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171 1 he Southwestent Naturadst <br />to several years after colonization, although long-term coexistent y ,i kA <br />occur. On a geographic scale, topminnows have declined only in the un>r <br />of overlap of these two species (Meffe et al., 1983). In northwestern Mexicn, <br />where G. affinis has only recently colonized a few localities, P. occidentattu <br />remains abundant (Hendrickson et al., 1981). <br />HYPOTHESES.-Although several topminnow populations have beret <br />extirpated by habitat destruction (Miller, 1961; Minckley and Deacon, ISN, <br />Minckley, 1973; McNatt, 1979; Meffe et al., 1983), many others have btrly <br />lost with no apparent environmental change other than introduction (? <br />mosquitofish. It is unlikely that subtle physical or chemical fluctuation, <br />contributed to decline of this species, as the fish can tolerate a wide ranr;r <br />of conditions (Schoenherr, 1974; Meffe et al., 1983). These observation, <br />suggest that interaction with G. affinis is a major factor in local extirpation <br />of P. occidentalis. Specific mechanisms of replacement are not known, <br />although several hypotheses were advanced (Miller, 1961; Minckley, 1975- <br />Schoenherr, 1974, 1981). The more plausible of these are summaiirrd <br />below, and I present experimental evidence for predation as,a major (an,r <br />of topminnow losses. <br />Parasites and Disease.-Introduced organisms may harbor parasites of <br />diseases that can decimate native species (Elton, 1958, and refer(-u(n <br />therein), and transfer of parasites to native fishes by introduced fish vector? <br />has been documented in some instances (Hoffman, 1970; Bauer anti <br />Hoffman, 1976; Deacon, 1979). In the present case no new parasites Of <br />diseases of P. occidentalis are evident and the fish appears healthy w•herr <br />sympatric with introduced species. Mpoame (1981) found only the ricalaurcle <br />Camallanus sp. infecting P. occidentalis, with no evidence that parasites to <br />introduced fishes affected any native Arizona species. Therefore, hypothvwl <br />suggesting decline of Sonoran topminnows through introduction c>) <br />parasites or diseases are not supported. <br />Hybridization.-There is no evidence that P. occidentalis and G. affirrtt <br />hybridize. Hybrids were not observed in the field or laboratory (Schoenherr. <br />1974; Meffe, pers. observ.), and conspecific mate choice fidelity was high in <br />laboratory observations (Schoenherr, 1981). <br />Competition.-Because topminnows and mosquitofish are morphology <br />tally similar (Minckley, 1973), and competition is generally greatest among <br />similar species (Darwin, 1859; Gause, 1934; Miller, 1967), one might predict <br />intense interspecific resource competition; however, none can be <br />demonstrated between P. occidentalis and G. affinis (Schoenherr, 1981; <br />Meffe et al., 1983). The mosquitofish is largely a piscivore/insectivoir <br />(Harrington and Harrington, 1961; Myers, 1965), while topminnows ford <br />primarily on detritus and vegetation, with insect larvae and amphip od <br />crustaceans important in the diet of some populations (Minckley, 1975; <br />Deacon and Minckley, 1974; Schoenherr, 1974; Constantz, 1976; Gerking <br />and Plantz, 1980). Food choice also seems little influenced by sympau) <br />(Schoenherr, 1981). Additionally, trophic morphologies are quite different. <br />Mosquitofish have strong, conical teeth, firmly attached to the jaw, and <br />gut length about 0.69 times standard length (SL), whereas Sonoran <br />.1", r1w 1985 srcuc-„ . ............... ._.,,..............