Reproductive performance of the desert pupfish (Cyprinodon n. nevadensis) in relation to salinity

Synopsis The reproductive performance of the desert pupfish, Cyprinodon n. nevadensis, was tested by exposing breeding pairs to a range of salinities from 0.1 to over 40 In terms of eggs g^–1 body weight day^–1 , eggs per spawning and embryo viability. reproductive performance was optimal at 10 Performance decreased above and below 11 producing a plateau of statistically equivalent reproductive performance from 0.2 to 20 The upper salinity limit placed on reproduction is the tolerance of the eggs, since a few eggs are laid above 20but they do not hatch. These small fish probably never experience the upper salinity limit for reproduction in their native habitat. The tolerance limits for reproduction are compared with a lethal range from less than 0.1 to 53in a 96 h LD-50 test.     

Productivity of a herbivorous pupfish population (Cyprinodon nevadensis) in a warm desert stream

Production and food intake by an herbivorous pupfish population ( Cyprinodon nevadensis amargosae ) living in the outflow of a thermal artesian well (Tecopa Bore) near Death Valley, California is described. Water issues from the ground at 47.5° C and cools 8.12° C before leaving the study area 300 m from the source. High stream temperatures restricted the pupfish population to some 41 % of the study area, with a resulting mean density of 89 individuals m⁻² (range = 13 to 196m⁻²). Biomass estimates ranged from 7 kcal m⁻² to 42 kcal m⁻². The mean annual standing crop of pupfishes (24 kcal m⁻²) turned over about five times annually. Growth rates were highest in juveniles (♂= 9.5% day⁻¹) and slowest in large adults (♀= 08% day⁻¹). Monthly production ranged from 22 kcal m⁻² in September to 3 kcal m⁻² in July and August. Pupfish in Tecopa Bore fed on algae and detritus, ingesting 1941 kcal m⁻² yr⁻¹ or 17.5% of the annual net primary production. 119 kcal m⁻² yr⁻¹ was deposited in growth. This latter value is approximately ten times greater than values previously reported for large carnivorous fishes but is comparable lo values reported for herbivorous fishes under pond culture.     

Exogenous vasotocin alters aggression during agonistic exchanges in male Amargosa River pupfish (Cyprinodon nevadensis amargosae)

Pupfishes in the Death Valley region have rapidly differentiated in social behaviors since their isolation in a series of desert streams, springs, and marshes less than 20,000 years ago. These habitats can show dramatic fluctuations in ecological conditions, and pupfish must cope with the changes by plastic physiological and behavioral responses. Recently, we showed differences among some Death Valley populations in brain expression of arginine vasotocin (AVT). As AVT regulates both hydromineral balance and social behaviors in other taxa, these population differences may indicate adaptive changes in osmoregulatory and/or behavioral processes. To test whether AVT is relevant for behavioral shifts in these fish, here we examined how manipulations to the AVT system affect agonistic and reproductive behaviors in Amargosa River pupfish (Cyprinodon nevadensis amargosae). We administered exogenous AVT (0.1, 1, and 10 microg/g body weight) and an AVP V1 receptor antagonist (Manning compound, 2.5 microg/g body weight) intraperitoneally to males in mixed-sex groups in the laboratory. We found that AVT reduced the initiation of aggressive social interactions with other pupfish but had no effect on courtship. The effects of AVT were confirmed in males in the wild where AVT (1 microg/g body weight) reduced the aggressive initiation of social interactions and decreased aggressive responses to the behavior of other males. Combined, these results show that AVT can modulate agonistic behaviors in male pupfish and support the idea that variation in AVT activity may underlie differences in aggression among Death Valley populations.     

Predation by the desert pupfish,Cyprinodon macularius onCulex mosquitoes and benthic chironomid midges

Predation ofCulex mosquitoes and chironomid midges was demonstrated byCyprinodon macularius Baird & Girard in two types of shallow ponds. In large ponds (5.5×2.6 m) with fish, all mosquito breeding ceased 4 weeks after introduction ofC. macularius. A significant, density-dependent reduction of several chironomid biotypes was sustained, although an increase was evident for other chironomid biotypes. Fish biomass increased 2.43 fold in 76 days in m² ponds and 23.22 fold in 70 days in larger ponds. Behavioral studies showedC. macularius capable of feeding effectively on mosquitoes in floating algal mats. The desert pupfish deserves serious attention in the biological-integrated control of medically important aquatic arthropods, and may possess attributes making it a preferred, ecologically adapted substitute forGambusia affinis (Baird & Girard) in certain habitats.     

