Freshwater Fishes by EDWARD C. RANEY, Associate Professor of Zoology and Fishery Biology, Department of Conservation, Cornell University

HISTORICAL REVIEW AND SOURCES OF DATA O SYSTEMATIC SURVEY has ever been made of the fish fauna N of the James River nor of any of the rivers of . Much of the available information on distribution is taken from studies com- pleted sixty or more years ago. Since that time various ichthyologists in a number of scattered papers have added to our knowledge of James River fishes. However, the study of the ecology, distribution, and habits of these fishes, especially the small non-commercial , offers an interesting and fruitful field for local students. Since some of the upper reaches of the main James River as well as areas below large cities are now badly polluted the precise distribu- tion of the original fish fauna will never be known. Fortunately two early ichthyologists, Cope (1869) and Jordan (1890) collected in the mountains and in the upper Piedmont area. Their work was rather thorough especially when one considers the difficulties of transporta- tion in those days. For a period of almost fifty years little interest was shown in the distribution of the James River fishes. During this time Fowler (1918, 1923a and b, 1924) reported on several isolated collections made in the James, and on a re-examination of Cope's mate- rial. In the years 1933 to 1935 collecting was resumed by Dr. George S. Myers and assistants from the U. S. National Museum. These speci- mens are deposited in the collections of the National Museum and are available for study. However, they have not been reported on except incidentally in connection with several generic revisions and in the descriptions of new species. In 1937 and 1938 Dr. Leonard P. Schultz, Curator of Fishes, and Earl D. Reid, formerly Scientific Aid, in the U. S. National Museum, made fifty-two collections in the several river systems of Virginia. Seven of these were from the James or its tributaries. A short summary of these explorations was published by Schultz (1939). Dr. Schultz encouraged the writer, then embarking on a large treatise intended to cover the freshwater fishes of the eastern United States, to continue the field work with the purpose of ultimately writing a Fishes of Virginia. To secure fishes for these projects col- lections were made during ten day periods in late March and early April in 1940, 1941, 1947, and 1948 and during mid-June 1946. A

151 152 THE JAMES RIVER BASIN total of 108 collections were made in Virginia and 20 were from the James River system. Considerable help toward meeting the expense of these trips was obtained from research grants from the Trustee- Faculty Research Committee of Cornell University. My associates, then graduate students, on these trips were L. James Kezer, Ernest A. Lachner, Roman A. Pfeiffer, Robert D. Ross, and Royal D. Suttkus. Much credit is due them for their untiring efforts and thoughtful co- operation in the field. Others became interested in the distribution of the fishes of the state and furnished specimens. Among these are Dr. Lincoln C. Pettit, then on the biology staff of Washington and Lee University, who made 19 collections in 1940 and 1941 in Rockbridge Co. Dr. Horton H. Hobbs and Richard L. Hoffman, University of Virginia, made available 7 small collections taken during 1946 from the tribu- taries of the James and Appomattox rivers. Hoffman also made avail-. able 4 small collections taken in 1946 in Alleghany Co. These collections of Schultz, Raney, Pettit, Hobbs, and Hoffman total 57 and serve as the original source of distributional data men- tioned in the species accounts that follow. Other hitherto unreported material from the James River deposited in the U. S. National Museum, as well as many of Jordan's (1890) specimens have also been used in this study. Besides the literature sources mentioned above several other works have appeared. Burton and Odum (1945) made a study of the dis- tribution of fishes found in the headwater streams in the neighbor- hood of Mountain Lake Biological Station. Included in this study are 16 collections from Johns Creek and 12 collections from Craig Creek, both tributaries of the James River. This type of study is recom- mended as the kind that students of vertebrate can carry on with great profit once the local fauna is learned. To this end the writer is glad to offer his services in making identifications of fishes. In 1942 Dr. Carl L. Hubbs drew up and made available a manu- script check list of the fishes of the James River system which was very helpful especially in the early stages of this paper. Fowler's (1945) monograph of the southern Piedmont and Coastal Plain fishes lists the species known from the James River system in a distribution table. Little new material is included and various com- ments on his records are found below. The saltwater fishes of have been studied for many years. Hildebrand and Schroeder (1928) give detailed descrip- tions, information on the life histories, and data on the distribution of the fishes of Chesapeake Bay. This fine work includes many fresh- water fishes that occasionally are found in brackish water. It also discusses saltwater fishes that occasionally wander into freshwater or that typically migrate into rivers during their spawning runs. Pearson FRESHWATER FISHES 153

(1941) also studied the young of marine fishes found in lower Chesa- peake Bay. The facilities of the Virginia Fisheries Laboratory, of the Col- lege of William and Mary and the Virginia Commission of Fisheries, located at Williamsburg, with a field station at Yorktown, have been devoted to the study of the tidewater fisheries of the lower James. Most of the published material from this agency deals with com- mercially important and crustacea. However, lenzel's (1945) study of the industry of Virginia was supported by this station. Studies of other fishes are now under way. For descriptions and keys to the various James River fishes the reader is referred to the general standard works such as Jordan and Evermann (1896), and Jordan (1929). Additional keys and illustra- tions to some of the species are given in Hubbs and .Lagler (1947). When available, references to more recent studies on the systematics and life histories are given under the species accounts.

ANNOTATED LIST OF JAMES RIVER FRESHWATER FISHES The freshwater fish fauna of the James River is classified under 21 families, 52 genera and 73 species. One species, Boleosoma nigrum, is represented by two distinct subspecies making a total of 74 kinds of fishes. In addition 14 species are given on a hypothetical list of fishes that probably will be found in the drainage when an exhaustive survey is made.

LAMPREY FAMILY PETROMYZON IDA E Sea Lamprey Petromyzon niarinus Linnaeus This parasitic species has recently been taken on fishes captured in the lower James River. Several such specimens are deposited in the U. S. National Museum. Although adults are normally found in salt water, it is an anadromous species. About May it migrates up- stream into small tributaries to deposit the eggs in a circular nest built by one or several males. The mature adults reach a large size and are upward of 18 inches long. It rasps circular holes in the side of its prey which sometimes result in death. The eastern brook lamprey, Lampetra aepyptera (Abbott), is a freshwater species that has been found in the Rappahannock and Patuxent River systems, two more northern tributaries of Chesapeake Bay, as well as to the south in the Neuse River system, (Raney, 1941:318). Adults of this non-parasitic species should be looked for in small nests in riffles of brooks where they in April and May. The larvae, like those of the sea lamprey, burrow 154 THE JAMES RIVER BASIN in the sand and silt of the stream bottom in quiet places for a period of about five years. Seldom exceed 8 inches in length. The larvae are of value as fish bait in some localities.

STURGEON FAMILY ACIPENSERIDAE Atlantic Sturgeon Acipenser sturio oxyrhyncbus (Mitchill) An anadromous species that enters Chesapeake Bay in April and May. They make their way up the larger rivers including the James and spawn in freshwater. At present they are not common especially in the larger size groups. They reach a weight of well over 100 pounds and 108,900 pounds were taken in the James River in 1880, but their numbers have seriously declined in recent years. They are valued chiefly for the roe from which caviar is made, but the flesh is eaten either fresh or smoked. Most are taken in pound nets. Hilde- brand and Schroeder (1928:72) give a detailed account of the Amer- ican sturgeon in Chesapeake Bay. Fowler (1945:21) records it from the James without listing a definite locality. The shortnose sturgeon, Acipenser brevirostris Le Sueur, has been recorded by Hildebrand and Schroeder (1928:76) from Chesapeake Bay but if present in the James River it is probably uncommon.

GAR FAMILY LEPISOSTEIDAE Longnose Gar Lepisosteus osseus osseus (Linnaeus) Actual records of its occurrence in the lower James basin are rather nebulous but it is expected in the Coastal Plain area. It is known from Chesapeake Bay and other rivers tributary to the Bay. Hilde- brand and Schroeder (1928:78) have summarized the locality records. It has been reported from the "vicinity of Norfolk" and from Lynn- haven Roads. Jordan (1890:114) mentions that the longnose gar is found in Lake Drummond, Dismal Swamp, although no specimens were taken there by him. The gar is a fish eater that is usually found in sluggish water and which often reaches a length of 4 feet. Because of its predatory habits it is usually in bad repute with fishermen. The flesh is of some slight value as food but the eggs are poisonous.

BOWFIN FAMILY AMIIDAE Bowfin Amia calva Linnaeus A relict freshwater species known definitely from the Chicka- hominy River, twenty miles from its mouth, James City County, where two specimens were secured on November 29, 1944, by R. W. Menzel. These specimens as well as several from Dismal Swamp are deposited in the U. S. National Museum. This predacious species is probably FRESHWATER FISHES 155 limited to sluggish areas in the Coastal Plain and lower Piedmont re- gions. Although sometimes eaten it is not considered a good food fish.

HERRING FAMILY CLUPEIDAE This large family of mostly saltwater food fishes is represented in the James River Basin by only one freshwater species, the gizzard shad, Dorosoma cepedianzim. Even this species is frequently found in brackish and occasionally in saltwater. Three species are common anadromous fishes that regularly enter the freshwater to spawn in early spring. They are important commercial species, namely, the American shad, Alosa sapidissima, the glut herring, Pomolobus aestivalis and the alewife, Pomolobus pseudoharengus. Two other marine species, the hickory shad, Pomolobus mediocris (Nlitchill) and the Atlantic menhaden, Brevoortia tyrannus Latrobe although normally saltwater fishes are often found in Chesapeake Bay and frequently wander into freshwater. Two marine fishes related to the herrings, the ten pounder, Elops saurus Linnaeus, and the tarpon, Tarpon atlanticus ( Valenciennes), are rarely found in the lower James. The anchovy, Anchoa mit- chilli nzitchilli ( Valenciennes), of the family Engraulidae which is one of the most abundant fishes in Chesapeake Bay also occasionally enters freshwater.

Glut Herring Pomolobus aestivalis (Mitchill) A saltwater fish that regularly migrates into freshwater rivers a short distance to spawn in April and May. An important commer- cial herring which attains a length of 15 inches and almost one pound in weight according to Hildebrand and Schroeder (1928:88). They are fairly common in the lower part of the James River.

Alewife Pomolobus pseudoharengus (Wilson) Like the glut herring it enters the rivers to spawn. Hildebrand and Schroeder (1928:90) observed that they usually ran in March, three or four weeks earlier than the glut herring, and somewhat be- fore the first run of shad. They move up to the small tributaries to lay their eggs. Most of the young return to the sea in the autumn of their first year. Adults attain a length of about 15 inches and average a little less than one pound. An important commercial species. Jordan (1890:15) found young alewives in Shingle Creek near Suffolk.

American Shad Alosa sapidissima (Wilson) An important commercial fish that enters Chesapeake Bay in March, migrates up the rivers only a short distance to spawn, and then 156 THE JAMES RIVER BASIN wanders back down to the saltwater, usually by June 1. The young usually move back to saltwater by November or December. Most adults in Chesapeake Bay weigh from 4 to 6 pounds and are 19 to 22 inches long, the females being slightly heavier than the males. The top weight is about 12 pounds. According to Hildebrand and Schroeder (1928:100) from whom the above summary is taken, the shad enters virtually all streams tributary to Chesapeake Bay. Jordan (1890:111) found young in Swift Creek north of Petersburg.

