W. I Minckley

Man and The Waters of the Upper Ohio Basin

A Symposium held at Linesville, Pennsylvania, under the joint auspices of the Pymatuning Laboratory of Field Biology, Department of Biological Sciences, The College, and the Department of Public Health Practice, The Grad- uate School of Public Health, University of Pittsburgh.

Special Publication Number 1, Pymatuning Laboratory of Field Biology FEB 1956 THE CHANGING FISH FAUNA OF THE UPPER OHIO BASIN

Ernest A. Lachner U. S. National Museum

Our fish heritage in the upper Ohio Basin was extremely rich. Dense schools of fishes existed in these waters. Early explorers and pioneers traversing the Alleghenies found the streams teeming with life and these provided them with a ready source of excellent food.

The work of man has considerably altered this once abundant food source. Many crystal-clear, constant-flowing streams with lush growths of aquatic plant life, clean beds of gravel, rock and sand have been laid waste. We have plundered the great forests, drained and ditched lowlands, dredged stream beds, sprayed forests and fields, and intensified agriculture to the extent that storms now erode quantities of soil into our streams. Streams became common disposal pipes for municipal sewage, industrial and agricultural wastes of all types, such as mill acids, mine drainage, cannery and dairy wastes. Many of the numerous springs which fed a constant supply of water to our streams have been reduced to a trickle. The groundwater level has been lowered considerably. Many streams are seasonally dry, likened to desert desolation. The large rivers show great fluctuations in water flow. Once clear streams carry a large load of silt today. Water tempera- tures have generally risen.

The general consequence is that the streams are no longer suitable for the fish life that once inhabited them. Only the tolerant and hardy species survive generally. In many areas of the Basin the fish fauna has been completely elimin- ated. The huge migratory runs characteristic of several species in former days cannot be found today. The density of our fish population has been generally re- duced to an ineffective resource. Some species face extinction. The angling pressure has risen considerably and in the immediate future we forecast even a larger army of fishermen. There are probably more than 3 million anglers of all ages in this Basin each year contributing to an aggregate business enterprise totalling millioqs of dollars in gear, tackle, food, clothing, lodging, and L transportation.

Character and Composition of the Fauna of the Basin

The fauna of the upper Ohio Basin, restricted herein to include waters of the Ohio River above the mouth of the Kanawha River, West Virginia, is comparatively rich in species. More than 125 species of fishes belonging to 25 families occur here. The representatives of several families in these waters are found nowhere else outside of the North American faunal area, as the gar (Lepisosteidae), bowfin (Amiidae), mooneye (Hiodontidae), North American freshwater catfish (Ictaluridae),

'Charles A. Dambach, "Multiple Use of Water Reservoirs," paper presented 17th Ann. Conf. Ohio Sec. Am. Waterworks Assoc., Columbus, Ohio. Excerpt from: "Fishing in the Drink," Bull. Sport Fishing Inst., 48:1-2, 1955.

64

63 Indications are that a sound land use program can do much to increase infil- tration of water i14o the soil and reduce e amount of run-off and erosion. We cannot propose tl1t the entire Upper 0 io Basin be reforested. Many acres must be kept in cult ation, pastures an other essential uses. Fortunately, those land surfaces t t should be in ag icultural use are much less critical than the steep broken a eas.

The first step in a soùd land util zation program is land classification. With intensive soil survey as a basic, the land that has been determined to be unfit for agriculture shoul be used or growing trees for timber, recreation, and water supply protection p poses The land that is clearly suited for per- manent agriculture should be dòyeloed as highly as possible through the adoption of agricultural practices which i keep erosion at a minimum and per-acre crop yields as high as economically fe ible.

Ad itio al References

The Nation Looks at its Res rces. ction III - Water Resource Problems, Report of the Mid-Century Conf rence on sources for the Future, 1953, Resources for the Future, Inc., Washing on, D.C.

Hoyt, William G., and La helm, Walter B., Floods, Princeton University Press, 1955.

Carhart, Arthur H., er in Your Life, J. B. ippincott Co., 1955.

J. FredericA and Associates, America's eds and Resources, The Twentieth Century Fund, ,N ew York, 1955. 65 sunfish (Centrarchidae) and troutperch (Percopsidae). The Basin fauna contains forms of several archaic groups as the gar and paddlefish (Polyodontidae) families. The gars, "living fossils," are as old geologically as the famed African coela- canths and rival them in biological intrigue. Only a few species are found in North American waters. The paddlefish, with a long, paddle-like snout, also represents a primitive family known only by one species in the Mississippi River fauna of North America and with one giant relative in the large rivers of China.

Unlike the higher groups of terrestrial vertebrates, most of the North American freshwater fish families are not represented in the South American fauna. The several hundred species of North American freshwater fishes in the (), perch (Percidae), sucker (Catostomidae) and the freshwater catfish (Ictaluridae) families have no immediate relatives in the rivers of South America, the number decreasing progressively in the Mexican fauna. In contrast, the vast South American freshwater fauna totalling several thousand species is most closely related to those of Africa. The several families of characins of South America, probably numbering about 1000 species, and the cichlids, also decrease in numbers in the Mexican fauna, and each is represented in southern Texas by a single species. The South American freshwater catfish group comprising about eight families and more than 1000 species is not found in waters draining the United States.

