraging.^[42] Non-native fish, such as tilapia (Oreochromis spp.) and mosquitofish (Gambusia affinis), prey on pupfish eggs and juveniles or outcompete them for resources; tilapia introductions in Mexican drainages have displaced desert pupfish populations by altering food webs and increasing predation pressure.^[77]^[43] Specific events underscore the vulnerability of pupfish to both natural and human-induced catastrophes. In late 2024 and early 2025, earthquakes in Nevada generated waves in Devils Hole that dislodged essential algae and eggs from the shallow spawning shelf, reducing the pupfish population to just 38 individuals by spring 2025—the lowest count in decades.^[6] Historically, 19th- and 20th-century water developments led to widespread extirpations, including the Tecopa pupfish (Cyprinodon nevadensis calidae), declared extinct in 1981 due to habitat desiccation from irrigation diversions, and the desert pupfish from over 90% of its former range in the Salton Sea basin by the mid-20th century.^[78]^[79]

Recovery Initiatives and Current Status

Recovery initiatives for pupfish species, primarily coordinated by the U.S. Fish and Wildlife Service (USFWS) and partner agencies like the National Park Service (NPS), focus on habitat protection, captive propagation, reintroduction programs, and ongoing population monitoring to address their endangered status. These efforts stem from species-specific recovery plans established under the Endangered Species Act, emphasizing the establishment of self-sustaining populations in protected refuges and the mitigation of threats such as groundwater depletion and invasive species. For instance, the 1993 Desert Pupfish Recovery Plan, amended in 2019, outlines criteria for delisting, including the protection of at least three secure habitat sites with viable populations exceeding 1,000 individuals each, though progress has been incremental due to persistent habitat degradation. The Devils Hole pupfish (Cyprinodon diabolis), one of the most critically endangered pupfish, exemplifies intensive recovery actions. Confined to a single geothermal pool in Nevada's Devils Hole within Death Valley National Park, its population peaked at 191 individuals in spring 2024 but plummeted to just 38 by spring 2025 following sediment disturbances from two earthquakes in December 2024 and February 2025, which dislodged food sources and eggs from the shallow limestone shelf. In response, a multi-agency team, including USFWS and NPS, released 19 captive-raised juveniles from the Ash Meadows Fish Conservation Facility in March 2025 to bolster numbers, supplemented by targeted feeding to restore algae-based diets. Biannual surveys, conducted since 1972 using surface observations and SCUBA dives, continue to track recovery, with fall 2025 counts anticipated to show spawning improvements amid stabilizing habitat conditions.^[80]^[5] For the Desert pupfish (Cyprinodon macularius), listed as endangered since 1986, recovery emphasizes reintroduction into restored habitats across Arizona, California, and Mexico. Recent efforts include the March 2025 reintroduction of more than 450 individuals to Ayer Lake at Boyce Thompson Arboretum under a Safe Harbor Agreement with the Arizona Game and Fish Department, aiming to create replicate populations while controlling invasive mosquitofish. In October 2025, 48 desert pupfish were released into a constructed stream at the University of Arizona's Biosphere 2 as part of efforts to establish additional refuge populations.^[81] Natural populations persist in California's Salton Sea and Sonora, Mexico, but remain vulnerable; USFWS's June 2025 status update confirms extirpations in parts of Arizona, with reintroductions in artificial ponds and streams yielding small but stable groups of 50–200 fish per site.^[30]^[82] Other species, such as the Comanche Springs pupfish (Cyprinodon elegans), benefit from similar strategies outlined in their 1981 recovery plan, including maintenance of refuge populations at Dexter National Fish Hatchery for potential reintroductions if wild numbers decline further. The Pecos pupfish (Cyprinodon pecosensis) faces a proposed uplisting to threatened status in November 2024 by USFWS, with conservation focusing on barrier construction to exclude non-native predators in New Mexico and Texas rivers. Overall, while pupfish populations remain precarious— with many below 500 individuals— these initiatives have prevented extinctions and fostered genetic diversity through captive programs, though full recovery requires sustained habitat restoration amid climate pressures.^[83]^[84]

References

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FAMILY Details for Cyprinodontidae - Pupfishes
### Summary of Cyprinodontidae (Pupfishes) and Genus Cyprinodon
[2]
[PDF] Thermal and Developmental Ecology of Pupfish, Cyprinodon
Pupfish from the genus Cyprinodon are often cited as having some of the widest thermal tolerances of any fish species. Sheepshead Minnow, Cyprinodon ...
