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.     

Survival and reproductive performance of the desert pupfish, Cyprinodon n. nevadensis (Eigenmann and Eigenmann), in acid waters

Desert pupfish, Cyprinodon n. nevadensis , were exposed to pH levels of 8·3 (control), 7·0, 6·5, 6·0, 5·5 and 5·0 to determine the effects of acidity on reproduction. Egg production was significantly reduced at every pH level tested below the control. Egg-laying virtually ceased at pH 5·0, while egg viability was reduced to less than 50% of the control value at pH 6·5, when eggs were tested at the same pH at which they were laid. The 96 h LC₅₀ for this species was pH 4·56, considerably below that for successful reproduction. Larvae were less tolerant to acid stress than were adults.
Acclimation of reproductive performance to increased acid levels did not occur. Reproductive performance did not fully recover to control levels when the fish were placed in more favourable conditions after prolonged exposure to low pH.
Reduction in the number and development of oocytes was observed in the ovaries of acid stressed fish resulting in the decreased reproductive potential.     

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.     

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.     

Sexual selection is stabilizing selection in pupfish [Cyprinodon pecosensis)

One of the predictions of the 'good genes' model of sexual selection is that reproductively successful males with well-developed indicator traits should show smaller variances for non-indicator traits, that are not directly associated with mating success, when compared to non-breeding males and females. Thus sexual selection should reinforce stabilizing natural selection in reducing the variance in quantitative traits. This prediction is tested by analysing variation in eight morphological traits of breeding males, non-breeding males, and females of pupfish (Cyprinodon pecosensis). Breeding males tended to be less variable than non-breeding males for all principal component factors, and for all morphological traits except for depth, although these differences were statistically significant only for PC2, and PC5 and for pelvic fin length, number of pelvic fin rays and number of preopercular and preorbital pores. Similarly, breeding males tended to be less variable than females for all principal component factors and for all morphological traits except for number of preopercular pores. These differences were statistically significant for PC2, and for depth, pelvic fin length, number of preorbital pores and pectoral fin rays. The overall pattern of reduced variability in independent traits of breeding males revealed by principal component analysis is very consistent and highly significant (P<10⁵). These results support the prediction of the 'good genes' model and show that reproductively active males are subject to more severe stabilizing selection for several quantitative traits than non-breeding males and females. Thus sexual selection, through male-male competition, female choice, or an interaction of both selective processes, results in stabilizing selection on quantitative morphological traits.     

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.     

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.     

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.     

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.     

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)]     

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.     

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.     

Metabolism of a desert stream

SUMMARY. Rates of photosynthesis and community respiration were determined for benthic assemblages in Sycamore Creek, a Sonoran Desert stream in Arizona. Benthos in this stream can be separated into (1) mats of Cladophora glomerata and associated epiphytes and (2) assemblages of epipelic diatoms and blue-green algae. Community respiration and net photosynthesis were measured for these assemblages using submerged light-dark chambers in situ . Multiple regression analysis was used to predict (1) gross photosynthesis as a function of photosynthetically active radiation, temperature and chlorophyll-α concentration; and (2) community respiration as a function of temperature and biomass.
Calculations suggest that Sycamore Creek is autotrophic during the summer ( P/R = 1.7) and that the rates of gross photosynthesis ( P =8.5 g O₂ m⁻² day⁻¹) and community respiration ( R = 5.1 g O₂ m⁻² day⁻¹) are high for a small stream. Considerable difference exists between the Cladophora mat assemblages, in which mean P is 12.5gO₂m⁻² day⁻¹and the P/R ratio is 2.3, and the epipelic assemblages in which mean P is 4.4 g O₂m⁻² day⁻¹ and P/R is 0.96. The high rate of gross photosynthesis, low litter inputs, high biomass of algae and the intermittent but severe floods that characterize Sycamore Creek indicate that this stream and other similar desert streams are net exporters of organic matter and are, thereby, truly autotrophic stream ecosystems.     