Gizzard Shad Dorosoma cepedianum (Le Sueur) A freshwater species found in the Coastal Plain area and which is often found in brackish water but rarely in strictly saltwater. Al- though they may attain a length of 15 inches and a weight of 1 Y2 or 2 pounds they are of little value as food., However, they are of im- portance as forage for other more desirable fishes. Jordan (1890:111) reported it as common in Swift Creek and lowland streams and also took it commonly in Shingle and Spring Creeks near Suffolk. Al- though no recent specimens have been reported this species is prob- ably still present in the lower James River.

SALMON FAMILY SALMONIDAE The eastern brook , Salvelinus f. fontinalis, is the only native trout present in the upper James. It is quite probable that both the brown trout, Salmo trutta Linnaeus, and the rainbow trout, Salmo gairdnerii Richardson, are also present through introductions. Cer- tainly the lower courses of some of the mountain streams should pro- vide adequate habitat for the rainbow which is now stocked. Eastern Brook Trout Salvelinus fontinalis fontinalis ( Mitchill) A superb game fish found natively in mountain streams in the headwaters of the James River. Cope (1869:238) noted its presence in streams rising in the mountains and Jordan (1890:111) reported it from all suitable tributaries of the James River in the Blue Ridge and Allegheny Mountains. Fowler (1923b:9) records it from the head of the James River. More recently Burton and Odum (1945:186 and 187) found it at 6 out of 16 stations in Johns Creek and at 4 out of 12 localities in Craig Creek. The writer recently took a juvenile speci- men in Muddy Creek, a tributary of Jackson River, Bath County.

SUCKER FAMILY CATOSTOMIDAE Four species of sucker are known from the James River and it is quite probable that two or three others also will be found inhabiting its waters. The common white sucker, Catostomus commersonni corn- FRESHWATER FISHES 157 mersonnii, the hog sucker, Hypentelium nigricans, and the rosyside sucker, Thoburnia rhothoeca, are common to abundant in the James River Basin. A fourth species, the eastern creek chubsucker, Erimvzon oblongus oblongus, is less common, but may be present in numbers greater than are now known in sluggish Coastal Plain creeks. There is a very good chance that two others, the eastern lake chub sucker, Erimyzon sucetta sucetta Lacepede, and the eastern red- horse, Moxostoma macrolepidotum (Le Sueur), also will be recorded soon. Both are known from Chesapeake Bay tributaries to the north. The former should be found in the sluggish ponds and creeks of the Coastal Plain. The eastern redhorse is a river species which the writer has captured recently in the Rappahannock River at Fredericksburg. The lake quillback, Carpiodes cyprinus cyprinus (Le Sueur), is to be looked for in the river channel. Hubbs and Lagler (1947:50) gave its range on the Atlantic Coast as from the Susquehanna to the Roanoke system. There are a number of specimens in the U. S. National Museum from the and from Havre de Grace but I know of no specimens from the James. There is also a possibility that another member of this occurs in the James since Jordan (1890:115) recorded Moxostoma papillosum from Shingle Creek near Suffolk. Additional fresh material is needed to check this record. The Atlantic Coastal species of Moxostoma are in need of revision. The spotted sucker, Minytrema melanops (Rafinesque), is of doubtful oc- currence both to the north and to the south of the James. The north- ern jumprocks, Moxostoma cervinum (Cope), has been erroneously re- ported as occurring in the James and probably does not. (See note under Thoburnia below.) See Hubbs (1930) for the most recent re- vision of eastern suckers.

Common White Sucker Catostomus commersonnii commersonnii (Lacepede) Fairly common in the Piedmont and the lower courses of mountain streams. Replaced in part in the small mountain brooks by the hog sucker and rosyside sucker. Found in 12 out of 57 recent collections and also recorded by Cope (1869), Jordan (1890), and Fowler (1923a and b and 1945). Prefers the quieter waters of creeks and small rivers except in the spring, April and May, when it migrates upstream and spawns in riffles. Burton and Odum (1945) did not find the common white sucker in their survey of the headwater streams, Craig and Johns creeks. Most adult specimens found in the smaller streams range from 1 to 2 feet in length. Hildebrand and Schroeder (1928:120) found that in the large tributaries of Chesapeake Bay they may reach 2 feet in length and up to 5 pounds in weight. The flesh is good especially in the early spring. 158 THE JAMES RIVER BASIN

Common Hog Sucker Hypentelium nigricans (Le Sueur) Common in small and moderate sized streams in the mountains and in the Piedmont region. It prefers the faster waters of riffles, but it is often seen in quiet areas in small streams. Spawns in riffles in April and May. Found in 14 out of 57 recent collections. In the mountain tributaries it shares the riffle habitat with the rosyside sucker. Burton and Odum (1945:186 and 187) collected it in 3 out of 16 places in Johns Creek and 2 out of 12 in Craig Creek. Cope (1869:240), Jordan (1890:109) and Fowler (1923a and b, and 1945) have recorded it. Raney and Lad-flier (1946a and 1947) have given its habits and charac- ters, and have compared it with a recently described hog sucker, Hypenteliztm roanokense Raney and Lachner, a species that is known only from the headwater streams of the Roanoke River system. Sel- dom reaches a length of more than 15 inches and is edible although rather bony.

Eastern Creek Chubsucker Erimyzon oblongus oblongus (Mitchill) Normally found in sluggish streams, backwaters, and ponds. It is probably most common in the Coastal Plain area where the least col- lecting has been done. Present in only 3 out of 57 recent collections made mostly in the Piedmont and mountain streams. Schultz and Reid took it in Big Lickinghole Creek west of Goochland; Pettit found one in a small pond, a half mile northeast of Buena Vista, Rock- bridge County; and Ross, Suttkus, and Raney collected an adult male in Tye River, three miles northwest of Roseland, Nelson County. Jordan (1890:115) recorded it as common in Shingle Creek near Suf- folk, and Fowler (19236:12) reported it from Richmond. Most speci- mens are under one foot in length although in lakes they sometimes reach a larger size. Of little value as food for humans but a fairly good forage fish for ponds. The eastern lake Chubsucker, Erimyzon sucetta sucetta (La- cepede), has also been recorded from Virginia (locality not given) by Hubbs (1930:35). Since it is widely distributed along the Coastal Plain from to it very probably occurs in the lower James River Basin.

Rustyside Sucker Thoburnia rhothoeca (Thoburn) Abundant in riffles of small streams in the mountain region of the James. Also found in headwater streams in the upper Piedmont in- cluding small tributaries of the Appomattox River. A map showing its distribution, as well as that of the closely related Thoburnia hamil- toni Raney and Lachner which is known only from the Roanoke River system, is given by Raney and Lachner (1945c:221). These writers FRESHWATER FISHES 159

(1946b:675) have also discussed the age, growth and habits of this montane form. Fowler (1923a:9 and 1923b: 12) records Thoburnia rhothoeca from Midway Mills and from Catawba Creek at Kyle. Hubbs (1930:43) has reviewed its taxonomic history. Its abundance is attested by the fact that it has been taken in 32 out of 57 recent collections. In addition Burton and Odum (1945:186) found it at 13 out of 16 stations in Johns Creek and at 7 out of 12 in Craig Creek. A small species that rarely attains eight inches in length. Spawning occurs in early April in riffles. Both Cope (1869) and Jordan (1890) failed to correctly identify this form. Both confused it with the northern jumprocks, Moxostoma cervinum, which does not occur in the James River as far as is now known. Fowler's (1923b:13 and 1945:24) records of Moxostoma cervinunz are based on Cope's records. As Raney and Lachner (1945c: 219) have pointed out, Cope (1869:236) confused these two species in the original description. M. cervinum is found in the mountain streams of the Roanoke River system and southward.

MINNOW FAMILY A large family of small fishes that contributes 21 known species to the fauna of the James. Known locally under various names such as chubs, dace, , and shiners. Seven species of the genus Notropis are known and another, the iron-color shiner, Notropis chalybaeus (Abbott), is expected on the Coastal Plain where it has been taken both to the north and south of the James. Two others are probably present as introductions, namely, the carp, Cyprinus carpio Linnaeus, and the goldfish, Carassius auratus (Linnaeus). The largest native in the James is the fallfish which under favorable conditions may reach 18 inches in length. Most of the other species do not exceed 10 inches in maximum length and many are less than 5 inches long. In many of the minnows the males build interesting nests at spawning time which is usually spring or early summer. The fallfish, river chub, Carolina chub, and cutups gather stones and pile them in circular heaps. Others, the creek chub and stoneroller, build one or several cup-like depressions above the riffles. Some kinds such as the common shiner, rosyface shiner, and redbelly dace use the nests built by other minnows or spawn elsewhere in or above riffles. Interspecific and intergeneric hybrids often result from the spawning in close proximity of several species. The bluntnose fastens her eggs on the underside of a stone or board and they are guarded by the male. The satinfin shiner uses the top side of submerged logs or rocks. The minnows are excellent subjects for observation and much remains to be learned of their behavior. 160 THE JAMES RIVER BASIN

Few cyprinids are of direct importance although the fallfish, creek chub, river chub, and golden shiner are sometimes taken on hook and line. They are more often used as bait for large and more desired fishes, and in nature they also serve as forage for other species.

Fallfish Leucosomus corporalis (Mitchill) The fallfish is the largest native eastern minnow and it is rather widely distributed in the upper half of the James River Basin. Usually it prefers the pool areas in medium and large streams and is less common in the smaller headwater brooks. Present in 18 out of 57 recent col- lections. Collected by all earlier ichthyologists who have studied James River fishes, and Burton and Odum (1945:187) took it in 7 out of 28 collections in Craig and Johns creeks. The fallfish may exceed 15 inches in length. In May and early June the male builds a nest by picking up stones in the mouth and placing them in a large pile that sometimes is 5 or 6 feet in diameter and 2 or 3 feet high (See Raney 1940b: 128). Although they bite readily and are caught especially by anglers flyfishing for trout, they are not considered a particularly de- sirable species. The flesh is edible.

Northern Creek Chub Semotilus atromaculatus atromaculatus (Mitchill) A scarce but widely distributed form in the upper third of the James Basin. Found at only 10 out of 57 recently collected stations. The early collectors, Cope (1869:240) and Jordan (1890:111) also found it to be uncommon. Burton and Odum (1945) did not take it in the hcadwaters of the James. This is somewhat of a surprise in- asmuch as the creek chub is a common inhabitant of headwater streams in more northerly drainages, and is also found, although less com- monly, in more southern streams. It reaches a length of more than 10 inches under favorable conditions, but is of little importance except as a boy's fish. In May just above a riffle the male builds a nest that consists of a cup-shaped depression with a long low ridge of sand and gravel that extends upstream often for a distance of -10 feet.

Allegheny Pearl Dace margarimmargarita:(Cope) Listed by Fowler (1945:26) in his distribution table. If this min- now is actually present in the James River Basin it probably will be found in small cold spring streams. It is not uncommon in small tributaries of the Potomac River.