[3]
Desert Pupfish (Cyprinodon macularius) | U.S. Fish & Wildlife Service
Small (2 inches) smoothly rounded body shape with narrow vertical bars on the sides. Breeding males blue on head and sides with yellow on tail.
[4]
https://www.fishbase.se/identification/SpeciesList.php?genus=Cyprinodon
[5]
Devils Hole Pupfish (Cyprinodon diabolis) | U.S. Fish & Wildlife ...
The Devils Hole pupfish resides in the upper 80 feet (24.38 meters) of the body of water, with typically half of the population residing on the limestone shelf.
[6]
[PDF] Desert pupfish(Cyprinodon macularius) 5-Year Review - ECOS
Mar 31, 1986 · Naturally occurring populations of desert pupfish (Cyprinodon m. macularius or C. macularius) are now restricted in the United States to two ...
[7]
FAMILY Details for Cyprinodontidae - Pupfishes - FishBase
Nov 29, 2012 · Family Cyprinodontidae - Pupfishes. Order, : Cyprinodontiformes · Cyprinodontidae. Class, : Teleostei. No. in FishBase, : Genera : 10 ...
[8]
Fish Identification: Find Species - FishBase
Class: Teleostei Order: Cyprinodontiformes Family: Cyprinodontidae Pupfishes Subfamily: Cyprinodontinae Genus: Cyprinodon (See list of species below) ...
[9]
CAS - Eschmeyer's Catalog of Fishes - Genera/Species by Family/Subfamily
- **Valid Genera for Cyprinodontidae**: 19
[10]
The extinction of the Catarina pupfish Megupsilon aporus and the ...
Dec 3, 2018 · Here we report the extinction of an endemic freshwater fish of Mexico, the Catarina pupfish Megupsilon aporus, the sole species of the genus Megupsilon.
[11]
(PDF) Historical Biogeography of the New-World Pupfish Genus ...
Aug 6, 2025 · ican highlands). Approximately 30 of the 50 or so species of. Cyprinodon occur in arid regions of southwest-. ern North America where they ...
[12]
Species: Cyprinodon artifrons, Yucatan Pupfish
In shallow water with vegetated bottoms, freshwater to full seawater. Depth range: 0-2 m. N Yucatan peninsula to Belize.
[13]
Cyprinodon): Implications for the biogeography of insular Caribbean ...
Aug 7, 2025 · ... genus Cyprinella are abundant and broadly distributed in eastern and central North America. Thirty species are currently placed in the genus ...
[14]
A cryptic lineage within the pupfish Cyprinodon dearborni suggests ...
Its reported coastal distribution extends from the Guajira Peninsula (12° 25′ N; 71° 30′ W) in Colombia to Isla de Margarita, Venezuela (11° 00′ N; 64° 00′ W), ...
[15]
Life in a fishbowl: Prospects for the endangered Devils Hole pupfish ...
Aug 11, 2014 · The Devils Hole pupfish (Cyprinodon diabolis) is a federally listed endangered species living solely within the confines of Devils Hole, a geothermal pool ...
[16]
Predicting the distribution of an introduced fish in an endemic ...
Cuatro Cienegas, in Coahuila, Mexico is an oasis in the Chihuahua desert and an endemic hotspot. A common aquarium trade fish, the West African jewel cichlid ( ...
[17]
[PDF] A Remarkable Species Flock of Cyprinodon Pupfishes Endemic to ...
May 30, 2020 · ABSTRACT. Two new species of Cyprinodon are described, both restricted to saline lakes on a single island in the Bahamas.
[18]
Cyprinodon macularius - Desert Pupfish - NatureServe Explorer
Small range in southern California, Arizona (all natural populations have been extirpated), and northwestern Mexico; range greatly reduced due to loss of native ...
[19]
Whipsnade Zoo attempts to save the Mexican boxer pupfish - BBC
Nov 19, 2024 · It is native to Lake Chichancanab in Mexico, but experts there believe the local population is now extinct. Fifty boxer pupfish eggs have been ...
[20]
Explosive evolution in pupfish - Wainwright Lab
May 5, 2011 · The second remarkable place for pupfish diversification is Lake Chichancanab ... These specialized species are now functionally extinct in the ...
[21]
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[22]
[PDF] Three New Pupfish Species, Cyprinodon (Teleostei ...
Three new species of Cyprinodon (Teleostei, Cyprinodontidae) are described, each long recognized as distinct. Cyprinodon pisteri occupies a variety of ...
[23]
Amargosa River Pupfish - California Fish Website
Small, usually less than 50 mm TL · Blunt head, deep body (especially in reproductive males) · Small and oblique terminal mouth, complete row of tricuspid teeth; ...