Distribution,habitat and conservation status of desert pupfish (Cyprinodon macularius) in the Lower Colorado River Basin,Mexico

In order to better determine the currentstatus of desert pupfish populations (Cyprinodon macularius macularius) in the LowerColorado River Basin of Mexico, bimonthly fishsampling and habitat evaluations were carriedout from September 1996 to August 1997 in sixlocalities of Baja California and Sonora.Desert pupfish were collected using minnowseines and traps. Four sampling sites are inSonora, in shallow marginal habitats of theCiénega Santa Clara (Hunters' Camp, Outletof the Welton-Mohawk channel, El Doctor andFlor del Desierto) and two are in BajaCalifornia (Cerro Prieto and streams south ofCerro Prieto). The most abundant fish speciessampled was native desert pupfish (C.macularius, 59%) followed by sailfin molly(Poecilia latipinna, 19%), redbellytilapia (Tilapia cf. zilli, 10%) and western mosquito fish (Gambusiaaffinis, 7%). Significant temporalfluctuations in distribution and abundance ofdesert pupfish populations, as previouslyreported for these populations, was againdocumented. The main anthropogenic factorsaffecting distribution and abundance of desertpupfish populations in the Mexican portion ofthe Lower Colorado River Basin are progressivealteration of aquatic and riparian habitats, aswell as presence of exotic fishes that havecaused the displacement or elimination ofnative fish populations. Both habitat andpresence of the desert pupfish populations inthe study area are controlled by theperiodicity, quality and volume of dischargesinto the Mexican portion of the basin from theUnited States.     

Condition status of the endangered desert pupfish,Cyprinodon macularius Baird and Girard, 1853, in the Lower Colorado River Basin (Mexico)

The length‐weight relationship (LWR) was used to test differences in the fish somatic condition factor among four populations of the endangered desert pupfish, Cyprinodon macularius, from the Lower Colorado River Basin, Mexico. Bimonthly fish sampling and habitat assessments were carried out from September 1996 to August 1997 for four sites in Baja California and Sonora. Slope b of the WLR varied from 3.238 (Cerro Prieto population) to 3.613 (Welton‐Mohawk population), showing in all populations a positive allometric growth. Male b values were higher than those of females for Cerro Prieto (3.303 vs 3.071), Welton‐Mohawk (3.866 vs 3.579), and Flor del Desierto (3.357 vs 3.169) populations. The Cerro Prieto population showed the highest somatic condition (y‐intercept, a = ?11.759). Water depth and salinity were the ecological variables that better accounted for most of the variation in the somatic condition of this pupfish, with a negative and a positive effect, respectively.     

Interrelationships of dominance and territorial behaviour in the pupfish,Cyprinodon variegatus

Under laboratory conditions, Cyprinodon variegatus establishes either a dominance or territorial mating system. In the dominance system, one male usually has complete access to receptive females, while in the territorial system, several males have equal access. Four factors were observed to enhance a subdominant's ability to establish a territory and improve his reproductive success; increasing the number of conspecific intruders, increasing the available area, the presence of partial barrier, and the presence of a receptive female.     

The conservation genetics of Ash Meadows pupfish populations. I. The Warm Springs pupfish <Emphasis Type="Italic">Cyprinodon nevadensis pectoralis</Emphasis>

The Warm Springs pupfish (Cyprinodon nevadensis pectoralis) inhabits several low flow (<0.15 cfs) springs in an otherwise dry Mojave Desert landscape. Increasing demands on groundwater coupled with predictions of decreased precipitation from global warming make this species a sentinel for biodiversity dependent on springs. Here we examine mitochondrial DNA haplotypes and nuclear microsatellite genotypes for individuals sampled in 1998 and 2007 as a means of inferring the historical demography of the subspecies. Estimates of genetic effective population size from comparison of allele frequencies over time underscores that all spring populations support small populations, typically <100 individuals. Such small population sizes suggest spring populations may have heightened probabilities of extirpation. Despite small population sizes, all springs harbor relatively high levels of genetic variation for both nuclear and mitochondrial DNA. Unexpectedly high levels of variation may be explained by recent declines in population size coupled with rare episodes of gene flow between springs suspected of occurring during flooding events. The inferences gained from the genetic data provide the basis for evaluating current and future restoration plans. The genetic data suggest there is a need to balance the predicted positive effects of restoring hydrological connectivity with the potential negative impacts of providing avenues for the spread of exotic species detrimental to pupfish populations.     