River Chub Nocomis micropogon (Cope) A rather scarce species in the larger streams of the upper third of the James basin where it has been taken in only 9 out of 57 collections. FRESHWATER FISHES 161

It has been taken in company with the Carolina chub, N000711iS lepto- cephalus, but the latter appears to be more common. Cope (1869:240) recognized only one of the two species occurring in the James and recorded it incorrectly under the name Ceratichthys biguttatus. Jordan (1890:110) likewise did not separate the two kinds and re- corded them under the name Hy bopsis kentuckiensis. Burton and Odum (1945:186) found it in the headwater of Johns Creek. The river chub often attains a length of 10 inches, and many are caught on hook and line especially in April and May when they move upstream to spawning areas. The male builds circular nests often two feet in diameter of small pebbles. These nests are a favorite spawning place for other minnows (see Raney, 1940b:130, and 1947b:125).

Carolina Chub N000711iS leptocephalus (Girard) A common species in smaller streams in the mountains and it is especially common and a dominant member of the fish fauna in sand bottomed Piedmont streams. Collected recently at 28 out of 57 sta- tions. The Carolina chub is common in other southeastern river systems and reaches the northern limits of its range in the York and James River systems of Virginia. It is a somewhat smaller species than the river chub, and seldom exceeds 8 inches in length. The nest also consists of a heap of pebbles. A fairly good bait fish. The following key modified after Lachner (1947:207) will facili- tate the identification' of the two species of Nocomis which have often been confused in the past.

la. Snout somewhat pointed anteriorly and longer; in adults in length enters the standard length about 8 times. inserted farther forward; the distance from end of hypural to origin of dorsal when measured forward reaches to or in front of nostril. More scales below ; 5 to 6, average 5.7. Least suborbital width goes into postorbital head 1.7 to 2.0, average less than 2.0 and into the head length 3.5 to 4.2, averages less than 4.0. Pre- orbital into postorbital head 1.0 to 1.1 times. averages less than 1.1. Caudal spot diffuse and faint. Head longer and less deep. Body less deep. Lateral line less decurved. In nuptial males the tubercles are located primarily on the internasal area, on the sides and in front of snout, but do not approach the occiput. River Chub, Nocomis micro pogon (Cope)

lb. Snout somewhat bluntish anteriorly and shorter; in adults its length enters the standard length more than 9 times. Dorsal fin inserted more posteriorly; the distance from end of hypural to origin of dorsal when measured forward reaches the eye. Fewer scales 162 THE JAMES RIVER BASIN

below the lateral line less; 4 to 5, average 4.6. Least suborbital width goes into the postorbital head 1.9 to 2.5, average more than 2.0 and into head length 3.6 to 5.0, averages more than 4.0. Pre- orbital into postorbital head, 1.1 to 1.4 times, averages more than 1.1. Caudal spot round, small, and distinct. Head shorter and deeper. Body deeper, especially just behind the head. Lateral line more decurved. In nuptial males the tubercles are located primarily between the internasal and interorbital regions and also definitely approach the occipital area, and are sometimes developed on the sides and front of snout. Carolina Chub, Nocomis leptocephalus (Girard)

Eastern Blacknose Dace Rhinichthys atratulus atratulus (Hermann) A common and widespread dace in the small streams of the moun- tains and the Piedmont. Captured at 26 out of 57 recently collected stations and recorded by all earlier collectors. Burton and Odum (1945:186) found it at 13 out of 28 stations in Craig and Johns creeks. It is a small minnow that seldom reaches a length of 4 inches in this part of its range. Found in or near riffles, or in pools of small creeks, where it may serve as forage for other larger fishes. They spawn in May. The males do not build nests although each may hold a small territory.

Longnose Dace Rhinichthys cataractae (Valenciennes) This long snouted dace is somewhat less common than the related blacknose dace, and mostly limited to fast riffles in the mountain region. Found in 20 out of 57 places that were recently collected. Cope (1869:227) did not find it and Jordan (1890:111) records it only from Buffalo Creek near Lexington. Fowler (1923b:12) found it in Catawba Creek at Kyle. It does not often penetrate to the very headwaters of the higher mountain streams and Burton and Odum (1945) did not find it in Craig or Johns creeks. Grows somewhat larger than the blacknose dace but seldom exceeds a length of 5 inches. Following recent usage, Fowler (1945:26) records the longnose from the basin as the subspecies cataractae. The writer refrains from following this lead until a thorough study of the speciation of this widespread species is made. It spawns in riffles in May.

Cutlips maxillingua (Le Sueur) Distributed widely but not common in small and medium sized streams in the Piedmont and the mountains although usually not pene- trating to the headwaters of the higher mountain brooks. Captured in FRESHWATER FISHES 163

8 out of 57 recent collections. Recorded by Cope (1869:241), Jordan (1890:109) and Fowler (1923b:12, 1945:31) but not found by Burton and Odum (1945) in any of their 28 collections from Craig or Johns creeks. Adults seldom exceed 5 inches in total length. The male builds a nest in May that consists of a pile of small pebbles (see Raney 1940a:131).

Mountain Redbelly Dace Chrosomus oreas Cope A brilliantly colored small minnow that is found in the small streams in the mountains and to a lesser extent in Piedmont brooks. It has been captured in 12 out of 57 recent collections including the headwaters of the Appomattox River. Neither Cope (1869) or Jordan (1890) took this species although the former described it originally in the same paper, from the specimens taken from the upper Roanoke River system in Virginia where it is an abundant species. Fowler (1918:17, 1923:9 and 1945:25) records it from the basin. It was not taken in Craig or Johns creeks by Burton and Odum (1945). The red- belly is much more common in the Roanoke River system and has also been taken across the mountains in the New River drainage (see Hubbs and Raney, 1944:12), and in the Potomac (Miller, 1946: 207 ). The known range is now extended southward to include the upper Neuse River system of North Carolina where the writer took it on April 1, 1941, in Flat River, and in a tributary of the same stream, a few miles northwest of Timberlake, Person County, North Carolina. Four inches is near the maximum size for this dace. Raney (1947b: 126) has given an account of its spawning behavior. Males were seen holding territories and breeding over Nocomis nests, but are also known to utilize the sand and gravel areas just above rifles for egg laying.

Southern Redside Dace Clinostomus vandoisulus ( Valenciennes) This pretty dace is a common species in both the Piedmont and mountains. Usually it is found in the pools of small or medium sized streams and was taken in 22 out of 57 recent collections. Found by all earlier collectors such as Cope ( 1869: 229), Jordan ( 1890: 111), Fowler (1923a:9, 1923b:9) and Burton and Odum ( 1945: 186 ). The latter found it at 11 out of 28 places in Craig and Johns Creeks. The southern redside dace is a small minnow which rarely reaches a length of five inches. The males and to a lesser extent the females are red sided during June when they are spawning. Fowler (1944:52) recently created a new genus, Rhodopleuriscus, for this species. The characters used are of a type that barely indicate specific separation from the other species in the genus, Clinostomus elongatzts (Kirtland), and the two obviously are closely related. It is 164 THE JAMES RIVER BASIN this writer's opinion that a generic level of differentiation has not been reached.

Eastern Golden Shiner Notemigonus cry soleucas crysoleucas (Mitchill) A quiet water species found to date at only three localities. Jor- dan ( 1890: 115) records it from Spring Creek near Suffolk and Fowler (1923:9) reports three specimens from Midway Mills, Nelson County. Schultz and Reid took 21 specimens in Big Lickinghole Creek west of Goochland. It prefers quiet waters and is expected to occur in !ponds and sluggish weedy creeks of the Coastal Plain. Of value as a forage fish in farm ponds and other impoundments. Some reach 12 inches in length especially in lakes and occasionally are caught by anglers. An important bait fish. Spawns in late spring and summer when the eggs are spread broadcast on aquatic vegetation or other objects.

Attractive Shiner Notropis amoenus (Abbott) This stream-lined shiner has been taken in only 5 out of 57 recent collections. Twice it was captured with the related rosyface shiner, Notropis rub ellus, a species with which it often has been confused. Cope (1869) did not recognize it or may have confused it with Notropis ardens ardens. Jordan (1890:110) reports it from 4 localities including Shingle Creek near Suffolk. He found it to be common in the channels of large streams. Fowler's (1923a:9 and 1923b:10) re- cords of Natropis photo genis amoenus from Midway Mills and Catawba Creek near Kyle probably refer to Notropis rubellus. He (1945:31) reports Notropis rubellus amoenus in his distribution list. The recent collections that include this species were those by Schultz and Reid from Chickahominy River north of Richmond, Big Licking- hole Creek west of Goochland, and Byrd Creek east of Columbia. The writer has taken it in a tributary of the Appomattox River, 14 miles northeast of Appomattox, and in South Anna River, 5 miles northeast of Charlottesville. Occasionally these two species may be found together at a given locality but they generally show a habitat preference. Notropis amoenus likes the pools and channels while rubellus is more often found at the base or at the head of riffles. The former is more com- mon on the Piedmont and even on the Coastal Plain region. Al- though rubellus is also found in the Piedmont, it is fairly common in the mountains but apparently does not occur on the Coastal Plain. They probably spawn at different times although there is little avail- able data on the spawning behavior of amoenus.. It retains its pale coloration and most likely spawns in early summer. On the other hand rube/ins breeds in May in riffles or over nests of other fishes, FRESHWATER FISHES 165 and at this time the head and body are suffused with red (Raney, 1940c:361). The common hybrid combination, Notropis cornutus cornutus X Notropis rubellus, may at times be confused with Notropis amoenus.

Rosyface Shiner Notropis rztbellus (Agassiz) A rather widely distributed shiner found mostly in the upper Piedmont and in mountain streams. It prefers riffles and spots in pools near an entering current. Not typically found in small brooks. Cap- tured at 10 out of 57 recent stations. Cope (1869) did not record it but perhaps confused it with Notropis ardens. Jordan (1890:110) reports it and gives some of the characters used to identify it. Fow- ler (1923:11) had this form but confused it with amoenus and dis- poses of it in his most recent work (1945:31) by considering it a sub- species, Notropis rztbellus amoenus. Maximum size is about 4 inches. Fowler (1923b:11) recorded Notropis atherinoides from the head- waters of the James. This record is regarded as extremely doubtful since atherinoides is not a headwater species in any part of its known range and is not known nor expected east of the Appalachians. Per- haps this record was based on Notropis rubellzts, N. ardens, or N. amoenus. To aid in the identification of these species a key is given below. This comparison was drawn largely from adult specimens of both species taken together below a dam in Rapidan Creek, Madison Mills, Madison County, Virginia, but other specimens collected at other places along the Atlantic Coast were also used. la. Body compressed; the lower two-thirds of sides almost vertical. Body thinner and deeper; the greatest body thickness at the level of the origin of the dorsal fin, when measured downward from the dorsal origin, does not reach the lateral line. Snout, as viewed from above, is broadly U shaped. Extended anal fin is falcate on posterior border. Depressed dorsal fin; the posterior tip of the first ray extends beyond the tip of the last ray. An- terior tip of premaxillary smaller; it gradually increases in width until the symphysis is reached. Anal rays usually 11, often 10, and sometimes 12 (counts were 10 in 13 specimens, 11 in 41 specimens, 12 in 5 specimens). Scales on the back before dorsal fin crowded, indistinct in outline, and greater in number. General body color light. Males, in spawning condition, pale colored. Both species have dark lips; in amoenus they appear darker be- cause the head, in contrast, is much lighter. Top of head with rather evenly distributed melanophores; heart-shaped mark on posterior head is indistinct. Melanophores form a band on the an-, 166 THE JAMES RIVER BASIN