It is of great zoogeographic interest that our fauna shows closest resemblance to that of Eurasia. Such families as the sturgeon (Acipenseridae), minnow, pike (Esocidae), mudminnow (Umbridae) and perch have representatives in both North America and Eurasian continental areas. Our sucker family is almost entirely North American, with only two species known so far in Asiatic waters. The sculpin family (Cottidae) is typically a marine group with many species in the cooler waters of the North Pacific, some in the Arctic, and three species in the colder waters of the southern hemisphere. We have the brook silverside, a fresh- water representative of the silverside family (Atherinidae), a group commonly represented in the temperate and tropical marine waters of the globe. The primitive lamprey family (Petromyzontidae) is present in the Basin represented by parasitic and non-parasitic species. These inhabit only fresh water and are related to the parasitic sea lamprey in the Great Lakes, a land-locked marine form which today is causing tremendous havoc to the commercial fisheries.

Our daces, chubs, shiners and are all members of the minnow family which is one of the largest of the world, containing more than 1,000 species in North American, Eurasian and African waters. It reaches its greatest abundance in species in southeastern Asia. This family, with the suckers and catfishes, forms a fascinating group for study because their distribution provides con- siderable evidence of the past history of the continents. For further discussion of this topic see Norman (1948), Myers (1938) and Gosline (1944) and numerous references listed therein. 1

1 J. R. Norman, A History of Fishes, Frederick A. Stokes Co., N.Y., 1948.

George S. Myers, "Fresh-water Fishes and West Indian Zoogeography," Smithsonian Rept. (1937): 339-364.

William A. Gosline, "The Problem of the Derivation of the South American and African Fresh-water Fish Faunas," An. da Acad. Brasileira de Ciencias., 16(3): 211-223, 1944. 66 Origin of the Fauna

Questions often presented to the ichthyologist are: Where did our fishes come from? How long have they been in the Basin? Do they really have relatives else- where on the earth? There has been considerable speculation about the origin of our freshwater fauna. Several chapters would be required for an adequate discus- sion of this subject. Let it suffice to say that long before the last great ice masses existed in northern United States, probably more than 100 million years ago, North America and Asia were joined. During the periods of time that followed, many groups of fishes evolved at different times and were able to surmount the existing barriers and populate various river systems over extensive areas. This ancient continental area was joined and severed on several occasions. In the millions of years which ensued, with long periods of warm weather, probably an exchange of various faunas gradually took place. It was during these periods that some representatives of the Eurasian fauna arrived in North America and vice versa. During a portion of these ancient times the Allegheny River, with a southerly direction today, flowed northward into an area which today is partly occupied by Lake Erie.

Although this discussion may seem fantastically unreal to the student not yet initiated to the methods of distribution of freshwater inhabitants, it seems quite ordinary to the ichthyologist and zoogeographer. They vision each river and its numerous branching tributaries as excellent highways for transportation and dis- persal of fishes. Nature built its own bridges, too, for these waterways. It may be in the form of an ancient land connection, as mentioned above, bridging large geographical areas, 1 or it may be a flooding of,marine coastal areas with fresh water, thus ecologically bridging the mouths of many rivers with a waterway for the dispersal of freshwater inhabitants.

When the last massive ice sheet came down over northern Pennsylvania and portions of Ohio, the tremendous quantities of cold water killed or forced many of our warmer water species southward. 2 As the glacier receded the salmonid fishes, trouts and chars, which are probably marine in origin, were able to move into numerous cold-water streams from the unglaciated areas and oceanic waters, forming an abundant trout population. Many of these salmonid fishes became land- locked; their access to the sea was blocked primarily by the disappearance of the broad glacier connections and later by warmer downstream waters. These salmonid fishes established widespread populations over North America.

A familiar phrase and song in eastern United States are the words, "Far above Cayuga's Waters." Above this lake and others are numerous fishes which are isolated today by swift cataracts and impassable falls. Students of glacial geology have

1Charles Schuchert, "Gondwana Land Bridges," Bull. Geol. Soc. Amer., 43: 875- 916, 1932.

Bailey Willis, "Isthmian Links," Bull. Geol. Soc. Amer., 43:917-952, 1932.

2Frank Leverett, "Glacial Formations and Drainage Features of the Erie and Ohio Basins," U.S. Geol. Surv. Mono. 41:1-802, 1902.

"Glacial Deposits outside the Wisconsin Terminal Moraine in Pennsylvania," Pa. Geol. Surv. Bull. G7:1-123, 1934. 67 carefully plotted the glacial lake shores above the present lake water level. It was in these post-glacial times, when the lake water level was considerably higher than today, that the fishes got into such upland, isolated waters.

