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.
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