tenor tip of the dentary and connect with a median dark stripe on the anterior half of the gular region. Dark lateral body stripe is poorly if at all developed on anterior half of body; it begins to fade near middle of the dorsal fin and is almost absent at the origin of the dorsal fin, and in front of the dorsal is represented by fine, short, vertical dark lines. Few melanophores extend onto the base of the caudal fin (beyond the dark lateral body stripe). Mid- dorsal dark streak behind dorsal fin is faint and diffuse. . . . Attractive Shiner, Notropis amoenus (Abbott) lb. Body subterete; the sides are definitely rounded. Body thicker and less deep; its greatest thickness at the level of dorsal origin, when measured downward, reaches to or below the lateral line. Snout V-shaped. Extended anal fin is straight on posterior bor- der. Depressed dorsal fin; the posterior tip of the first dorsal ray reaches only to the tip of the last ray. Premaxillary massive; the bone increases abruptly in width near the symphysis. Anal rays usually 10, but occasionally 11 (counts were ten in 27 specimens, 11 in 3 specimens). Scales before dorsal more distinct, less crowded, and fewer (the extremes of rubellus approach amoenus in this character). General body color dark. Spawning males red on head and body. Lips dark, but only slightly more so than the rest of the head. Dark heart-shaped mark on posterior part of head is pronounced. Dark on anterior dentary not expanded, and does not connect with the few scattered melanophores in the gular region. Dark lateral stripe fades somewhat on the anterior half of the body but reaches the posterior border of the opercle. Many melanophores extend beyond the posterior tip of the dark lateral stripe onto the base of the caudal fin. Mid-dorsal streak behind dorsal fin dark and discrete . . . Rosyface Shiner, Notropis rubellus (Agassiz)

Eastern Rosefin Shiner Notropis ardens ardens (Cope) This slender shiner was identified by Dr. Carl L. Hubbs and re- ported by Burton and Odum (1945:187) from 3 out of 12 collections in Craig Creek in the upper James River system. Cope (1869:229) reports that he saw this species only in the Roanoke although a year later (1870:460) he said that he took it in both the Roanoke and James rivers. However, Cope did not report Notropis rubellus or Notropis amoenus from the James and may have confused them with N. ardens. The rosefin is common in the Roanoke River system where its habits were studied by Raney (1947:126). A small shiner that seldom is longer than 3 inches. The habitat is usually moderately fast water. FRESHWATER FISHES 167

Eastern Common Shiner Notropis cornutus cornutus ( Mitchill) A common species in the Piedmont and mountains which was captured at 32 out of 57 recently seined localities. Occurs usually in moderate sized and small streams. Burton and Odum (1945:186) found it at 10 out of 28 localities in Craig and Johns creeks. All early col- lectors also report this species. However, Fowler (1923a:9, 1923b:10, and 1945:29) has identified it as a subspecies, N. cornutus cerasinus, or a full species N. cerasinus. However, the true N. cerasinus is found in the Roanoke River system where it, along with Notropis albeolus, replaces Notropis cornutus. Of some importance as a bait fish although it does not usually reach a length of more than 7 or 8 inches. In May, - when the red nuptial color is found on the sides of the body as well as the fins of the male, it is often called redfin shiner. Prefers moving water and resorts to the riffles or to the nests of other fishes to spawn. Raney (1940:8) gave an account of its reproductive activities. The common shiner hybridizes readily with several other species of Notropis as well as with other genera of minnows. Most of these hybrids are a result of a fortuitous meeting of egg and sperm as two or more species spawn at the same time at a given place, usually over the nest of a river chub, a fallfish, or a cutups minnow. One of the most common hybrids is that between the common shiner and rosyface shiner, Notropis cornutus cornutus X Notropis rubellus. This hybrid was once described as a new species from the Shenandoah River, Vir- ginia and was named Notropis macdonaldi Jordan and Jenkins (see Jordan 1890:110). The common shiner less frequently crosses with the mountain redbelly dace, Notropis cornutus cornutus X Chrosomus oreas, and the redside dace, Notropis cornutus cornutus X Clinostomus vandoisulus. Other hybrids occur less frequently in the James. See Raney (1940c:362 and 1947b:129) for additional information and fur- ther references to this interesting subject.

Southern Spottail Shiner Notropis budsonizts saludanus ( Jordan and Brayton) Although this shiner has only been taken at 6 out of 57 recent stations it is probably fairly common in river channels. It ranges from the mountains where Burton and Odum (1945:187) record it from only 2 out of 28 collecting stations (under the subspecific name amarus) to the Coastal Plain where it was recorded by Jordan (1890:110 and 115) from Swift Creek near Petersburg and Shingle Creek near Suf- folk. Cope did not report it. Fowler (1923:91) took it at Midway Mills. The spottail is a small species, and usually measures less than 5 inches in length. It spawns in riffles in early May. On the basis of the comparatively few specimens available for analysis the spottail of the James has been assigned tentatively to the 168 THE JAMES RIVER BASIN southeastern subspecies saludanus, which seems to differ from the more northerly subspecies amarus, mostly in having a longer head and a more inferior mouth. Perhaps the latter subspecies is present in the upper James. Considerable additional specimens and study are needed to solve this problem.

Satinfin Shiner Notropis analostanus (Girard) A pretty shiner that is found mostly in the Piedmont and in the larger mountain streams. It may also occur more sparingly on the Coastal Plain. Taken in 13 out of 57 recent collections. It was re- ported by Cope (1869:229), Jordan (1890:110), and Fowler (1923b: 10, 1945:28) but was not captured in the mountain headwaters by Bur- ton and Odum (1945). Often taken in the quieter waters below mill dams. It usually lays the eggs on the surface of logs but when not available they utilize the surface of stones. Spawning occurs in June. At this time the fins, especially of the male, become satiny white. Jordan (1890:115) records the snowy shiner, Notropis niveus (Cope), from Shingle and Spring creeks near Suffolk. Some of his material from these localities, deposited in the U. S. National Museum, was reexamined recently and proved to be Notropis analostanus. Notropis niveus is normally a southeastern species and while it may reach the northern limit of its range in the Dismal Swamp area its certain presence there needs to be demonstrated. Although N. niveus is obviously related to N. analostanus it differs greatly in general ap- pearance. Its body is slenderer, the snout bluntish, the mouth inferior, and it has 8 anal rays while N. analostanus has 9.

Northern Swallowtail Shiner Notropis procne procne (Cope) A widespread species found especially in the larger streams in each of the three physiographic regions of the James. Captured at 12 out of 57 stations worked recently. Taken by earlier ichthyologists except by Burton and Odum (1945). It apparently avoids the moun- tain headwater streams but shows a liking for sand bottom and is well adjusted to life in Piedmont streams. The studies of Hubbs and Raney (1947:3) and Raney (1947a:103) show that the subspecies inhabiting the James is procne. Notropis procne longiceps ranges through the Roanoke system and more southerly drainages. Cope (1869:231) based the original description of longiceps largely on Roanoke River speci- mens although he also states that he found it in the James in Giles County. His records of longiceps in the James have been repeated by Fowler (1923b:10 and 1945:27) but its occurrence needs to be demon- strated. Few swallowtail shiners grow to be more than 2 Vz inches long. Like most other minnows they are of some importance as forage fish FRESHWATER FISHES 169 but are not a particularly good bait fish. Their interesting spawning behavior which occurs over sand and gravel in moving water in mid- June is described by Raney (1947a: 105).

Eastern Silvery Minnow Hy bognathus nuchalis regius Girard Apparently a scarce species since it is represented in only two out of 57 recent collections. Perhaps it is more common in the deeper waters of river channels where it was found by Jordan (1890:109). He captured it in James River at Lick Run, east of Clifton Forge, and in Shingle and Spring creeks near Suffolk. Fowler (1923:9) reported it from Midway Mills. The recent collections in the U. S. National Museum made by Schultz and Reid are from Big Lickinghole Creek west of Goochland, and from Byrd Creek, east of Columbia. A small minnow that rarely exceeds 5 inches in length. Raney (1939:674) has given an account of its spawning which takes place in early spring.

Stoneroller Minnow Campostoma anomalum (Rafincsque) subsp. Fairly common in the mountain headwaters where Burton and Odum (1945:186) found it at 10 out of 28 stations. In 57 other recent collections it was captured at only 5 places. Neither Cope (1869) or Jordan (1890) reported it, but Fowler (1923:9, 1945:25) recorded it from Midway Mills. The suspecies present in the James is probably not Campostoma anomalum anomalum since the lateral line scale count and the number of scales around the body just anterior to the dorsal fin are much too high. In these respects it is similar to the population of Campostoma in the Roanoke River system. However the allocation of a subspecific name is deferred awaiting the final results of a study now being made. Seven or 8 inches is near maximum size for this species in the James while most do not exceed 6. It is of some im- portance as a forage fish. In feeding it scrapes algae and other small organisms from rocks and its rather general scarcity in the sandy Pied- mont region may be attributed in part to this fact. The eggs are de- posited in cup-like depressions made by the male usually in an n roa just above a riffle. Raney (1947b:125) gives some data on the hybridi- zation of the stoneroller as well as other species in Virginia.

Bluntnose Minnow Hy borhynchus notatus Rafinesque This small bottom loving minnow is generally scarce except locally. East of the Appalachians the James River is apparently the southern limit of its range. Dr. Lincoln C. Pettit found it in 5 out of 19 places in the vicinity of Lexington. Burton and Odum (1945:187) took it at only one out of 28 stations in Craig and Johns creeks. The writer caught it at only two places; in Buffalo Creek southwest of 170 THE JAMES RIVER BASIN

Lexington, and in a tributary of the James River near Eagle Rock. None of the early ichthyologists reported it. The bluntriose rarely exceeds 4 inches in length. The male guards a nest hollowed under a flat stone or board and the eggs are fastened to the underside. An important forage and bait fish in more northerly waters.

NORTH AMERICAN CATFISH FAMILY AMEIURIDAE Six members of this family are known from the James River. Four species, 2 large , and 2 bullheads, are of commercial importance and are more commonly found in the lower river and its larger tribu- taries. Menzel (1945:364) who has studied the catfish industry of Virginia states that they are more important than the shad or striped bass in the James and its tributaries. Two species of small madtoms are also known. One, Schilbeodes mollis, is a Coastal Plain form and the other, S. m. marginatus, is found in riffles of medium and small streams. In catfishes in general, the male builds a nest consisting of a circular cleared area on the bottom, a hole in a bank, or a hollow under a stone or log, usually in June. The eggs are guarded and young are accom- panied by a parent for part of their first summer of life. See Breder (1935:143) for an account of the spawning habits of some members of this family.

White Catfish lctalurus catus (Linnaeus) An important commercial fish in the James River and its larger tributaries. In appearance the white cat resembles the yellow bullhead, natalis, but differs most notably externally in having the tail shallowly forked. Cope (1869:487) reports it as Ameiurus lynx, Jordan (1890:109) as Ameiurus albidus, and Fowler (1923:9 and 1945: 32) as Ameiurus catus and lctalurus catus catus. According to Menzel (1945:365) this catfish reaches a size of somewhat over two pounds. They eat fishes, crustaceans, and aquatic insects. Most of them are taken commercially in cylindrical pots that are baited with dead fish or "bean cakes".