I After the periodic recession of the tremendous ice mass numerous water con- nections were in existence, linking the Mississippi valley with the historical Great Lakes. 2 The Wabash River had an ancient connection with western Lake Erie, the Illinois River with Lake Michigan near the present site of Chicago, and there were numerous other historical connections. Through these passages and the borders of the Pleistocene Great Lakes the Mississippi fish fauna passed to repopulate the Great Lakes area, reaching as far as central New York. As the ice receded from eastern Ohio and northwestern Pennsylvania the fishes moved into the upper Allegheny River system. From here some of them were able to pass into the Genesee River system using the Post-Pleistocene outlet near Olean and at a later date when this outlet was closed, some got into the Susquehanna River system by way of the Canisteo outlet. When the water flow from the Finger Lakes area was southward, the Susquehanna River had a number of glacial connections with it. These con- nections were closed when this water flow was east and south by the Hudson River through the Mohawk-Hudson connection. With continued glacial recession the latter connection remained closed when the St. Lawrence River became uncovered. The St. Lawrence connection having the lowest altitudinal outlet exists into recent times.

Thus our present fauna received its origin, prolific speciation and distribu- tion in various periods of historical times. Through intensive field work in various sections of our country, we see today clear-cut evidence of the dispersal which took place in the fish fauna in the immediate post-glacial times. 3 As examples, such widespread species of the Basin as the Northern rainbow darter (Etheostoma caeruleum caeruleum) and the Eastern tonguetied minnow (Parexoglossum . laurae) penetrated into the upper Genesee River probably from the upper Allegheny River via the Olean outlet. There is no evidence that natural connections existed . in these areas in recent historical times.

Factors Affecting the Distribution of our Fishes in Recent Times

We have touched briefly on the historical factors which affected the distribu- tion of our fishes such as the glacial connections. In recent times there have been two types of influences on distribution: (a) man-made, such as dams, pollutions, canals, and (b) natural causes. The aggregate of several of these influences has wrought dire effects on some of the fish life.

'Richard Foster Flint, Glacial Geology and the Pleistocene Epoch, John Wiley and Sons, New York, 1947.

2 Frank Leverett and F. B. Taylor, "The Pleistocene of Indiana and Michigan and the History of the Great Lakes," U.S. Geol. Surv., Mon. 53:1-529, 1915.

3Carl L. Hubbs and Robert R. Miller, "The Great Basin with Emphasis on Glacial and Postglacial Times," Bull. Univ. Utah, 38:18-166, 1948. 68 I. Recent Physiographic Connections. -- There is some evidence that in recent years the headwaters of some of our major river systems were joined in certain areas during periods of heavy rains. It is easy to conceive that during such periods the general flooding of certain present-day marsh sites could lead to an exchange of some of the typical headwater inhabitants from each system. Observa- tions by several individuals point out the feasibility of such a transfer. 1

II. Topography and Hydrography. -- The general topography and hydrography of our streams affect the general distribution of our fishes considerably, especially stream gradient. The areas of glacial drift in the upper Allegheny, having streams with relatively clear water and gravelly, sandy bottoms are preferred by the follow- ing species:

, Hybopsis biguttata eastern tonguetied minnow, Parexoglossum laurae northern redfin shiner, Notropis umbratilis cyanocephalus eastern bigmouth shiner, Notropis dorsalis keimi mountain madtom, Noturus eleutherus northern longear sunfish, Lepomis megalotis peltastes

Various other kinds of fishes are characteristic of certain streams depending on their size, bottom and flow within the general area of the Basin. We have species characteristic of large rivers; some inhabit the riffle portions of the large rivers. Others reach their greatest abundance in the small creeks. Various species exhibit distributional patterns that may be generalized or considerably localized.

The river inhabitants are as follows:

lake sturgeon, Acipenser fulvescens paddlefish, Polyodon spathula mooneye, Hiodon tergisus skipjack, Alosa chrysochloris smallmouth buffalo, Ictiobus bubalus blue sucker, Cycleptus elongatus spotted sucker, Minytrema melanops blue catfish, Ictalurus furcatus channel catfish, Ictalurus punctatus

Inhabitants of the swiftest riffles of our larger rivers:

gilt darter, Hadropterus evides bighead darter, Hadropterus macrocephalus spotted darter, Etheostoma maculatum bluebreast darter, Etheostoma camurum Tippecanoe darter, Etheostoma tippecanoe

Species characteristic of brooks and creeks: blacknose dace, Rhinichthys atratulus creek chub, Semotilus atromaculatus hornyhead chub, Hybopsis biguttata redside dace, Richardsonius elongatus sculpin, Cottus bairdi

'Shelby D. Gerking, The Distribution of the Fishes of Indiana, Invest. Ind. Lakes and Streams, Ind. Dept. Cons., 3(1):8, 1945. 69

III. Barriers. -- A number of barriers exist, man-made and natural, which affect the distribution of our fishes such as falls, dams, reservoirs, extensive polluted areas, siltation, water temperature, and various biological ones, as competition for food and spawning sites, and predation among species.