Sexual selection, stabilizing selection and fluctuating asymmetry in two populations of pupfish (Cyprinodon pecosensis)

Fluctuating asymmetry of morphological traits is thought to reflect the capacity of a genotype to produce an integrated, functional phenotype. I tested three predictions. (1) In a polygynous breeding system, under intense sexual selection on males, breeding males should show greater symmetry in bilaterally symmetrical traits than non-breeding males or females. (2) If these traits are under stabilizing selection, highly symmetrical individuals also should be modal phenotypes, thus near the mean value for that trait, whereas individuals with increased asymmetry should represent marginal phenotypes, near the extremes of the distribution for that trait. (3) Differences in the intensity of sexual selection should be reflected in differences in the degree of fluctuating asymmetry between sexes among populations. I examined the relationship between male breeding status and the degree of fluctuating asymmetry of four bilaterally symmetrical- traits, preorbital and preopercular pores and pectoral and pelvic fin rays, in two populations of Pecos pupfish which differed in the intensity of sexual selection. These traits do not function in male-male competition or female choice, thus are not directly affected by sexual selection. In Mirror Lake breeding males, as a group, were most symmetrical for all four traits, while non-breeding males and females showed higher levels of fluctuating asymmetry. Similarly, symmetrical individuals also represented modal phenotypes for four traits (breeding males), and for three traits (non-breeding males and females). These patterns were not seen in the Lake Francis population, where breeding males were as asymmetrical as non-breeding males and females, and the degree of fluctuating symmetry did not differ between modal and marginal phenotypes for any of the four traits. When ecological conditions favour intense sexual selection, either through female choice, male-male competition, or both, breeding males represent the most fit phenotypes. Thus sexual selection reinforces the effects of stabilizing selection on characters that do not function as secondary sexual traits. However, when sexual selection is relaxed, differences between sexes disappear.     

Characterization of microsatellite markers in a threatened species,the White Sands pupfish (Cyprinodon tularosa)

We report the characterization of nine new microsatellite markers for a threatened species, the White Sands pupfish (Cyprinodon tularosa), using an enriched library method. These markers show moderate levels of variation (two to five alleles per locus) in the two native populations of this species and reveal substantial divergence between these two populations as indicated by a high percentage of private alleles. These markers will prove very useful in the conservation management of this rare species.     

Thermoregulatory activity in the Tecopa pupfish,Cyprindon nevadensis amargosae,an inhabitant of a thermal spring

Synopsis The final thermal preferendum of Tecopa pupfish Cyprinodon nevadensis amargosae collected from a thermal spring was determined in the laboratory in spatial and temporal thermal gradients. Temperatures chosen centred around 30°C — a value close to those reported for other warmwater species in the literature. Upper limits to final preferenda and thermal tolerance in all species of fish are proposed.     

Sexual selection promotes hybridization between Pecos pupfish, Cyprinodon pecosensis and sheepshead minnow, C. variegatus

Rapid and extensive genetic introgression has occurred between Pecos pupfish (Cyprinodon pecosensis) and sheepshead minnow (Cyprinodon variegatus) in the wild. We studied both female mate choice and male-male competition for mates among C. pecosensis, C. variegatus, and their F1 hybrids to determine what role these behaviours played in the formation of the hybrid swarm. Female C. pecosensis preferred male C. variegatus to conspecific males, C. variegatus females displayed no significant preference when given a choice between purebred males, and neither C. pecosensis nor C. variegatus females discriminated against F1 hybrid males. We found no evidence for female olfactory recognition of mates. Male F1 hybrids and C. variegatus were more aggressive than C. pecosensis males, achieving greater reproductive success under two different experimentally-induced mating systems. Hybrids were superior to C. variegatus when only two males competed (dominance interactions), but the two types were competitively equivalent in a territorial mating system. Our results indicate that active inter- and intra-sexual selection contributed to the accelerated hybridization between these two species. By including the possibility that some aspects of a hybridization and introgression event may be under positive selection, researchers may better understand the dynamics that lead to hybrid zone stability or the spread of introgressed genetic material.     