Southern Channel Catfish lctalurus lacustris punctatus (Rafinesque) This large catfish is native to the drainage system and was introduced into the James River as well as some other eastern rivers. It is now present in considerable numbers and is of commercial importance. Usually it is most often taken in or near the channels of rivers and in the moving water of power outlets below dams. It has been found as far Upstream as Midway Mills by Fowler (1923a:9) and by the writer in South Rivanna River, 5 miles northeast of Charlottes- ville. It reaches a larger size than the white catfish. Specimens of 5 FRESHWATER FISHES 171 pounds are not uncommon and it may exceed 20 pounds in weight. The deeply forked tail and the usual presence of dark spots on the sides of the body are characters that separate it from the white cat- fish. An omnivorous feeder but insects and fish serve as staple foods.

Northern Brown Bullhead Ameiztrus nebulosus nebulosus (Le Sueur) Common in the river and in sluggish streams. It more often makes its way up into the smaller Piedmont and mountain streams than does the yellow bullhead. In the lower James River it is taken commercially along with the yellow bullhead and the 2 larger catfishes. Recorded by Cope (1869:242); Jordan (1890:109), Fowler (1923b:9 and 1945: 32) and Menzel (1945:364). Most of the larger brown bullheads are 12 to 15 inches long.

Southeastern Yellow Bullhead Ameiztrus natalis erebennus Jordan A common species in the lower river. It contributes considerable to the commercial fishery according to Menzel (1945:364), but by most fishermen it is not usually recognized as different from the brown bullhead. Jordan (1890:114) found it in Jericho Canal and Shingle Creek near Suffolk and says it is the most common catfish in Dismal Swamp. The subspecies erebennus is tentatively applied to this form of yellow bullhead pending a badly needed study on the systematics of eastern bullheads. Most are under 15 inches in length.

Tadpole Madtom Schilbeodes mollis (Hermann) Fowler (1945:32) records it in his distribution table under the name S. gyrinus but the earlier workers and the writer have not seen it from the James. It is certainly expected to be present on the Coastal Plain inasmuch as it is found both to the north and to the south. To be looked for in sluggish weedy areas with muddy bottom. A small madtom that rarely is more than five inches long. Both the common eastern madtom and this species have a poison gland at the base of the pectoral fin. The venom penetrates through a puncture of the skin caused by the pectoral spine and results in a rather severe pain. The eggs are laid during late June in a clump in under a stone, or board, and have even been found in old tin cans.

Common Eastern Madtom Schilbeodes marginatus marginatus (Baird) Common in riffles in the upper Piedmont and mountain region where it has been taken in 19 out of 57 recently made collections. Cope 172 THE JAMES RIVER BASIN

(1869: 237 ) , Jordan (1890:104) record it under the name S. insignis (Mitchill). Hubbs and Raney (1944) have discussed the systematics of this madtom and have given detailed records of its distribution as well as a map showing its range. Its nest is a depression made by the male under a stone in moving water. Seldom attains a length of more than 7 inches. Of some value as food for the smallmouth bass, and it often is used as a bait fish in angling for this species.

MUDMINNOW FAMILY UMBRIDAE Eastern Mudminnow Umbra pygmaea (DeKay) A Coastal Plain species which is probably not uncommon in the sluggish weedy habitats in the lower James. Reported by Cope (1869: 242) from near Richmond, by Jordan (1890:116) in Jericho Ditch near Suffolk, and by Fowler (1923b:13 and 1945:33) from the lower James. A small, dark, pike-like fish with horizontal light and dark stripes along the sides. Four to five inches is its maximum size. Its eggs are de- posited in early spring. A fair bait fish.

PIKE FAMILY ESOCIDAE Only two members of this family of game fishes are found in the James. Both prefer quiet waters and are largely carnivorous, feeding on fishes, crayfishes, insects, and other larger aquatic animals. Bulldog Pickerel americanus Gmelin This small redfin species is probably fairly common in the Coastal Plain although actual records of its presence are few. Cope (1869:242) reports taking two kinds of Esox near Richmond, one of which prob- ably was this species. Jordan (1890:116) found it commonly in Jericho and Shingle creeks near Suffolk, and there are other recently collected specimens in the U. S. National Museum taken in the Dismal Swamp region. Most specimens are less than 10 to 12 inches, and because of its small size it is of little importance as a food fish. It is often caught on hook and line by boys. Chain Pickerel Esox niger Le Sueur This species is larger and perhaps more common than the bulldog pickerel. It also is more widespread and has been found in such diverse habitats as pools in mountain streams to brackish water in Chesapeake Bay. Burton and Odum (1945:186) took it in 8 out of 28 collections in the Craig and Johns creeks. Also recorded by Cope (1869:242), Jordan (1890:111) and Fowler (1928:134). Recently collected by Schultz and Reid in Tuckahoe Creek above Gayton, and by the writer in a tributary of Slate River, 5 miles southwest of Rockingham. Of FRESHWATER FISHES 173 considerable importance as a sport fish and is sometimes seen in the commercial market. Reaches a length of two feet and a weight of 4 to 5 pounds in the James. Spawning occurs in the early spring. The eggs are broadcast over vegetation in quiet water and are not deposited in jelly like strings, reports in the literature notwithstanding.

FRESHWATER EEL FAMILY ANGUILLIDAE American Eel Anguilla bostoniensis (Le Sueur) A common species in the lower river and its tributaries. Although Midway Mills in Nelson County is as far upstream as it has been re- corded by Fowler (1923a:10), probably no medium sized stream in the James system is without the eel at some season or other. Pools below mill dams are a favorite haunt. Occasionally they are found landlocked in isolated ponds where they have been introduced. The adults return to the sea to spawn. They move down stream in autumn and migrate to the spawning ground south of Bermuda. The young eels enter all the coastal streams of Virginia and are found in great abundance near the mouths of rivers in early spring. At this time they are about 4 to 6 inches in length. Adults consistently attain a length of 3 to 4 feet and may reach 6 feet. They are of considerable commercial im- portance and are captured in eel pots set especially for them and also by hook and line.

KILLIFISH FAMILY CYPRINODONTIDAE A big family of small egg laying fishes that are widely distributed along the coast in salt and brackish water. Several species regularly enter freshwater and some like the eastern banded killifish, Fundulus diaphanus diaphanus, and the mummichog, Fundulus heteroclitus mac- rotlepidotus, often get landlocked and thrive in inland lakes and ponds. A number of other cyprinodonts are chiefly marine in the latitude of Virginia but are expected to be found at one time or another in the freshwaters of the lower James River. They are the shecpshead killi- fish, Cyprinodon variegatus Lacepede; rainwater fish, Leucania parva (Baird and Girard); May killifish, Fundulus majalis (Walbaun); and spotfin killifish, Fundulus luciae (Baird). Hildebrand and Schroeder (1928:134) have given a general account of each of the above. They are mostly gregarious species of shoal waters, are of some importance as bait and forage fishes, and perhaps also aid in mosquito control.

Mummichog Fundulus heteroclitus macrolepidotus (Walbaun) Occasionally found in the freshwaters of the lower James River although its typical habitat is brackish or saltwater. According to Hildebrand and Schroeder (1928:138) it is very common along the shore of Chesapeake Bay. 174 THE JAMES RIVER BASIN

Eastern Banded Killifish Fundulus diaphanus diaphanus (Le Sueur) Found commonly in Chesapeake Bay and in the lower James River. Reported by Cope (1869:242) near Richmond and Jordan (1890:116) from Shingle Creek near Suffolk.

LIVEBEARER FAMILY POECILIIDAE Eastern Mosquito Fish Gambusia affinis holbrookii (Girard) Found on the Coastal Plain in quiet waters. Jordan (1890:116) found it to be very common in the Suffolk area, and Schultz and Reid took it recently in Tuckahoe Creek above Gayton. Related to the killifishes but it bears living young. The modified anal fin serves as an intromittent organ. A small species that does not often exceed 3 inches in length but which is of considerable importance because of its destruction of mosquito larvae. Distributed along the Atlantic coast as far north as .

CAVE FISH FAMILY AMBLYOPSIDAE Rice Fish Chlogaster cornutus Agassiz A small species related to the killifishes found in quiet waters along the Coastal Plain. It is recorded here as occurring in the James River system upon the strength of Jordan's (1890:116) records of Chlogaster avitus Jordan and Jenkins, from the Jericho Canal near Suffolk where he said it was abundant.

PIRATEPERCH FAMILY APHREDODERIDAE Eastern Pirateperch Aphredoderus sayanus say anus (Gilhams) A small aggressive fish found in sluggish waters of the Coastal Plain and less often in the Piedmont. Jordan (1890:16) captured it in Jericho Canal near Suffolk. Schultz and Reid found it in Tuckahoe Creek above Gayton, and the writer took it as far upstream as Willis River, 10 miles west of Cumberland, Buckingham County. A dark colored fish that is characterized by having the anus located far for- ward in the throat region. Rarely exceeds 5 inches in length. An in- teresting aquarium fish.

BASS FAMILY SERRANIDAE Striped Bass Roccus saxatilis (Walbaun) An important game and food fish that lives in the sea but which enters the river during its spawning run in May. There are many records of its occurrence in the lower James and it has been recorded FRESHWATER FISHES 175 by all early ichthyologists except Cope. Hildebrand and Schroeder (1928:247) have given a general account of its distribution and eco- nomics in Chesapeake Bay.

White Perch Morone americana (Gmelin) Found both in Chesapeake Bay and in lower James River efe it seems to be equally at home in brackish or freshwater. Jordan (1890:117) found it commonly in Shingle and Spring creeks near Suf- folk, and Menzel (1945:367) notes its occurrence in a catfish stomach taken in the lower James River. It is of considerable commercial im- portance and is frequently netted in large numbers. Spawning occurs from April to June. Attains a length of 12 inches and weight of about 1.5 pounds.

PERCH FAMILY PERCIDAE Eight kinds of small fishes belonging to this family are known from the James River. They are classified into 7 species, and one species, Boleosoma nigrum, has 2 subspecies. Many are eaten by larger fishes especially by bass. The yellow perch, Perca flavescens (Mitchill), a Coastal Plain species is not actually recorded from the James River but it un- doubtedly occurs in the lower reaches of this system. It is known from the Coastal Plain waters both to the north and south of the James. Hildebrand and Schroeder (1928:236) gave several records of its occurrence in the freshwater tributaries of Chesapeake Bay. An important pan fish.

Stripeback Darter Hadropterus notogrammus Raney and Hubbs Fairly common and widespread in the mountain and Piedmont streams. Raney and Hubbs (1948:4) have recently described this darter and have mapped its distribution. It reaches the southern limit of range in the James River. Those from the upper James are extreme in several characters and probably represent a different subspecies, a matter that will be settled as additional specimens become available for study. Usually found in small and medium streams. Attains a length of about three inches.