A. Falls, dams, reservoirs -- There are few natural falls of any height in the larger streams of the area but numerous dams and reservoirs made by man have certainly inhibited movement and migration of many of our species. Kirtland remarked more than a hundred years ago that the migratory species of the Mahoning River were eliminated by the construction of dams on the Beaver River. Grain mills with their small dams were constructed by the hundreds as the land of the Basin became settled and farmed. Many of these are gone today but they undoubtedly effected a reduction in many migratory species. The large reservoirs cptemplated today for flood prevention will further change the life of our rivers.

B. Pollution -- Various pollutants harmful to fish life in a section of a river may form an impassable barrier against them. This pollution may be only of a temporary nature, in which case little harm may be caused. Certain permanent effluents, whether locally in a river or widespread, form impassable barriers as effective as the most precipitous falls. The intensive mining operations with their resultant pollutants have practically wiped out the once rich fauna of the Monongahela River. Evermann and Bollman recorded 40 species of fishes from this river near the Pennsylvania-West Virginia state line in 1886 and such reductions following pollution have been observed widely from many areas.

C. Siltation - The original fauna of the Basin was generally adapted to clear water streams with gravelly, sandy bottoms. Our intensive agricultural practices and many other human endeavors 2 in the past fifty years have tremendously accentuated the erosion of our land, resulting in high siltation in our streams. Siltation has its direct and indirect harmful effects on the fish fuuna. Directly it clogs and handicaps the respiratory mechanism and may cause the extirpation of some species. Indirectly, it has wrought havoc to the ecological habitats of many other aquatic forms. It has reduced or eliminated the spawning areas for many species. The vital and often sensitive food chain of many species has been considerably altered by high, constant siltation through the reduction or elimina- tion of many invertebrate forms as well as the algae and higher aquatic plants. Aquatic plant retreats required by certain young and adult fishes have been seriously reduced by siltation and shifting stream beds. As a general consequence, the environment has been rendered unfit for many species of the original fauna in many portions of the Basin. 3 Milton B. Trautman, Franz Theodore Stone Laboratory,

1 Milton B. Trautman (Ms.) has an excellent detailed review of this subject for Ohio.

2 , "The Effects of Man-made Modifications on the Fish Fauna in Lost and Gordon Creeks, Ohio, between 1887-1938," Ohio Jour. Sci., 39(5): 275- 288, 1939.

3 C. Willard Greene, The Distribution of Wisconsin Fishes, Wisc. Cons. Comm., Madison, 235 pp., 1935. 70 Put-in-Bay, Ohio, recently presented several papers on the great changes and effects of siltation and turbidity on fishes, particularly in Ohio. 1

D. Temperature - The impetus on agriculture and lumbering during the past century has substantially reduced our forests and stream cover, thereby causing rapid water run-off in our streams and diminishing their supply of cold, spring-fed water. These factors coupled with the increased industrial and domestic consumption of water have seriously lowered our ground-water level. The water temperatures of our streams during the critical summer months are generally higher than in former days for there is less water volume, greater exposure and a re- duced supply of spring water. Some streams, once productive, are today dry gravel beds.

These conditions have restricted the cold water fauna, such as the native brook trout which was formerly widely distributed in the area, to a few, scattered, headwater streams in the Alleghenies. These mountain populations are isolated today by the warmer waters in the river courses below them.

E. Biological barriers - These barriers are often complex, and to the inexperienced observer of fishes, many may appear to be purely exaggerations. That they actually exist can be readily demonstrated but their true significance will require considerable future study. The aquaria hobbyist frequently observes various antagonisms and behavioral antics when certain species are placed to- gether. Various species in their natural environment also may show interactions with others. Some, in the presence of competition, acquire a restricted distribu- tion. The competition for food or spawning sites and predation among species are common occurrences. Dense populations may retard reproduction or excite movement.

Some species apparently exhibit characteristically favorable inter- actions when occurring together to the extent that their association is correlated with successful reproduction and high populations. For example, the , Hybopsis micropogon, and the central common shiner, Notropis cornutus chrysocephalus, successful species in the Basin, are frequently observed and captured together. This applies to them generally throughout their ranges where they occur together, and to their close relatives as the hornyhead chub, Hybopsis biguttata of the upper Mississippi and Great Lakes area, or the Piedmont chub, Hybopsis leptocephalus of the Atlantic coast, and the northern and eastern subspecies of the widely distributed common shiner, or the Roanoke Shiner, Notropis cerasinus. The chubs build large, dome-shaped nests of pebbles. The male chub works fast and hard for several days seeking stones which he carries in his mouth to his nest site. Sometimes more than 10,000 stones are gathered. 2 He thus provides an excellent site over which the

1"Effects of Turbidity and Siltation upon the Freshwater Faunas and Fisheries of the Eastern U.S?, paper presented at Pac. Div., A.A.A.S., 31st Ann. Meet., Univ. Utah, Salt Lake City, June 19-24, 1950; "Effects of Turbidity and Siltation upon the Distribution and Abundance of Fishes of Ohio," paper presented Meet. Ohio Acad. Sci., Oxford, Apr. 6, 1951; "Changes in Fish Distribution and Abundance in Ohio since 1800, Caused Primarily by Increased Turbidity and Siltation," paper presented Amer. Soc. Ichthys. and Herps., Chicago, June 27-30, 1951.