Morphology and gonad development of normal soldiers and reproductive soldiers of the termite Zootermopsis nevadensis nevadensis (Isoptera, Archotermopsidae)

Reproductive or neotenic soldiers of the Archotermopsid Zootermopsis nevadensisnevadensis (Hagen) are compared to sterile soldiers and primary male reproductives. Several head capsule morphometrics correlate significantly with gonad size across all forms and both sexes of soldiers. The easily observed field character of ratio of mandible length to labrum length is a consistent and reliable feature of head capsule external morphology for predicting gonad development and reproductive potential of soldier forms regardless of age, sex, or live weight.     

Effects of sex-ratio manipulation on territoriality and spawning success of male pupfish,Cyprinodon pecosensis

The sex ratio of Cyprinodon pecosensis was manipulated by varying the number of females but not of males. This resulted in some subordinate males changing their breeding tactics from satellites to territorials. The spawning success of breeding males was directly related, and the intensity of male-male competition was inversely related, to the number of available females. Time-budget data indicated a tradeoff between courtship and agonistic behaviour. In the male-biased treatment, males engaged in a greater number of agonistic interactions and fewer courtship bouts; conversely, in the female-biased treatment courtship behaviour was more frequent than aggression. A greater proportion of males adopted the satellite breeding tactic in the male-biased and even-sex-ratio treatments. Smaller, competitively inferior males were most sensitive to changes in the sex ratio. In the female-biased treatment, with an increase in mating opportunities and a decrease in the level of male-male competition for females, average mating success increased, more males were able to defend territories, and some satellite individuals were able to adopt the territorial breeding tactic. Temporal or spatial changes in the operational sex ratio of a population may have profound effects on the dynamics of the breeding system by affecting (1) the proportion of males adopting primary and secondary breeding tactics, (2) variance in reproductive success, and (3) intensity of sexual selection.     

Role of male-male competition and female choice in the development of breeding coloration in pupfish (Cyprinodon pecosensis)

The function of male breeding coloration and the conditionsunder which it is expressed were investigated using two experimentalprotocols that differed in the degree of control of social factors.In tank experiments, focal males were separated by opaque orclear partitions or allowed to interact freely without partitions.In lake experiments, males were introduced into opaque or clearenclosures in the lake from which they were collected. Thisdesign tested the relative importance of breeding substrate,social interactions with other males, and presence of femalesand juveniles on the expression of breeding coloration and agonisticand mating behavior in pupfish. Breeding coloration was notexpressed in the absence of conspecifics, even in the presenceof suitable breeding substrate, or in the presence of juveniles.Breeding coloration was poorly developed when only males werepresent, was more intense in the presence of females, and darkenedsignificantly after a spawning bout Males in clear enclosuresin their natural lake habitat were in visual contact with conspecificsof both sexes and developed more intense blue breeding colorationthan males in the tank experiments that were in visual contactonly with other males. Uncon-fined males with territories inthe lake developed the most intense breeding coloration. Breedingcoloration of male pupfish is a facultative trait that is welldeveloped when breeding females are present and plays a majorrole in the attraction and courtship of mates. However, maximumdevelopment of breeding coloration by territorial males suggeststhat intrasexual and intersexual selection act concordantlyon this trait.[Behav Ecol 7: 431-437 (1996)]     

Evaluation of the role of refugia in conservation efforts for the Devils Hole pupfish,Cyprinodon diabolis Wales

The Hoover Dam Refugium was built to help insure the survival of the Devils Hole pupfish during an especially critical period. It served that purpose well. The Amargosa Pupfish Station added an additional, nearly identical artificial habitat to use in conservation and management of the Devils Hole pupfish. Contemporary concerns of conservation biology suggest that: 1) Genetic diversity or homogeneity of refugium populations should, when possible, be established by biochemical and morphological techniques; 2) protection of the natural habitat has been effected, and population levels have returned essentially to normal, therefore, management intervention in the natural population is not now appropriate; 3) refugium populations represent a unique opportunity to develop information useful in conservation and management of small populations of endangered fishes, if legal restrictions to such investigations can be accommodated; and 4) refugia, dependent on human activities for construction and maintenance, are less reliable than natural habitats.