The American Midland Naturalist Published Quarterly by the University of Notre Dame, Notre Dame, Indiana
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The American Midland Naturalist Published Quarterly by The University of Notre Dame, Notre Dame, Indiana Vol. 109 APRIL, 1983 No. 2 Multivariate Assessment of Environmental Preferences of Cyprinid Fishes of the Illinois River, Oklahoma JAMES D. FELLEY1 and LOREN G. HILL Oklahoma Biological Survey and Department of Zoology, University of Oklahoma, Norman 73019 ABSTRACT: Cyprinid fishes were collected from the Oklahoma drainage of the Il- linois River in all seasons of 1978 and 1979. The Illinois River and its tributaries drain the Ozark Plateau and the Boston Mountains and support an Ozarkian fish fauna. Species included in this analysis were Campostoma anomalum, Nocomis asper, Notropis boops, N. nubilus, N. pilsblyi, N. rubellus, N. whipplei, Pimephales notatus, Phoxinus erythrogaster and Semotilus atromaculatus. Fishes were collected and 17 environmental variables measured from each of 15 localities sampled in every season. Species distribu- tions were characterized in reference to each variable as follows: the value of a variable at each location was weighted by the number of individuals of a species present, and the mean of these weighted values was taken as that species' preferred state for that variable. Principal components analysis, with rotation of the component solution to simple structure, was used to elucidate the relationships among variables in terms of species distributions. In most seasons, components related to species preferences for upstream vs. downstream locations, different foods, and slow- vs. fast-water habitats. For most species, differences in habitat preference between seasons were related to breeding (in the spring) and to preference for warmer locations during the rest of the year. INTRODUCTION The distribution of a species is constrained by its environmental tolerances and by interactions, such as competition and predation, with other species (Smith and Powell, 1971). Within these limits, distribution is related to preferences for given environmen- tal condition. The relationship between environmental conditions and distribution has been shown for cyprinids in particular (e.g., Matthews and Hill, 1979 a and b; Mat- thews and Maness, 1979). Different tolerances and preferences of separate species may result in differing patterns of distribution, and seasonal or daily changes in en- vironmental conditions may lead to movement of a species. One way to characterize the distribution of a species relative to an environmental variable is by the mean of the variable over all collected individuals (Felley, 1980). This mean is the state of that variable found at locations where individuals are most likely to occur, and can be used as an estimate of a species' preference for the variable. Though species distributions may be so characterized for any number of variables, not all of these will be of equal importance in affecting a species' choice of habitat. A given environmental variable may directly influence a species' habitat choice. Alter- natively, a variable may have no effect on choice of habitat but may be functionally related to a variable that does have an effect (for example, if a species' occurrence is af- Present address: Department of Biology, McNeese State University, Lake Charles, Loui- siana 70609. 209 210 THE AMERICAN MIDLAND NATURALIST 109(2) fected by current speed, variables such as substrate type or amount of debris present would be correlated with it). Finally, a variable may be unrelated to habitat choice. In- vestigation of species distribution according to different environmental variables must take into account the possible confounding relationships among the variables themselves. Baker and Ross (1981) investigated cyprinid distributions in reference to different environmental variables and related distributional patterns to habitat segregation among the species, along a number of ecological axes. They showed that analysis of species preferences (as suggested above) and identification of patterns of covariance among the variables can point to trends in habitat partitioning among the species. In this case, covarying variables are those reflecting a common trend in species distribu- tions. Multivariate procedures (such as multivariate analysis of variance and com- ponents analysis) are uniquely suited to identification of sets of covarying variables. We used components analysis to investigate distribution of members of the family Cyprinidae in different seasons in the Oklahoma drainage of the Illinois River. MATERIALS AND METHODS The Illinois River drains portions of the Boston Mountains and the Ozark Plateau in Arkansas and Oklahoma. In Oklahoma, the Illinois River drains a limestone/sand- stone