2Ernest A. Lachner, "Studies of the Biology of the Cyprinid Fishes of the Chub Genus Nocomis of Northeastern United States," Am. Midi. Nat., 48(2):435, 1952. 71 eggs are spawned. The common shiner takes advantage of this spawning site, too, as well as other daces, minnows and darters. Hankinson 1 remarked on the possible commensalism of the chub and the shiner; the former species built the nest while the latter chased away intruders. I have observed chubs active at nest building drive away intruding, small suckers seeking the eggs that were deposited on the nest, but they were not alarmed at the presence of adult shiners directly over the nest. Thompson 2 and Thompson and Hunt 3 commented on the "extreme exactness" of their association, the "two species being found together in 99 per cent of the collections in which either one of them occurred." Although the actual significance of such an association requires further study, it is certainly more than co- incidental, for the adult chubs and shiners do not have altogether the same habits.

IV. Stocking. -- The occurrence of several species in the Basin is due to introductions. The present distribution of a number of native species has been maintained or they have become greatly extended by introductions. These intro- ductions include:

1. Salmonid fishes, the trouts and chars. Three species have been stocked in recent years, the brown trout, Salmo trutta, which is an introduction to America from the European fauna, the rainbow trout, Salmo gairdneri, intro- duced from the Pacific Coast, and the brook trout, Salvelinus fontinalis fontinalis, a native inhabitant of the Basin. The fishing pressure is much greater than the capacity of our available waters to maintain these cold-water forms.

2. Carp, Cyprinus carpio. A native of Asia, it was introduced to America about 80 years ago from Europe where it was reared for centuries as a food fish in small ponds somewhat similar to the farm ponds today in America. We consider it largely as a pest and detrimental to other fishes. The Pymatuning Reservoir has an abundant population which could enter the diet of our anglers much more than at present. Prejudice toward its name and the lack of knowledge of the proper cooking methods have led to its neglect. Our comparatively abundant hoofed meats and sea food products have also contributed to its neglect. The carp becomes over-populated in some areas and causes a reduction of the more desirable food and game species.

3. Goldfish, Carassius auratus. A common aquaria species of America today, this native of eastern Asia has been reported in several localities in the Basin but nowhere is it common. It prefers warm, quiet, weedy waters. Its presence in streams may be associated with discouraged aquarists, or it may escape from garden pools and ponds.

'Thomas L. Hankinson, "Observations on the Breeding Behavior and Habitats of Fishes in Southern Michigan," Pap. Mich. Acad. Sci., Arts & Letrs., 15(1931):418.

2David H. Thompson, "A Survey of the Fishes of Champaign County, Illinois," Trans. Amer. Fish. Soc. 59:5, 1929.

3David H. Thompson and Francis D. Hunt, "The Fishes of Champaign County," Bull. Ill. Nat. Hist. Surv. 19(1):66-69, 1930. 72 4. Ohio muskellunge, Esox masquinongy ohiensis. Recent attempts have been made to build up the population by stocking this rapidly diminishing, native form for it is one of the angler's choice species.

5. Killifish family, Cyprinodontidae. Two species have been introduced into the area as forage or bait fishes, the eastern banded killifish, Fundulus diaphanus diaphanus and the mummichog, Fundulus heteroclitus macrolepidotus, which has a very limited distribution in the Basin. These forms were originally inhabitants of the Atlantic coastal streams.

6. North American catfish, bass, perch, and sunfish families. Many native warm water fishes of the area have been stocked in recent years due to their excellent qualities as game, pan or forage fishes. The northern smallmouth bass, Micropterus d. dolomieui and the northern largemouth bass, Micropterus s. salmoides have received widespread stocking in streams, lakes or ponds. The current interest in farm ponds has partly accounted for the wide distribution maintained in several species. The northern largemouth bass and the northern bluegill, Lepomis m. macrochirus are most frequently stocked in farm ponds but others along with these have been tried, some of which are: northern brown bull- head, Ictalurus n. nebulosus; northern black bullhead, Ictalurus m. melas; yellow bullhead, Ictalurus n. natalis; white bass, Roccus chrysops; pumpkinseed, Lepomis gibbosus; white crappie, Pomoxis annularis; black crappie, Pomoxis nigromaculatus; and yellow perch, Perca flavescens.

V. Canals. -- The upper Ohio, Allegheny and Susquehanna Rivers were connected by interlacing canals with Lake Erie and Lake Ontario, all of which were completed more than 100 years ago. It is doubtful if they had much effect on the general distribution pattern of the fishes of this area. Certainly more intensive field work is required in all areas where the Basin was connected by canals with other drainages in an attempt to acquire further evidence. No example of a species is as obvious as the transfer through canal connection of the sea lamprey, Petromyzon marinus, from Lake Ontario, where it was formerly land-locked, to Lake Erie, by- passing Niagara Falls via the Welland Canal.