Saltcedar (Tamarix ramosissima) invasion alters organic matter dynamics in a desert stream

1. We investigated the impacts of saltcedar invasion on organic matter dynamics in a spring‐fed stream (Jackrabbit Spring) in the Mojave Desert of southern Nevada, U.S.A., by experimentally manipulating saltcedar abundance. 2. Saltcedar heavily shaded Jackrabbit Spring and shifted the dominant organic matter inputs from autochthonous production that was available throughout the year to allochthonous saltcedar leaf litter that was strongly pulsed in the autumn. Specifically, reaches dominated by saltcedar had allochthonous litter inputs of 299 g ash free dry mass (AFDM) m^?2 year^?1, macrophyte production of 15 g AFDM m^?2 year^?1 and algal production of 400 g AFDM m^?2 year^?1, while reaches dominated by native riparian vegetation or where saltcedar had been experimentally removed had allochthonous litter inputs of 7–34 g AFDM m^?2 year^?1, macrophyte production of 118–425 g AFDM m^?2 year^?1 and algal production of 640–900 g AFDM m^?2 year^?1. 3. A leaf litter breakdown study indicated that saltcedar also altered decomposition in Jackrabbit Spring, mainly through its influence on litter quality rather than by altering the environment for decomposition. Decomposition rates for saltcedar were lower than for ash (Fraxinus velutina), the dominant native allochthonous litter type, but faster than for bulrush (Scirpus americanus), the dominant macrophyte in this system.     

Nitrogen fixation in a desert stream ecosystem

Few measurements of nitrogen fixation exist for streams. Desertstreams are warm, well lighted, and often supportabundant cyanobacterial populations; thus N₂ fixationmay be significant in these N-poor ecosystems. N₂fixation was measured in situ by acetylene reductionfor two patch types (Anabaena mat and anepilithic assemblage). Patch-specific rates were highcompared with published values (maximum 775 µgN₂ [83 µmol C₂H₄]mg chl a ^-1 h^-1or 51 mg N₂ [5.4 mmol C₂H₄] m_-2 h_-1).Daytime fixation was higher than nighttimefixation, and temperature, light and inorganic Nconcentration explained 52% of variance in hourlyrates over all dates. Diel input-output budgets wereconstructed on five dates when cyanobacteria werepresent in the stream. Diel N₂ fixation rates weremeasured for comparison with reach-scale diel nitrogenretention, to assess the importance of this vector to Neconomy of the stream. Fixation accounted for up to85% of net N flux to the benthos, but its importancevaried seasonally. Finally, we applied biomass-specificfixation rates to 1992 and 1993 biomass data to obtainseasonal and annual N₂ fixation estimates.Cyanobacteria were absent or rare during winter andspring, thus most of the annual N₂ fixation occurredduring summer and autumn. Annual rates of nitrogenfixation for 1992 and 1993 (8.0 g/m₂ and 12.5g/m₂) were very high compared to other streams,and moderately high compared to other ecosystems.Like other phenomena in this disturbance-proneecosystem, nitrogen fixation is strongly influenced bythe number and temporal distribution of flood events.     

Genic diversity in Red River pupfish Cyprinodon rubrofluviatilis (Atheriniformes: Cyprinodontidae) and its implications for the conservation genetics of the species

Starch gel electrophoresis of proteins was used to study geographic variation at 26 gene loci in the Red River pupfish ( Cyprinodon rubrofluviatílís ), a species restricted to west Texas and Oklahoma. Marked differences were detected between populations in the Red and Brazos river drainages, with fixed or nearly fixed differences occurring at five gene loci. In addition, mean heterozygosity was uniformly high for the Red River form ( = 0·076–0·101) while samples of the Brazos River form were genetically depauperate ( =0·00–0·017). Introduced populations in the South Canadian and Colorado river drainages appear to have been derived from the Red River drainage. The presence of alleles diagnostic of the Red and Brazos river forms supports the suggestion from previous work that they may represent cryptic species. Regardless of taxonomy, however, the presence of two genetically distinct forms must be taken into consideration by those concerned with maintenance of biotic diversity.