Shielded Darter Hadropterus peltatus peltatus (Stauffer) This darter is much less common than was formerly supposed. Most early records are based on Hadropterus notogrammus as was pointed out by Raney and Hubbs (1948:3). Jordan (1890:113) record from Swift Creek near Petersburg (specimen reexamined) and 176 THE JAMES RIVER BASIN a recent single specimen taken by Schultz and Reid in Chickahominy River are the only good records' of peitatus known to me from the James. Jordan's (1890:117) specimen from Dismal Swamp, Shingle Creek near Suffolk, was also reexamined and found to be peitatus. Raney and Suttkus (1948) have recently made a study of the sub- specific variation in this species. The James River specimens are the southernmost examples of the typical subspecies. Fowler's (1945:37) record of Hadropterus peltatus crassus (Jordan and Brayton) from both the James and Roanoke systems is in error. Hadropterus crassus is a southern species. The shielded darter is small species that rarely attains a length of four inches. It spawns in the riffles in May and thereafter is more at home in pools.

Tessellated Johnny Darter Boleosoma nigrum olmstedi (Storer) This common Coastal Plain form has small body scales and also has scales on the cheek, opercle, nape, and breast. It prefers quiet waters and has been found nesting by Neil D. Richmond in the inter- tidal area of Chickahominy River, in New Kent Co. Hubbs and Raney (1946:19) have given the characters, and Raney and Lachner (1943: 229) have studied the growth rate of this species. In May the eggs are attached to the underside of a flat rock, shell, or similar object.

James River Johnny Darter Boleosoma nigrum (Rafinesque) subsp. The mountain streams and at least part of the Piedmont brooks are populated by a Johnny darter that is at least subspecifically dif- ferentiated from olmstedi. It differs in having a lower scale count (lateral line scales 35 to 44, mean about 39, in contrast with 41 to 54, mean 48 in olmstedi), a low dorsal spine and soft ray count, and a low anal ray count. Furthermore it is a much smaller species and seldom reaches 2 inches in length while olmstedi is often 3 inches long. The application of the proper subspecific name to this darter must await the results of studies, now in progress, of the Johnny darters through- out their range. Taken in 16 out of 57 recent collections made in the Piedmont and mountains. Fowler's (1945:38) identification of this species with Boleosoma effulgens (Girard) (equals Boleosoma nigrum effulgens) is unacceptable inasmuch as the characters of the effulgens of the Potomac tributaries are considerably different.

Longfin Darter Boleosoma longimanum (Jordan) A large species that is probably found only in the James River system. It occurs in the same geographic area With Boleosoma nigrum subsp. but seems to prefer the faster water. It penetrates to the moun- tains headwaters where it was captured by Burton and Odum (1945: FRESHWATER FISHES 177

186) at 4 out of 16 localities in Johns Creek. Taken in 17 out of 57 other recent collections in the Piedmont and mountain regions. Fowler (1918:18) reports it from Midway Mills. but later (1945:37) records it as Cottogaster longimanus Agassiz or Putnam, from both the Potomac (almost certainly an error) and from the James rivers. His (1945:37) record of Cottogaster podostemone ( Jordan and Jenkins) from the James is also thought to be an error since this species, Boleosoma podostemone, is probably limited to the Roanoke River system. Raney and Lochner (1943:235) have studied the growth of the long fin darter. It rarely exceeds 3 inches in length and is short lived. This species can easily be separated from the other two Boleosoma in the area by the presence of two anal spines (rather than one) and by the darker coloration. All members of this genus, as well as other darters, serve as important items in the diet of many game fishes.

Glassy Darter loa vitrea (Cope) This elongate light colored darter is almost strictly limited to sand bottom in both the Coastal Plain and Piedmont. Recorded by Jordan (1890:117) as common in sandy places in Shingle Creek near Suffolk. Schultz and Reid took a single specimen at Big Lickinghole Creek west of Goochland, and in Byrd Creek east of Columbia. The writer col- lected a series in a tributary of 'Willis River, 10 miles west of Cumber- land, and in South Rivanna River, 5 miles northeast of Charlottesville. Is seldom more than 3 inches long.

Fantail Darter Poecilichthys flabellaris (Rafinesque) Widespread in mountain and Piedmont streams. Usually found in or close to riffles. Burton and Odum (1945:187) collected it in 2 out of 12 collections from Craig Creek. The other ichthyologists have also recorded it from the James system and it was taken in 29 out of 57 recent collections. Very likely it is the most common darter in the upper James. However, collectors have undoubtedly sampled its riffle habitat better than that of some of the other darters. The fantail is of some importance as forage for bass. The eggs are laid under a flat stone in the late spring and are guarded by the male.

Swamp Darter Hololepis fusiformis Girard Known from the James River system only from 11 specimens taken by Schultz and Reid in the Piedmont in Big Lickinghole Creek, west of Goochland. It prefers quiet waters and perhaps will be found on the Coastal Plain. When Hubbs and Cannon (1935) revised the genus Hololepis, no specimens were available from Virginia. However, the above specimens from Big Lickinghole Creek and additional series re- cently taken from other localities on the Coastal Plain in Virginia will 178 THE JAMES RIVER BASIN make it possible to work out the one or several subspecies that inhabit this region. The swamp darter is a small bottom dweller that rarely exceeds 2.5 or 3 inches in length.

SUNFISH FAMILY This family, which includes the basses and sunfishes, is the second largest in the James River Basin. It is represented by 12 species, 2 of which, the smallmouth bass and northern rock bass, were introduced. It is highly probable that another, the northern blackbanded sunfish, Mesogonistius chaetodon chaetodon (Baird), will also be found on the Coastal Plain since it has been taken both to the north and south of the James. The white crappie, Pomoxis annularis Rafinesque, is also prob- ably present, and the green sunfish, Lepomis cyanellus Rafinesque, may also be present, both as introductions. The green sunfish is now estab- lished in several other Chesapeake Bay tributaries such as the Rappa- hannock and Potomac rivers. Many of the larger species are fine game fishes and practically all except the small Enneacanthus and A canthar cus are good pan fish. All are carnivorous and eat such animals as fishes, crayfishes, and insects. A nest that consists of a cleared area on the stream bottom is swept off by the male. Their interesting spawning habits have been sum- marized by Breder (1936:1). Hybrids are not uncommon between the several species in the genus Lepomis and occasional intergeneric hybrids also are found.

Northern Smallmouth Bass Micro pterus dolomieu dolomieu Lacepede An important game species which was introduced from the Ohio River. Cope (1869) did not observe it in the upper James and it is probable that it was planted after his explorations. By 1890 it was established and Jordan took it in several headwater streams. It is now rather widespread in the larger streams of the upper Piedmont and mountains. A pool and riffle habitat is preferred in contrast to the largemouth bass which likes weedy muddy ponds, backwaters, or slug- gish rivers. It reaches a length of more than 20 inches and a weight somewhat greater than 5 pounds under favorable conditions. The in- teresting nesting habits and other biological information are summarized by Hubbs and Bailey (1938 and 1940). The smallmouth subsists largely on fishes and crayfishes.

Northern Largemouth Bass Micro pterus salmoides salmoides (Lacepede) A highly regarded game fish which may not be native to the James River. It is common southward along the Coastal Plain, and FRESHWATER FISHES 179 has been stocked rather widely in similar situations to the north. Also found in Piedmont rivers and ponds where it prefers quiet, weedy habitats. The largemouth was present in the James River at least as early as 1869 when it was reported by Cope and it was also recorded by Jordan (1890:112). A weight of 8 to 10 pounds is not unusual under favorable conditions. The male hollows out a nest in the silty stream bottom and guards the eggs. Fishes, crayfishes, and other large aquatic animals serve as food.

Warmouth Chaenobryttus coronarius (Bartram) This sunfish is a characteristic species of weedy, sluggish waters of the Coastal Plain. Known definitely only from Cope's (1869:225) record under the name Lepomis gillii -Cope. His specimen was cap- tured in Tuckahoe Creek near Richmond. This record was again re- ported by Fowler (1923b:13) and is apparently Fowler's (1945:41) basis for inclusion in his distribution table. Although it is highly prized as a pan fish, it seldom weighs more than 1.5 or 2 pounds. A carnivor- ous species that devours other fishes, insects and other large animals.

Pumpkinseed Lepomis gibbosus (Linnaeus) A common and generally distributed sunfish of the Piedmont, espe- cially in mill ponds, and is found also in the sluggish waters of the Coastal Plain. Jordan (1890:112) found it to be common especially in the lower James and he took it as far upstream as Clifton Forge. In recent collections it was taken in only 4 out of 55 collections. How- ever, the stations seined were usually not the typical habitat of this sunfish. Although it rarely exceeds 7 or 8 inches in length it is of some importance as a pan fish. Eats insects and other small animals. Although Jordan's (1890:117) specimens of Lepomis holbrooki (Cuv. and Val.) [equals Lepomis microlophus (Gunther)] from Dis- mal Swamp have not been found for reexamination, it is extremely doubtful if this species ranges that far north. See Hubbs (1932:7) for an account of this southern representative of Lepomis gibbosus.

Common Bluegill Lepomis macrochirus macrochirus Rafinesque Although only one record is available this sunfish is probably not uncommon along the Coastal Plain and in ponds of the Piedmont. Schultz and Reid found it in Tuckahoe Creek above Gayton. A com- bination of the common bluegill and the largemouth bass have proved a desirable pair in many southern farm ponds. A good pan fish that rarely exceeds 8 or 9 inches in length. 180 THE JAMES RIVER BASIN

Yellowbelly Sunfish Lepomis auritus (Linnaeus) The most common and widespread of the sunfishes in the James.. Found both in streams, where it often occurs in the faster waters, and in sluggish rivers and ponds. In the latter habitat it reaches a large size and specimens of from 9 to 10 inches are caught occasionally. The adult Males get a redbelly in the early summer during the spawning season and are often known locally as "robins". Recorded by Cope (1869:224), Jordan (1890:112), and Fowler (1923a:10; 1923b:13; 1945: 42). Richmond (1940:329) gave an interesting account of their nest- ing habits in the tidal zone of the Chickahominy River. In the head- waters of Craig Creek, Burton and Odum (1945:187) found it in one out of 12 collections. Also taken in 9 out of 57 recent collections.

Banded Sunfish Enneacanthus obesus (Girard) A small sunfish which is closely related to the bluespot sunfish. Both are found together on the Coastal Plain. Recorded by Cope (1869:219) from Tuckahoe Creek near Richmond (as E. guttatus), by Jordan (1890:116) as common in Jericho Canal near Suffolk, and by Fowler (1945: 41). It does not often exceed 3.5 inches in length. An attractive sunfish that lives well in an aquarium.

Bluespot Sunfish Enneacanthus gloriosus (Holbrook) Another small sunfish that is similar in appearance to the banded sunfish. It also occurs on the Coastal Plain but has not been reported in recent years from the James. It was collected in Tuckahoe Creek near Richmond by Cope (1869: 217 ) who called it Hemioplites simu- lans. Jordan (1890:117) found it to be scarce in Shingle Creek near Suffolk, and Fowler (1923b:13) records it from the lower James. It rarely exceeds 4 inches in length and makes a fair aquarium fish.