VI. Climate. -- There is little visible direct relationship between climate and the distribution of fishes in the Basin. The correlation here is one of creat- ing a differential in the rate of growth, time of reproduction, productivity, and other conditions among the species in various years, just as we may find in agricultural crop production. Indirectly, unsuccessful reproduction can alter the local distribution of any species considerably.

Changes in the Fauna

We can construct a picture of the distribution of many of our fishes, past and present, primarily from the efforts of a number of individuals studying the fishes of various areas of the Basin. The early attempts to study the fauna give us today a comparative measure of the changes that have occurred and their type and amount. 73

1 The garliestL men to study the fauna of the Basin were LeSueur and Rafin- esque. These men named new forms from the Ohio River. Several species re- ported in the Ohio as far as Pittsburgh by Rafinesque, among which were the shovelnose sturgeon and shortnose gar, have not been seen since. Kirtland 3 published valuable descriptions and accounts of the fishes of Ohio. The type locality of the hornyhead chub, Hybopsis biguttata (Kirtland) and the beautifully colored darter, Etheostoma variatum (Kirtland), was the Mahoning River, long since almost entirely void of fish life because of heavy industrial pollution. Cope published several valuable papers on the distribution and habits of fishes of Pennsylvania; among these were his "Synopsis of the Cyprinidae of Pennsylvania" (1869) and "The Fishes of Pennsylvania" (1881). The report of 40 species of fishes from the Monongahela River by Evermann and Bollman (1886) is a valuable historical faunal treatise of an area badly polluted for many years. 4 The fishes of the upper Monongahela in West Virginia were later treated by Goldsborough and Clark (1908). 5 Bean's (1892) account of Pennsylvania's fishes was too generalized to be of much value. 6 Fowler published a number of papers recording the distribu- tion and habits of fishes in Pennsylvania; those of especial value to the knowledge of the fishes of the Basin are his studies of 1909, 1919, 1927, and 1940.

1C. A. LeSueur, "A new Genus of Fishes of the Order Abdominales, Proposed under the name of Catostomus and the Characters of this Genus, with Those of its Species Indicated," Jour. Acad. Nat. Sci. Phila. 1(1):88-96 and 102-111, 1817. Also, "Descriptions of Several New Species of North American Fishes," Jour. Acad. Nat. Sci. Phila. 1(2):222-235 and 359-368, 1818.

2 C. S. Rafinesque, "Ichthyologia Ohiensis, or Natural History of the Fishes Inhabiting the River Ohio and its Tributary Streams," edit. by R. E. Call, The Burrows Bros. Co., Cleveland, 1820.

3 Jared Potter Kirtland, "Report on the Zoology of Ohio," Ann. Rept. Geol. Surv. St. Ohio, 2:157-200, 1838; "Descriptions of the Fishes of the Ohio River and its Tributaries," Boston Jour. Nat. Rust., 3:338-352, 1840; and "Descriptions of the Fishes of Lake Erie, the Ohio River, and their Tributaries (concluded), "Boston Jour. Nat. Hist., 5:330-344, 1847.

4 Barton W. Evermann and Charles H. Bollman, "Notes on a Collection of Fishes from the Monongahela River," Ann. N.Y. Acad. Sci., 3:335-340, 1886.

5 Edmund Lee Goldsborough and H. Walton Clark, "The Fishes of West Virginia," Bull. U. S. Bur. Fish., 27:29-39, 1908.

6 Tarleton H. Bean, "The Fishes of Pennsylvania," Rept. State Comm. Fish. Pa., 1889-91:1-149.

7 Henry W. Fowler, "A Synopsis of the Cyprinidae of Pennsylvania," Proc. Acad. Nat. Sci. Phila. 60(1908):517-553; "A List of the Fishes of Pennsylvania," Proc. Biol. Soc. Wash. 32:49-74, 1919; "Fishes from McKean, Potter and Cameron Counties, Pennsylvania," Proc. Biol. Soc. Wash. 40:65-74, 1927; and "A List of the Fishes Recorded from Pennsylvania, Pa. Bd. Fish. Comm., Bull. 7:1-25, 1940. 74 McConnell (1906) reported new records and Keim (1915) listed the fishes at Port Allegany. 19 2 The Allegheny River in New York was surveyed by the New York Conservation Department, and the fishes of the area were reported by Greeley. 3 A recent and detailed study of the distribution of the Ohio drainage Basin of western Pennsylvania is yet unpublished (Raney, ms.), but his abstract (1939) lists the species involved. 4

Following Kirtland's early studies of Ohio's fishes, Klippart (1877) 5 pub- lished a catalogue and Osburn (1901) The Fishes of Ohio. These studies were continued for Ohio by Osburn, Wickliff and Trautman (1930), and Trautman (1950). The most detailed study of the fishes of Ohio is also yet unpublished (Traut- man, ms.).