topography, and headwaters are heavily influenced by springs. Figure 1 il- lustrates the Oklahoma drainage of the Illinois River and our collection localities. The fish fauna of the Illinois River is typical of Ozark streams; cyprinid species regularly collected and included in our analysis were as follows: Campostoma anomalum stoneroller; Nocomis asper redspot chub; Notropis boops bigeye shiner; N. nubilus Ozark minnow; N. pilsbgi duskystripe shiner; N. rubellus rosyface shiner; N. whipplei steelcolor shiner; Pimephales notatus bluntnose minnow; Plwxinus erythrogaster southern redbelly dace, and Semotilus atromaculatus creek chub. All of these species were collected in at least seven of our eight collecting seasons. Fish were collected at 15 localities in the Illinois drainage in spring, summer, autumn and winter of 1978 and 1979. In every season (calendar-defined, e.g., "spring" was 21 March to 21 June) we collected fish once at each locality. Collections were ob- tained by seine (3.7 x 1.5 m, 3 mm mesh) over small discrete areas (never more than 40 m2, often smaller) representing microhabitats. A microhabitat was an area determined in the field to be homogeneous for the following parameters: water clarity, substrate type, current speed, presence or absence of cover (structure or vegetation in which fish might hide), presence or absence of debris (leaves and sticks on the bottom), and presence or absence of emergent vegetation. Thus we attempted to minimize the possibility of seining through environments recognized as different by the fishes. So defined, a microhabitat was our sampling unit, and only one microhabitat was sampled at a location in a season. Water clarity was measured as Secchi disc depth, in meters. Water clear to the bottom was arbitrarily coded as 2 m. Substrate type was coded from 0-5 representing mud, sand, gravel, rubble, boulders, bedrock, respectively. Current speed was measured as seconds required for an object to float 10 m. Still water was ar- bitrarily assigned a value of 180 sec. Cover, debris and vegetation were each coded 1/0 for presence/absence. Additional variables measured at each location were stream width (m), maximum stream depth at the location sampled (m), depth of the stream at the sampled microhabitat (termed capture depth, measured in m), conductivity, pH and temperature. Fishes were preserved in 10% formalin. Up to 10 individuals of each species col- lected at a location were examined for gut contents. Gut contents were divided into four categories (terrestrial invertebrates, aquatic invertebrates, filamentous algae and detritus; Minckley, 1963) and percentages of volume of these categories were estimated 1983 FELLEY & HILL: FISH ENVIRONMENTAL PREFERENCES 211 for each individual. Stream order (which ranged from 2-4 at our collection localities) was determined from 1/250,000 topographic maps. Treatment of variables. —The values of environmental variables at each location were weighted by the number of individuals of a species collected there. For example, if 15 individuals of a species occurred at one location, and a single individual at a second, the values of the environmental variables at the first location would be weighted 15 times more heavily than those of the second location. For data coded 1/0 as presence/absence (debris, cover, vegetation) the result of this weighting procedure gave the percentage of individuals of a species that were collected at locations where that variable was coded "1." Temperature was averaged in the following manner for each species in a season: The average temperature for all locations was computed for that season and this value was subtracted from the temperature mean of each species col- lected in that season. So expressed, negative temperature averages indicated a preference for temperatures lower than the drainage mean; positive values indicated preference for temperatures higher than the average. Because pH is a power function (negative logarithm of hydrogen ion concentration), geometric means of pH values Fig. 1. —Locations sampled for cyprinid fishes in the drainage of the Illinois River, Oklahoma 212 THE AMERICAN MIDLAND NATURALIST 109(2) were calculated for all species (Sokal and Rohlf, 1969). Means for food type percent- ages were calculated for each species. The categories of terrestrial and aquatic prey were combined into the variable "prey" for the analyses described below. This differen- tiated among cyprinids eating animal prey, filamentous algae or detritus. Statistical analyses were performed on the means of the variables for all species. Statistical analyses. —Statistical analyses were done using the Statistical Analysis System (SAS; Barr et al., 1976) on a 370 IBM computer. Variable means were cor- related across species in each season. The resulting correlation matrices were then sub- jected