Northern Rock Bass Ambloplites rupestris rupestris (Rafinesque) An introduced species that is now scattered throughout the upper James River system. It appears to be somewhat more common in the upper Piedmont and in the mountains. Burton and Odum ( 1945: 186) found it in 6 out of 28 collections. Fowler (1923:13) reports it from the lower James. However, it has been collected only once in 57 recent collections. A fair pan fish that is usually found in the same habitat as the northern smallmouth bass. It occasionally reaches a length of 10 inches and a weight of 2 pounds. There is a remote possibility that another rock bass, Ambloplites eawifrons Cope, may be found in the James River. This rare species is now known with certainty only from the headwaters of the Roanoke FRESHWATER FISHES 181

River in Montgomery County, Virginia. Cope (1869:277) first de- scribed it and Jordan (1890:123) took 2 specimens and emphasized its main structural features. A large adult, 175 mm. in standard length, was taken June 21, 1946, by E. A. Lachner, R. D. Ross, and the writer, in the Roanoke River, 2 miles north of Alleghany Springs. It is a much different fish than Ambloplites rupestris rupestris which is also present in the Roanoke River, apparently as an introduction. The Roanoke rock bass is a dark fish with narrow and slightly irregular light stripes along the side of the body. These extend high on the back and are also present on the dorsal fin. The cheeks and opercles are marked by discrete round light spots. Its main structural difference lies in the very small and at times embedded scales on the cheek and opercle. The breast scales are also very small and more or less embedded. Fowler (1945:41) recognizes the Roanoke Rock Bass a sub- species, Ambloplites rupestris cavifrons Cope and (1945:253) created a new genus, Bartramiolus, for the southern rock bass. The latter is a Gulf Coast form which the writer and others recognize as a good sub- species, Ambloplites rupestris ariomnrus Viosca. The latter subspecies differs mostly in having a larger eye but was found by the studies of Hubbs reported in Hubbs and Lagler (1947:94) to intergrade with the northern subspecies, in southeastern Missouri and eastern Arkansas.

Flier Centrarchus macro pterus (Lacepede) A Coastal Plain species whose distribution in the lower James is as yet unknown. Jordan (1899:116) claimed it was common in Jericho Canal near Suffolk. An attractive species that seldom exceeds a length of 6 inches.

Black Crappie Pomoxis nigro-maculatus (Le Sueur) A probable introduction that was reported by Jordan (1890:116) as common in Jericho Canal at Suffolk and by Fowler (1923b: 13) from the James River. Cope's (1869:24) record of Pomomys sp. may have referred to this or to the white crappie, Pomoxis annularis, a species expected but not yet recorded from the area. A good game and pan fish that reaches a weight of about 2 pounds.

Mud Sunfish Acantharcus pomotis (Baird) Although the mud sunfish is found in other parts of the Coastal Plain there is only one record from the lower James area. Jordan (1890:116) found 2 specimens in Jericho Canal at Suffolk. Six inches is about the maximum length. 182 THE JAMES RIVER BASIN

ATHERINIDAE Glassy Silverside Menedia beryllina (Cope) Occurs in brackish water throughout Chesapeake Bay. It readily enters freshwater and occasionally is found upstream almost to the Fall Line but its distribution in the lower James is imperfectly known (see Hildebrand and Schroeder 1928:191). Attains a length of 3 inches. The eggs are laid during the summer and adhere to objects on or near the bottom by means of adhesive threads. Of some value as forage for larger fishes.

SCULPIN FAMILY COTTIDAE Two kinds of sculpin or muddlers are found in the James. The northern muddler is widely distributed in small streams and springs in the mountains. The common slimy sculpin has only been taken at one locality. It is a northern species that apparently has been able to survive only in cold springs in the southern part of its range. In both species the male guards a nest hollowed out under a stone in early spring. The yellow mass of eggs is fastened to the underside of the stone.

Northern Muddler Cottus bairdii bairdii Girard Common in many of the riffles of headwater streams and in springs in the mountains and upper Piedmont. Taken by Cope (1869: 226) who said it abounded in cold springs, and by Jordan (1890:113) who reported it from Elk Creek opposite Natural Bridge. More re- cently it was taken by Burton and Odum (1945:186) in 9 out of 28 localities in Johns and Craig creeks. Also captured at 7 out of 57 other recently collected localities. It lives on or near the bottom and seldom gets more than 4 or 5 inches long.

Common Slimy Muddler Cottus cognatus gracilis Hackel Recorded by Hubbs and Lagler (1947:98) on the basis of speci- mens taken by Dr. Lincoln C. Pettit from Cove Run, a tributary of Maury River, in Rockbridge County. Five specimens were obtained. The palatine teeth were absent in 3 specimens, present as rudiments on one side and fully developed on the other in a fourth specimen, and fully toothed on both sides in a fifth specimen. The pelvic rays were I, 3 on both sides in 3 specimens, I, 3-4, I in one, and I, 4-4, I in an- other. The anal rays were 12 in 4 specimens and 13 in a fifth, the 13th ray in the latter was a rudiment. This represents the southernmost ex- treme of the known range. This normally cold water species has en- FRESHWATER FISHES 183 dured as a relict and should be looked for in other springs in the same area. STICKLEBACK FAMILY GASTEROSTEIDAE Fourspine stickleback Apeltes quadracus (Mitchill) A common species in the brackish waters of Chesapeake Bay and according to Hildebrand and Schroeder (1928:181) they enter prac- tically all the creeks and bays in weedy areas. Found at several places in the lower James River but its detailed distribution has not as yet been worked out. The writer has taken it in the Rappahannock River at Fredericksburg and it is probable that it occurs commonly through- out the lower courses of all the streams entering Chesapeake Bay. A small fish that seldom reaches a length of more than 2.5 inches. The eggs are deposited in a spherical nest made of plant parts glued together by a kidney secretion of the male. Of some importance as a forage fish. The threespine stickleback, Gasterosteus aculeatus Linnaeus, has been taken at Cape Charles and in the vicinity of Hampton (see Hilde- brand and Schroeder, 1928:180). Since this species often enters and may live permanently in freshwater it is also expected to be found at times in the lower James.

OTHER FISHES EXPECTED IN THE JAMES It is probable that the fishes listed below occur in the James River Basin since they are found in river systems both to the north and south of the James.

LAMPREY FAMILY PETROMYZONIDAE Eastern Brook Lamprey, Lampetra aepyptera (Abbott).

STURGEON FAMILY ACIPENSERIDAE Shortnose Sturgeon, Acipenser brevirostris Le Sueur.

SALMON FAMILY SALMONIDAE Brown Trout, Salmo trutta Linnaeus. Rainbow Trout, Salmo gairdnerii Richardson [stocked].

SUCKER FAMILY CATOSTOMIDAE Lake Quillback, Carpiodes cyprinus cyprinus (Le Sueur). Eastern Lake Chubsucker, Erimyzon sucetta sucetta (Lacepede). Eastern Redhorse, Moxostoma macrolepidowm (Le Sueur). 184 THE JAMES RIVER BASIN

MINNOW FAMILY CYPRINIDAE Carp, Cyprinus carpio Linnaeus. Goldfish, Carassius auratus (Linnaeus). Ironcolor Shiner, Notropis chalybaeus (Abbott).

PERCH FAMILY PERCIDAE Yellow Perch, Perca fiavescens (Mitchill).

SUNFISH FAMILY CENTRARCHIDAE Green Sunfish, Lepomis cyanellus Rafinesque. Northern Blackbanded Sunfish, Mesogonitsius chaetodon chaeto- don (Baird). White Crappie, Pomoxis annularis Rafinesque.

FISHES PROBABLY ERRONEOUSLY REPORTED FROM THE JAMES The fishes listed below have been reported from the James River at one time or another but there are grave doubts about the validity of the identifications. Each species has been discussed in the proper place above.

CATOSTOMIDAE Moxostoma cervinum (Cope).

CYPRINIDAE Notropis atherinoides Rafinesque. Notropis cerasinus (Cope). Notropis procne longiceps (Cope).

Notropis macdonaldi Jordan and Jenkins (is a hybrid Notropis cornutus cornutus X Notropis rubellus).

PERCIDAE Hadropterus crassus (Jordan and Brayton). Boleosoma nigrum efftdgens (Girard). Boleosoma podostemone (Jordan and Jenkins). FRESHWATER FISHES 185

ORIGIN AND RELATIONSHIPS OF THE JAMES FISH FAUNA Only two of the 74 kinds of fishes now known from the James River system seem to be endemic. Boleosoma longimanum has been reported from the Potomac River system (see discussion under the species account) but this record is regarded as probably erroneous and its distribution in drainages other than the James remains to be demonstrated. The other form limited to the James is an undescribed subspecies, Boleosoma nigrum subsp. It is related to but shows a con- siderable degree of differentiation when compared with the typical western Boleosoma nigrum nigrum. It is much different from the sub- species, Boleosoma nigrum olmstedi, which is present on the Coastal Plain, and also is separable from the Johnny darters of the Roanoke River system as well as those in more northern drainages. A study of the systematic relationships of eastern representatives of Boleosoma is now in progress. The James River fish fauna is derived from several geographic sources. A considerable element is marine in origin and all in this group have a more or less wide distribution along the Atlantic Coast. Most of the freshwater fishes now present in the James are distributed along the Atlantic Coast and often elsewhere. Of the species that have a more limited easterly distribution, those in the James are mostly related to the northern fauna. Generally, as far as freshwater fishes are concerned, one of the most distinct faunal breaks occurs between the James River on the north and the Roanoke (and Chowan) River on the south. The Dismal Swamp area has not been a barrier to the typical Coastal Plain species but many of the Piedmont and mountain types have not crossed this region. There is a small montane fauna which is represented in the James by such species as Thoburnia rho- thoeca and Crosomus oreas. In general the fish fauna west of the Allegheny divide is much different than that of the James except for certain species of rather wide distribution. A few species have been introduced by man. Elements of the James River fauna that have been derived from salt water are:

Petromyzon marinus Fundulus d. diaphanus Acipenser sturio oxyrhynchus Fundulus heteroclitus macrolepi-

Alosa sapidissima dotus Pomolobus aestivalis Roccus saxatilis Morone americana Pomolobus pseudoharengus Anguilla bostoniensis Meneidia beryllina Apeltes quadracus 186 THE JAMES RIVER BASIN

James River fishes that have a rather wide distribution in fresh-. waters along the Atlantic Coast or at least occur both to the north and south of the James River system:

Amia calva Schilbeodes rnarginatus DOrosoma cepedianum Umbra py gmaea Salvelinus f. fontinalis Esox americanus Catostonms c. commersonnii Esox niger Hypentelium nigricans Gambusia affinis Erimyzon oblongus Aphredoderus sayanus Semotilus a. atromaculatus Boleosoma nigrum olmstedi Clinostomus vandoisulus loa vitrea Notropis amoenus Hololepis fusiformis Notropis procne Poecilichthys flabellaris Hy bognathus nuchalis regius Chaenobryttus coronarius Cam postoma anomalum Lepomis gibbosus lctalurus catus Lepomis macrochirus Ameiurus nebulosus Lepomis auritus Ameiurus natalis Enneacanthus obesus Schilbeodes morns Enneacanthus gloriosus Acantharcus pomotis

Fishes found to the north (sometimes also to the west) and that apparently reach their southern limit of their range in the James River system.