-------- <br />vol. Sll <br />e ina 1 „ninnows have elongate, spatulate teeth, rnore weakly attached to the <br />7)L <br />ow, and a gut length 1.5 to 2.0 times SL (Schoenherr, 1981; Meffe et al., <br />i+g13). These trophic morphologies are characteristic of a carnivore and an <br />urbivore/detritivore, respectively (Al-Hussaini, 1949; Barrington, 1957; <br />lrgler et al., 1977). Furthermore, there is no evidence that food is limiting <br />n highly productive desert habitats (Schreiber and Minckley, 1981), and <br />ttowth rates also appear unaffected in sympatry (Schoenherr, 1981). <br />(irmpetition hypotheses therefore appear unsupported. <br />Predation.-Predation was suggested as a major factor in decline of <br />i.,noran topminnows (Minckley, 1973; Minckley et al., 1977; Schoenherr, <br />1481; Meffe et al., 1983), yet the hypothesis was not adequately tested. <br />r;antbusia affinis is a known piscivore and its introduction was correlated <br />.01 population reductions of more than 20 species of fishes (Schoenherr, <br />:981). It eats P. occidentalis in the field and laboratory (Minckley, 1973; <br />vhoenherr, 1981; Meffe et al., 1983) and is potentially capable of exerting <br />;sedation pressure on the species. I tested the hypothesis that predation on <br />pneniles causes local extinction of Sonoran topminnows. <br />Physiological Stress.-Schoenherr (1981), through observations of these <br />Iwo fishes in aquaria and an experimental pond, and with concordant <br />,observations by P. Winkler (in Schoenherr, 1981), concluded that <br />,nosquitofish physiologically stress topminnow adults by constant <br />Aggression. This resulted in reduced fecundity, cessation of feeding, and <br />increased mortality of P. occidentalis when sympatric with G. affinis. I <br />,ntcd Schoenherr's hypothesis through examination of those parameters in <br />i controlled laboratory experiment, and at a field site with natural <br />rxlxrimental and control groups. <br />%I,TEBtAIS AND METHODS.-Laboratory Coexistence Study.-I examined impacts of predation <br />1, G. affinis on population size of P. occidentalis under non-competitive conditions. The null <br />t,l.,thesis is that there is no difference in topminnow population growth rates, and hence <br />,owunent, when in allopatry or sympatry with mosquitofish. <br />smipatric and allopatric populations of each species were established in 75.0 and 37.5 1 <br />.imria, respectively. Six replicates of each were in simple habitats (sand substrate only) and 6 <br />n,,,mplex habitats (sand, rocks, and submergent vegetation 1Vesicularia sp., Hypnaceal). Eight <br />' -,, Ies and four males of either or both species were stocked on 16 July 1981, at a density of <br />';R fish%m' (natural densities vary widely from a few individuals to several hundred fish per m'). <br />I yuninnows were taken from a natural spring-seep in southern Arizona (Meffe et al., 1982) and <br />n ?uµnitofish were from an artificial pond at Arizona State University (ASU). <br />lisp were fed powdered trout chow twice daily, with frozen brine shrimp substituted three <br />.:ens weekly. Fish were fed to excess, thus eliminating food competition. Populations were <br />ennrd at regular intervals for a total of 63 weeks and (lead fish were removed and preserved <br />n 10% formalin. Adult fernales were dissected as they died, and mature ova and embryos <br />•?idit}) counted. <br />field coexistence study.-The above design was repeated in a natural habitat, a small spring <br />.n near Bylas, Arizona (habitat descriptions are in Meffe, 1983x, and Meffe and Marsh, 1983), <br />-1&00 ing the same null hypothesis. Open-bottomed cages of 15 mm-mesh hardware cloth lined <br />-ah nylon window screening were sunk into the substrate on 24 June 1981. Three allopatric <br />i :IN) r m') and three sympauic (2,400 em') covered cages were used. Seven female and three male <br />•w,minnows were stoked in the former, and seven males and three females of both species were <br />curd in the latter. All fish were collected in situ, and stocked at densities of I fish/120 an' <br />.alate area (water depth = 3-10 on). Only natural food was available. Individuals in cacti cage <br />-i, tounted 5, 13, and 24 days after stocking.