[24]
Cyprinodon diabolis (Devils Hole Pupfish) - Animal Diversity Web
Cyprinodon diabolis has been called one of the most geographically isolated species in the world. Its home range is a 3.0 foot (1 m) by 5.0 foot (1.67 m) ...Physical Description · Reproduction · Conservation Status<|separator|>
[25]
[PDF] desert pupfish, cyprinodon macularius
A description adapted from Moyle (1976) and Minckley (1973) is as follows: Males are larger than females, up to 75 mm total length (TL). The body is thick-.
[26]
Species Profile for Desert pupfish(Cyprinodon macularius) - ECOS
Two subspecies are recognized: Desert Pupfish (C.m. macularis) and Quitobaquito Pupfish (C.m. eremus). Critical habitat includes Quitobaquito Springs, Pima ...
[27]
Diabolical survival in Death Valley: recent pupfish colonization ... - NIH
Jan 27, 2016 · One of the most endangered vertebrates, the Devils Hole pupfish Cyprinodon diabolis, survives in a nearly impossible environment: a narrow ...
[28]
[PDF] Effects of oxygen variation on the ecology of Devil's Hole, Death ...
... Hole pupfish, Cyprinodon diabolis Wales, is endemic to Devil's Hole (fig. ... This riffle beetle is not known to survive under high temperature and low oxygen ...Missing: 94° | Show results with:94°
[29]
Spatial and temporal variation of dissolved oxygen and ecosystem ...
... Devils Hole pupfish,Cyprinodon diabolis. With consistently high water temperature (33.5 °C/93 °F) and low dissolved oxygen (O2) concentration (about 2.5 mg ...Missing: levels | Show results with:levels
[30]
Effects of temperature and salinity on gill Na+-K+ ATPase activity in the pupfish, Cyprinodon salinus
### Summary of Pupfish Osmoregulation and Adaptations
[31]
Osmotic acclimation in the Red River pupfish, Cyprinodon ...
Nov 1, 1971 · Abrupt transfer from fresh water to salt water showed that the pupfish could acclimate to increased salinity within 8 hr.
[32]
[PDF] Population divergence in plasticity of the AVT system and its ...
Apr 6, 2001 · In teleost fish, AVT regulates hydromineral balance by altering ion and water transfer in gill and kidney tissues. (Warne, 2002; Warne et al., ...
[33]
Swimming Performance and Energetics as a Function of ...
... blood hemoglobin oxygen affinities between populations, resulting in northern fish having a greater ability to deliver oxygen to tissues at low temperatures.
[34]
Genetic Differences in Physiological Tolerances of Amargosa ...
Amargosa pupfish from a constant-temperature spring have a narrower range of temperature and oxygen tolerances than pupfish from the much more variable ...
[35]
Natural thermal adaptation increases heat shock protein levels and ...
The present results provide evidence that physiologically increased HSP levels provide protection against oxidative stress and enhance cytoprotection.
[36]
Endocrine and metabolic impacts of warming aquatic habitats - NIH
Nov 3, 2016 · Pupfish in this region evolved some of the highest critical thermal maxima and broadest thermal tolerances recorded in freshwater fishes ( ...
[37]
[PDF] Native fishes of arid lands - USDA Forest Service
Native freshwater fishes of the desert Southwest are disappearing due to scarce water and human populations, and have received little attention.
[38]
[PDF] DESERT PUPFISH RECOVERY PLAN - ECOS
Behavioral ecology of two pupfish (Cyprinodontidae,. Genus Cyprinodon) from Northern Mexico. Doctoral Dissertation, Arizona. State University, Tempe. Bagley ...
[39]
Desert Pupfish (Cyprinodon macularis)
Naturally-occurring populations of desert pupfish have been extirpated in Arizona but still occur in the Salton Sink Basin of California and the Colorado River ...Missing: current | Show results with:current
[40]
Desert pupfish - Facts, Diet, Habitat & Pictures on Animalia.bio
Pupfish typically prefer clear waters with either rooted or unattached aquatic plants, restricted surface flow, or sand-silt substrates, and will do well if ...
[41]
Cyprinodontidae - an overview | ScienceDirect Topics
New World pupfish in the genus Cyprinodon are classified into five distinct species groups or clades that are believed to have diverged from a common ancestor 6 ...
[42]
Relation of desert pupfish abundance to selected environmental ...
Pupfish abundance was generally highest in habitats where water quality extremes (especially high pH and salinity, and low dissolved oxygen) seemingly limited ...
