A second group, composed of rosy-faced shiner, blunt-faced shiner, mimic shiner, slender madtom, freckled madtom, spotted bass, and fan-tailed darter, normally is characteristic of clear tributaries rather than the mainstream of rivers. These species probably used the mainstream as a refugium during drought; with the resumption of flow, conditions became unsuitable for these populations in the mainstream. At the same time, conditions probably became favorable to the re-establishment of these species in tributaries. Metcalf (1959:396) listed the rosy-faced shiner, blunt-faced shiner and mimic shiner as species that were characteristic of upland tributaries in the Flint Hills and Chautauqua Hills of Chautauqua, Cowley and Elk counties in Kansas. The slender madtom and fan-tailed darter are more common in clear streams of southeast Kansas than in other areas of the state (Cross, personal communication and data of the State Biological Survey of Kansas). Both species are recorded by Hall (1952:57-58) only in upland tributaries on the east side of Grand (Neosho) River in the Fort Gibson Reservoir area of Oklahoma. Neither species was taken in faunal studies of the Verdigris River in Oklahoma (Wallen, 1958), in the Verdigris and Fall rivers in Kansas (Schelske, 1957), or by Metcalf (1959).
The spotted bass is not so restricted in its distribution and its habitat-requirements as are other species in this group; but, in Kansas, spotted bass are most abundant in clear creeks in the southeast part of the state.
The freckled madtom was taken in most of the studies cited above and is most common in the smaller streams of the southeast one-fourth of Kansas and the northeast one-fourth of Oklahoma. Schelske (1957:47) reports that the freckled madtom was taken only in March, April, October and November in the Verdigris River, Kansas. My only record of this species was obtained in the Neosho River in April, 1958.
The third group is composed of channel catfish, flathead, and freshwater drum. This group represents that element of the population that responded most quickly to the resumption of continuous flow. The fact that adult channel catfish and flatheads live in pools and do not require flowing water to spawn gives these species a survival advantage as well as a reproductive advantage over obligatory riffle fishes (such as most darters) in the highly variable conditions found in Kansas streams. These factors resulted in unusually high reproductive success in 1957. Subsequent survival of fry was excellent; however, some mortality in the highly-dominant 1957 year-class became apparent in the 1958 and 1959 collections, accounting for a numerical decline in these species. The ability to respond immediately to increased flow is an adaptive feature that allows these species to maintain high levels of abundance in the highly fluctuating streams of Kansas.
The continuous flow that occurred in 1957 in the Neosho and Marais des Cygnes rivers, for the first time in four years, provided the necessary habitat for survival of young catfish hatched in that year. The nearly complete absence of other species on the riffles, and the abundant populations of riffle-insects that I observed in the summer of 1957, were undoubtedly factors contributing to the survival of young.
The decrease in abundance of the red shiner may be partially due to an increase in the numbers of other species that are well adapted to conditions of permanent flow. At the completion of my study, the red shiner was still the most abundant minnow in both rivers. In 1957 this species was common in many habitats, including swift riffles, that were later occupied by madtoms, darters, the gravel chub, mountain minnow and sucker-mouthed minnow.
The basic pattern of change was clearly an increase in the species that are characteristic of permanently flowing waters, and a decrease in the species that are characteristic of ponds or small, clear streams.
CONCLUSIONS
The fauna of the Neosho and Marais des Cygnes rivers is capable of a wide range of adjustment in response to marked environmental changes. As these rivers become low and clear they take on many of the faunal characteristics of smaller tributaries and ponds. Species such as black bullhead, spotted bass, largemouth, white crappie, red shiner, rosy-faced shiner, blunt-faced minnow, mimic shiner, and slender madtom assume a more prominent position in the total population. Other species such as channel catfish, flathead, freshwater drum, blue sucker, and such riffle-dwelling species as the gravel chub, Neosho madtom, and slender-headed darter hold a less prominent position in the total population.