Numerous extra-limital studies assist our understanding of the past and present distribution of the fishes of the Basin, some of which are those of Woolman (Kentucky), 6 Evermann (Kentucky and Tennessee),7 Forbes and Richardson (Illinois),6

1W. R. McConnell, "Preliminary Report of the Investigation of Certain Waters of Pennsylvania," Rept. Dept. of Fish. Pa. 1905:172-179, 1906.

2Thomas D. Keim, "Notes on the Fauna about the Headwaters of the Allegheny, Genesee and Susquehanna Rivers in Pennsylvania," Copeia, 24:51-52, 1915.

3John R. Greeley, "Fishes of the Area with Annotated List," in A Biological Survey of the Allegheny and Chemung Watersheds, Suppl. Ann. Rept. N.Y. St. Cons. Dept. 1937:48-73, 1938.

4Edward C. Raney, "The Distribution of the Fishes of the Ohio Drainage Basin of Western Pennsylvania," Cornell Univ. Thesis, (1938).

5 J. H. Klippart, "Catalogue of Fishes of Ohio," in Report Ohio State Fish Comm., 1875-76: 143-195, 1877.

6 A. J. Woolman, "Report of an Examination of the Rivers of Kentucky, with Lists of the Fishes Obtained," Bull. U. S. Fish Comm., 10:249-288, 1892.

7 Barton Warren Evermann, "The Fishes of Kentucky and Tennessee: A Distributional Catalogue of the Known Species," Bull. U.S. Bur. Fish., 35:293-368, 1918.

8 Stephen A. Forbes and Robert E. Richardson, The Fishes of Illinois, (second edit) Nat. Hist. Surv. Ill., vol. 3, 1920. 75 2 Hubbs (Great Lakes),1 Greeley (Genesee R., New York, and Susquehanna R., New York), Gerking (Indiana), and Hubbs and Lagler (Great Lakes).4

The recent group revisionary studies of our North American freshwater fishes, such as those completed by Reeve M. Bailey, Museum of Zoology, University of Michigan, Carl L. Hubbs, Scripps Institution of Oceanography, La Jolla, California, and Edward C. Raney, Cornell University, and their doctoral students and asso- ciates have also contributed substantially to our understanding of the fishes of this Basin.

We learn today from a careful comparison of these faunistic and revisionary studies that the primary changes in the fauna are two-fold: (1) in the complete extirpation of certain species, or in their reduction to greatly reduced popula- tional densities or in their limitation to minor local distributions; (2) in the resurgence of a replacement fauna.

Almost all of the following 18 species were reported before 1900 in the upper portion of the Basin in Pennsylvania and northward in some of the tributaries of the Monongahela or Allegheny Rivers, some of which were common or abundant, but hvve not been reported in recent years. Perhaps some of these species may still be found in the Ohio River of Pennsylvania below Pittsburgh but the general data warrant the thesis that there has been a great reduction in numbers and the ex- tirpation of many forms.

paddlefish, Polyodon spathula lake sturgeon, Acipenser fulvescens shovelnose sturgeon, Scaphirhynchus platorhynchus shortnose gar, Lepisosteus platostomus bowfin, Amia calva goldeye, Hiodon alosoides gizzard shad, Dorosoma cepedianum

'Carl L. Hubbs, "A Check-list of the Fishes of the Great Lakes and Tributary Waters, with Nomenclatorial notes and analytical Keys," Misc. Pub. Mus. Zoo. Univ. Mich. 15:1-77, 1926; also "Further Additions and Corrections to the List of the Fishes of the Great Lakes and Tributary Waters," Pap. Mich. Acad. Sci., Arts & Letrs., 11:425-436, 1930.

2 John R. Greeley, "Fishes of the Genesee Region with Annotated List," in A Biological Survey of the GEnesee River System, Suppl. 16th Ann. Rept. N.Y. St. Cons. Dept., 47-66, 1927; also "Fishes of the Area with Annotated List," in A Biological Survey of the Delaware and Susquehanna watersheds," Suppl. Ann. Rept. N. Y. St. Cons. Dept., 45-88, 1936.

3 Shelby D. Gerking, 5.1p cit.

4 Carl L. Hubbs and Karl F. Lagler, "Fishes of the Great Lakes Region," Bull. Cranbrook Inst. Sc., 26:1-109, 1947. 76 smallmouth buffalo, Ictiobus bubalus highfin sucker, Carpiodes velifer river carpsucker, Carpiodes carpio blue sucker, Cycleptus elongatus sturgeon sucker, Catostomus catostomus river shiner, Notropis blennius silvery minnow, Hybognathus nuchalis blue catfish, Ictalurus furcatus American eel, Anguilla rostrata sauger, Stizostedion canadense freshwater drum, Aplodinotus grunniens

The following species are only rarely captured in this area today:

Ohio lamprey, Ichthyomyzon bdellium shorthead redhorse, Moxostoma breviceps spotted sucker, Minytrema melanops flathead catfish, Pilodictis olivaris slenderhead darter, Hadropterus phoxocephalus warmouth bass, Chaenobryttus gulosus

The alligator gar, Lepisosteus tristoechus, was apparently last recorded in the Ohio near Cincinnati in 1838. i