[43]
Cuatro Ciénegas, Coahuila, México - Desert Fishes Council
Habitat & Ecology. Coahuilan pupfish occupy habitats varying from hypersaline (to >95 gm/l, and higher), seasonally hot, shallow and otherwise severe lakes ...
[44]
Inland blue holes of The Bahamas - chemistry and biology in a ...
Dec 9, 2020 · The majority of the studied blue holes were vertically stratified with oxic freshwater overlying anoxic saline groundwater of marine origin.Missing: pupfish | Show results with:pupfish
[45]
[PDF] Population ecology and monitoring of the endangered Devils Hole ...
The entire geographic range of the Devils Hole pupfish (Cyprinodon diabolis) resides in this cavern system.
[46]
A comparative study of prokaryotic diversity and physicochemical ...
The small (<30mm in length), energetic, iridescent blue Devils Hole pupfish survive on a diet composed of diatoms, cyanobacteria, filamentous green algae, and ...
[47]
Food Habits of the Amargosa Pupfish in a Thermal Stream
Food habits of an Amargosa pupfish {Cyprinodon nevadensis amargosae) population inhabit- ing a thermal stream were examined at monthly intervals from ...Missing: foraging | Show results with:foraging
[48]
(PDF) Trophic apparatus in cyprinodontiform fishes - ResearchGate
Aug 6, 2025 · Recent studies suggest that this group is well suited to capturing prey by "picking" small items from the water surface, water column, and the ...Missing: schooling | Show results with:schooling
[49]
Determinants of male reproductive success in pupfish (Cyprinodon ...
The importance of male nuptial coloration was demonstrated by increased male reproductive success with intensity of colour of the same individual males ('light' ...
[50]
Behavior of the Pupfish, Cyprinodon rubrofluviatilis - jstor
Correspondingly, males seem maxi- mally aggressive on days when spawning is frequent and minimally aggressive on days when spawning is absent or infrequent. The ...
[51]
[PDF] ' ' DISTIIIBIITION OF THIS OIJCI!Mf'IIS UNliM(J - OSTI
The limits for both growth and reproduction reflect the high temperature adaptation of the pupfish which lives in the Death Valley, California, region (Brown ...
[52]
[PDF] Egg Deposition of the Desert Pupfish, Cyprinodon Macularius, in ...
Crear and Haydock (1971) reported that female pupfish spawned 50 to 200 eggs once per week. The present study shows a similar level of production.Missing: batch | Show results with:batch
[53]
Comanche Springs Pupfish (Cyprinodon elegans)
Comanche Springs pupfish are found in spring pools, spring-influenced marshes (ciénegas), and irrigation canals carrying spring waters. They evolved to be the ...
[54]
The Phenotypic Plasticity of Death Valley's Pupfish
The Warm Spring pupfish (Cyprinodon nevadensis pectoralis) is found near Devil's Hole. Two other species, Cyprinodon nevadensis calidae and Cyprinodon ...
[55]
[PDF] The breeding system of an endangered pupfish (Cyprinodon elegans)
In the quiet pool, unlike in swift water, the breeding system was characterized by 3 different male mating tactics: territorial defense, satel- lite positioning ...Missing: combat | Show results with:combat
[56]
[PDF] Leon Springs pupfish (Cyprinodon bovinus) 5-Year Review - AWS
Aug 8, 2013 · Pupfish reach sexual maturity at 1 in (29 mm) standard length (Kennedy 1977, p. 97); maximum lifespan is 20-23 months in the wild (Kennedy 1977, ...Missing: larval | Show results with:larval
[57]
Thermal Limits for Growth and Reproduction in the Desert Pupfish ...
The overall mean was 75.3%. The same result was found at 36 C, feeding rations of 3.0%, 4.5%, and 6.0% body weight day−1. Assimilation rates at 12 and 16 C were ...
[58]
risk analyses to guide harvest of Devils Hole pupfish for captive ...
Sep 9, 2014 · The Devils Hole pupfish is restricted to one wild population in a single aquifer-fed thermal pool in the Desert National Wildlife Refuge ...Missing: 94° | Show results with:94°
[59]
Cyprinodontiformes (Killifishes and Live-Bearers) - Encyclopedia.com
The earliest fossils, the cyprinodontid Pachylebias and Prolebias, both found in Europe, date to the Oligocene epoch, 25–40 million years ago (mya). Most ...
[60]
(PDF) Multigene phylogeny of cyprinodontiform fishes suggests ...
The fossil record of both families is quite extensive ... Phylogenetic relationships of five species of Prolebias from the Oligocene of southern France (P.
[61]
Closing of the Tethys Sea and the phylogeny of Eurasian killifishes ...
Dec 11, 2002 · Sauvage (1874) described Prolebias, an Aphanius-like cyprinodont, from the middle Oligocene of Europe and Priem (1908) described middle Miocene ...
[62]
fossil fishes of death valley national park, california: reconstructing ...
Included was the only known record of the extinct pupfish Cyprinodon breviradius. ... The tuff beds most likely fall within the range for Miocene tuff beds ...
[63]
TROPHIC NOVELTY IS LINKED TO EXCEPTIONAL RATES OF ...
Mar 21, 2011 · Here, we show that rapid rates of morphological diversification are linked to the evolution of novel ecological niches in two independent ...
[64]
Rapid vertebrate speciation via isolation, bottlenecks, and drift - PNAS
May 21, 2024 · While rapid speciation is often driven by natural or sexual selection, here we show that isolation and drift have led to speciation within a few thousand ...
[65]
Diabolical survival in Death Valley: recent pupfish colonization ...
Jan 27, 2016 · Death Valley pupfishes shared a common ancestor with Northeastern Mexican species approximately 17 kyr ago (95% CI: 9–34 kyr ago) and ...
[66]
New evidence for the recent divergence of Devil's Hole pupfish and ...
Nov 17, 2017 · We estimated the median divergence time between the Owens pupfish and Death Valley pupfishes between 15.1 and 39.5 kya across all ten data sets ...
[67]
The Pleistocene history of the sheepshead minnow (Cyprinodon ...
Reproductive isolation among endemic pupfishes (Cyprinodon) on San Salvador Island, Bahamas: Microsatellite evidence. 2008, Biological Journal of the Linnean ...<|control11|><|separator|>
[68]
Adaptive introgression from distant Caribbean islands contributed to ...
Aug 10, 2017 · The adaptive radiation of San Salvador Island pupfishes provides an outstanding system to compare the contributions of different sources of ...
[69]
[PDF] DFW 5-year Status Review - CA.gov
The Desert Pupfish (Cyprinodon macularius Baird and Girard) is currently listed as endangered under the California Endangered Species Act (CESA) (Fish and ...
[70]
Climate change puts endangered Devils Hole pupfish at risk of ...
It is an extreme environment, with water temperatures and dissolved oxygen concentrations near their lethal limits for most fish. The iridescent blue, one-inch- ...
[71]
A tiny desert fish hits a 25 year population high in one of Earth's ...
May 30, 2024 · The critically endangered Devils Hole pupfish population has reached a 25-year high of 191 fish, offering hope for the species that lives in ...
[72]
Blue Tilapia - Texas Invasives
Aug 30, 2021 · Blue tilapia are also known to impact catfish and a number of minnows and topminnows such as shiners, chub, pupfish, and mosquitofish -- ...Blue Tilapia · Description · Similar Species
[73]
Devils Hole pupfish - spring 2025 count - National Park Service
Apr 15, 2025 · The spring 2025 count of Devils Hole pupfish was 38, down from 191 the previous year, after a population decline due to earthquakes.
[74]
[PDF] Extinctions of North American Fishes During the past Century
Extinction of the Tecopa pupfish oc- curred in late 1970 or soon thereafter. Six pupfish collected by D. V. Hemphill on 28 March 1954 from "Tecopa Hot.
[75]
[PDF] Deformation of Endangered Status and Critical Habitat for the Desert ...
Summary: The Service determines the desert pupfish (Cyprinodon macularius) to be an endangered species. Critical habitat is also designɛted for this species in ...Missing: preferences | Show results with:preferences
[76]
Devils Hole Pupfish Population Takes A Dive - National Parks Traveler
Apr 16, 2025 · The number suddenly declined to 35 fish in 2013. Population numbers climbed in recent years and reached a 25-year-high during the spring of 2024 ...
[77]
Endangered desert pupfish RETURN TO longtime refuge
Apr 2, 2025 · During breeding season, males turn bright blue with yellow tails and aggressively defend their territory. Desert pupfish are evolutionarily ...
[78]
[PDF] COMANCHE SPRINGS PUPFISH RECOVERY PLAN - ECOS
Apr 20, 1981 · A remarkable species flock of pupfishes, genus Cyprinodon, from Yucatan, Mexico. Copeia,. 1981:52-64. Hubbs, C. 1957. Distributional patterns ...
[79]
Threatened Species Status With Section 4(d) Rule for Pecos Pupfish ...
Nov 22, 2024 · We propose to list the Pecos pupfish as a threatened species with a rule issued under section 4(d) of the Act (“4(d) rule”).