In his studies of Ohio fishes Trautman (ms.) points out that the tremendous siltation in Ohio's streams in recent years has extirpated such forms as the bigeye shiner, Notropis boops. The creek chubsucker, Erimyzon oblongus, formerly widely distributed in Ohio, is also considerably reduced today. The hairlip sucker, Lagochila lacers, is probably extinct. In contrast to the depletion of the original fauna, he sees certain species increasing in abundance and distribu- tion. The suckermouth minnow, Phenacobius mirabilis, originally an inhabitant of the turbid plain streams, now reaches the Appalachian foothills. Two other species, the fathead minnow, Pimephales promelas, and the bluntnose minnow, Pimephales notatus, extremely tolerant to the silty streams, show a greatly increased dis- tribution in Ohio in recent years. Fishes such as the blacknose shiner, Notropis heterolepis, an inhabitant of clean, clear vegetative streams, also formerly widely distributed in Ohio, may be completely eliminated by siltation.

We thus witness the probable extinction of one form, the hairlip sucker, and the extirpation or reduction in distribution of many desirable food and game fishes, such as the pikes, walleyes, catfishes, buffalofishes, suckers, drums, sturgeons and salmonid fishes.

A Program of Research and Conservation

There is great need for a concerted interstate program to conserve our existing aquatic resources. We must rehabilitate many areas to increase the quantity, quality and productivity of many forms of life. Legislation and new methods of management must be sought in order to meet the tremendous demands of future recreation enthusiasts. In an effort to maintain an adequate supply of several prized game fishes in our waters such as the upper portion of the Basin in western

1 Kirtland, 1838, 22_1. cit. 77

Pennsylvania, the State has annually stocked numbers of fish. Yet we are generally aware of a considerable reduction in the abundance of a number of species over the past years. Generally, our procedures of (1) fishing legis- lation and enforcement and (2) stocking programs do not cope with the factors that are depleting our fish populations. We recognize that in order to main- tain adequate populations of desirable game and forage fishes their (1) intro- duction into new waters, (2) propagation for stocking, and (3) production through legal restriction and enforcement are not sufficient. Our freshwater fishes must be regulated as an agricultural crop, the abundance of which can be more efficiently maintained and controlled only through a "general management policy." This policy must embody certain fundamental principles already known concerning the biology of our fishes, their conservation and utilization. We must seek new procedures, tested -hrough research and investigation, and introduce those that prove desirable.

As we have reviewed above, the factors primarily responsible for the deple- tion of the various species are (1) humanistic, (2) environmental, and (3) biological.

Each decade we observe an increase in the fishing intensity of our waters. More sportsmen are afield and with the trend toward shorter working hours we can expect an ever growing army of anglers. A considerable fish crop is harvested annually, yet we have no satisfactory estimate of the abundance of stock in the streams.

The environmental changes caused by floods, soil erosion, deforestation, temperature changes and variable climatic conditions all take their toll directly or indirectly of a given species. Water impoundments have altered the abundance of our fishes through changes in the environment. While some of these impoundments have reduced certain fishes, their construction, as the large flood prevention reservoirs now underway in several states, may increase certain warm water game and pan fishes.

It is seen that the aggregate of these factors may reduce the biological productivity of a given species through reductions and changes in the supply of natural foods, encouraging predation, restricting migrations, reducing habital waters, decreasing spawning sites and many other causes.

We arrive at a critical stage in our fisheries program. The great value of our freshwater fish heritage should be generally realized. Numerous game fishes provide millions of sportsmen with recreation. Our food and pan fishes enter the daily diets of many people. Our forage fishes not only supply the natural food for game fishes but also serve as a supply source for business enterprises as bait dealers and aquarists. This fish resource is of great indirect value through the sale of millions of dollars of gear and equipment and in the build- ing and equipment of numerous resort areas. Probably the greatest value of this fish resource is the recreation enjoyed by so many, both young and old, through the "wholesome use of leisure time, an immeasurable value."

We must use and manage intelligently our game, pan and forage fishes. It is timely and definitely necessary to obtain a more complete understanding of their abundance, mode of living, and intricate relationships in this Basin. This can only be done through planned investigaiions toward solution of various objective problems. We cannot rely on biased assumptions and unfounded principles and 78

regulations for the maintenance of adequate fish populations. This natural resource has dwindled to a critical minimum in respect to the increasing number of anglers.

We need intensive research programs to obtain new procedures in fisheries management, and plans for problems of the future. We should arrange for the training of personnel for present and future management and conservation re- quirements. We should prepare and publish useful and vital fisheries informa- tion for public consumption. The basic knowledge of intelligent management and wise conservation must be brought before them. We require a broad, integrated, inter-state program combining the efforts of such departments as those of public health, sanitation, game, fish, forest, and education-recreation. We must intro- duce and teach conservation practices in our elementary and secondary schools, and have some of these regarded as curriculum requirements. The immediate and future problems of our streams must be brought before the public through radio, television and the press. Considering the best possible organization and suc- cess, we can only anticipate a hard, uphill grind for a partial rejuvenation of life in "our waters."