Thoburnia rhothoeca Notropois procne procne Rhinichthys cataractae Notropis .cornutus cornutus Nocomis micropogon Hadropterus notogrammus Hy borhynchus notatus Hadropterus peltatus peltatus Notropis rube//us Cottus bairdii bairdii LeliCOSOMUS corpora/is Cottus cognatus gracilis

Others that go south beyond the James at least to the Roanoke River are Exoglossum maxillingua and Rinichthys atratulus atratulus. Southern fishes that seem to reach the northern limit of their dis- tribution in the James are Notropis hudsonius saludanus, Chrosomus oreas, and Chologaster cornutus. The Carolina chub, Nocomis leptocephalus, apparently reaches the northern limit of its range in the York River system which is just north of the James. Notropis ardens ardens is found only in the James and Roanoke systems. It is closely related to Notropis ardens matutinus (Cope) of the more southern Tar and Neuse River systems. FRESHWATER FISHES 187

Among the southeastern fishes that reach their northern limit of range in the Roanoke or Chowan River systems are the following:

Moxostoma cervinum and Notropis albeolus perhaps others of this genus Notropis altipinnis chowanus Nocomis sp. Notropis procne longiceps Notropis niveus? Schilbeodes marginatus atrorus Notropis alborus Fundulus rathbuni Hadropterus roanoka

A number of species are known with certainty only from the Roa- noke. Included in this category are:

Hypentelium roanokense Percina rex Thoburnia hamiltoni Hadropterus peltatus subsp. Notropis cerasinus Boleosoma podostemone Schilbeodes gilberti Boleosoma nigrum subsp. Arnbloplites cavifrons

Fishes introduced by man are Ictalurus lacztstris punctatzts, Mi- cropterus d. dolomieu, Micro pterus s. salmoides?, Ambloplites r. rupestris, and Pomoxis nigro-maculatus, Salim gairdnerii.

DISTRIBUTION OF FISHES WITHIN THE SYSTEM The James River like other parallel river systems that enter Chesa- peake Bay or the Atlantic Ocean, flows through three rather well de- fined physiographic regions. It rises in the Allegheny Mountains, flows easterly crossing the Great Valley, and cuts through the Blue Ridge south of Lexington. East of Lynchburg it turns northeast and parallels the Blue Ridge as far as Scottsville where it turns easterly. This area is spoken of generally as the mountain region as far as the distribution of the fishes is concerned. It includes the region from the vicinity of Lynchburg to Charlottesville where many of the streams rise in the Blue Ridge and are fairly typical mountain brook types, of alternating riffles and pools and with considerable gravel, rubble, and bedrock. The fishes of this region are better known than those of the other two physiographic provinces. The mountain streams because of their beauty and varied fish fauna attracted the attentions of Cope (1869), Jordan (1890) and Burton and Odum (1945), and most of the recent collecting has been done here. East of this region is the Piedmont which extends to the Fall Line just west of Richmond. These streams are characterized by sand bottom and are often turbid due to the heavy silt load they typically 188 THE JAMES RIVER BASIN

carry. Little rubble or bed rock bottom is present and few riffles are to be found. The Piedmont Province as a whole has been worked very little. It is difficult to seine in these streams because of the soft sandy bottom. Also the known fish fauna is rather limited. Indeed it has been a rather general practice of naturalists along the east coast to go either to the mountains or the Coastal Plain for interesting collecting. The Coastal Plain begins below the Falls at Richmond. This section of the river is sluggish and with silt or sand bottom. The fauna is fairly large but is not well known. It is difficult to collect here using the traditional seining methods. Generally speaking the Coastal Plain freshwater fish fauna of the East coast is known from collections made in other systems. However, the probability of adding a number of species, now on the hypothetical list for this area, should attract the efforts of young zoologists. This region is also interesting because of the many saltwater species that regularly or occasionally enter the freshwater. The following discussion of the distribution of the James River fishes evolves mostly around these three physiographic provinces. Scant information is available on the fishes of the Piedmont and Coastal Plain so that the writer has drawn on his notes on the distribution of the fishes in the other stream systems of Virginia. It should be emphasized that much additional work must be done before a relatively complete distributional picture is obtained. Fishes that are usually common in the mountain region are:

Salvelinus f. fontinalis Clinostomus vandoisulus Hypentelium nigricans Notropis c. cornutus Tboburnia rhothoeca Campostoma anornalum Leucosomus corporalis Schilbeodes m. marginatus Nocomis micropogon Boleosoma nigrum subsp. Nocomis leptocephalus Boleosoma longimanum Rhinichthys a. atratulus Cottus b. bairdii

In addition to the above the following are found in the mountains but generally are not quite as common.

Chrosomus areas Esox niger Notropis rubellzts Micropterus d. dolomieu Notropis a. ardens Lepomis auritus N otro pis hit dsonius Ambloplites r. rupestris Hy borhynchus nOtatus Poccilichthys flabellaris Rhinichthys cataractae Cottus cognatus FRESHWATER FISHES 189

A few of the above species are probably to be considered more typical mountain species than others. Since they do not occur else- where in the system, except very occasionally in the headwaters of Piedmont brooks. In addition they occupy mountain type streams in other drainage systems. Examples of these fishes are Salvelinus f. fontinalis, Rhinichthys cataractae, Cottus bairdii bairdii, and Cottus cognatus. Others like Thoburnia rhothoeca and Chrosonnts oreas are often common in mountain streams but also frequently are abundant in Piedmont streams such as the headwaters of the Appomattox River. Another group of fishes drawn from several families seems to be at home in the upper Piedmont or in the mountain region. Included here are such species as

Catostomus c. commersonnii Hy borhynchus notatus Hypentelium nigricans Ameiurus n. nebulosus Leucosomus corporalis Boleosoma nigrum subsp. Semotilus a. atromaculatus Hadropterus notogrammus Nocomis leptocephalus Poecilichthys flabellaris Notropis rubellus Micro pterus d. dolornieu Exoglossum maxillingua Ambloplites r. rupestris

Fishes found often in typical Piedmont streams arc

Leucosomus corporalis Notropis c. corn utus Nocomis leptocephalus Notropis analostanus Clinostomus vandoisulus Hy bognathus nuchalis regizts Notropis p. procne Boleosoma nigrum subsp. loa vitrea

Others that are found regularly either in the lower Piedmont or on the Coastal Plain are

Erimyzon o. oblongus Anguilla bostoniensis Ictalurus catus Aphredoderus s. sayanus Ictalurus lacustris punctatus Hadropterus p. peltatus Ameiurus n. nebulosus boa vitrea Ameiurus natalis erebennus Micro pterus s. salmoides Notropis amoenus Lepomis macrochirus Notemigonus c. crysoleucas Pomoxis nigro-maculatus 190 THE JAMES RIVER BASIN

The freshwater component of the Coastal Plain fauna includes

Lepisosteus o. osseus Chologaster cornutus Amia calva Boleosoma nigrum olmstedi Dorosoma cepedianum Hololepis fusiformis Sc.hilbeodes mollis Chaenobryttus coronarius Umbra py gmaea Centrarchus macro pterus Esox arnericanus Enneacanthus obesus Esox niger Enneacanthus gloriosus Gambusia affinis Acantharcus pomotis

A few species are found throughout the river system. The chain pickerel, Esox niger, is mostly found on the Coastal Plain but ranges to the mountain headwaters. The redbelly sunfish, Lepomis auritus, has a similar distribution. Other widely distributed species are the eel, Anguilla bostoniensis, spottail shiner, Notropis hudsonius, swallowtail minnow, Notropis procne, and the silvery minnow, Hybognathus nuchalis regius. Many marine fishes enter the freshwaters of the lower James. Some are anadromous and regularly make their way upstream, usually in the spring of the year, to lay their eggs. Included in this category are

Petromyzon marinus Pomolobus pseudoharengus Acipenser sturio oxyrhynchus Alosa sapidissima Pomolobus aestivalis Roccus saxatilis

Another group of predominantly marine fishes seem equally at home in salt or freshwater. These fishes often become adjusted to life in freshwater and may not return to the sea. Fishes in this category which have been included in the species discussion above are

Fundulus d. diaphanus Morone americana Fundulus heteroclitus macro- Menedia beryllina lepidotzis Apeltes quadracus

Other marine fishes, not discussed in the annotated list above, which occasionally enter freshwaters but which apparently do not stay long are

Brevoortia tyrannus Fundulus luciae Pomolobus mediocris Leucania parva Fundulus majalis Cyprinodon variegatus Anchoa m. mitchilli FRESHWATER FISHES 191

Most of the species of Chesapeake Bay shore fishes may be ex- pected to be found in the river even as far upstream as the Fall Line. Such possible irregular visitors are various sharks and rays, the ten- pounder, Elops saurus; the tarpon, Tarpon atlanticus; billfish, Strongy- lura marina, and some flatfishes, especially the hog choker.

LITERATURE CITED Breder, C. M., Jr. 1935 The Reproductive Habits of the Common Catfish, Ameitt- rus nebulosus (Le Sueur), with a Discussion of Their Signif- icance in Ontogeny and Phylogeny. Zoologica, 19 (4):153-185, Figs. 12-23. 1936 The Reproductive Habits of the North American Sunfishes (Family Centrarchidae). Zoologica, 21(1):1-48, Text Figs. 1-6, Pls. 1-7. Burton, George W. and Eugene P. Odum. 1945 The Distribution of Stream Fish in the Vicinity of Moun- tain Lake, Virginia. Ecology, 26 (2) : 182-194, Figs. 1-2. Cope, E. D. 1869 On the Distribution of Fresh-water Fishes in the Allegheny Region of Southwestern Virginia. Journ. Acad. Nat. Sci. Phila., (2)6:207-47, Pls. 22-25. 1870 A Partial Synopsis of the Fresh Water Fishes of North Carolina. Proc. Amer. Phil. Soc., pp. 448-95. Fowler, Henry W. 1918 Fishes from the Middle Atlantic States and Virginia. Occ. Pap. Mus. Zool. Univ. Mich., 56:1-19, Pls. 1-2. 1923a Records of Fishes for the Eastern and Southern United States. Proc. Acad. Nat. Sci. Phila., 74:1-27, Pls. 1-2. 1923b Records of Fishes for the Southern States. Proc. Biol. Soc. Wash., 36:7-34. 1924 Notes on North American Cyprinoid Fishes. Proc. Acad. Nat. Sci. Phila., 76:389-416, Figs. 1-23. [19441 Notes on Fishes Obtained in 1940, 1941 and 1942 with an Account of a New Genus of Rosy-sided Dace. Biennial Rept. Pa. Bd. Fish Comm., 1942, pp. 50-55, Fig. 1. 1945 A Study of the Fishes of the Southern Piedmont and Coastal Plain. Monogr. Acad. Nat. Sci. Phila., 7:i-vi, 1-408, Figs. 1-313. Hildebrand, Samuel F. and William C. Schroeder. 1928 Fishes of Chesapeake Bay. Bull. U. S. Bur. Fish., 43 (1927), Pt. 1:1-366, Figs. 1-211. Hubbs, Carl L. 1930 Materials for a Revision of the Catostomid Fishes of Eastern North America. Misc. Pub. Mus. Zool. Univ. Mich., 20:1-47, Frontisp. 192 THE JAMES RIVER BASIN

Hubbs, Carl L. 1932 The Scientific Name of Two Sunfishes, Helioperca macro- chira (Rafinesque) and Eupomotis microlophus (Gunther).. Occ. Pap. Mus. Zool. Univ. Mich., 305:1-12. Hubbs, Carl L. and Bailey, Reeve M. 1938 The Small-mouthed Bass. Bull. Cranbrook Inst. Sci., 10: 1-92, Pls. 1-9. 1940 A Revision of the Black Basses ( Micropterus and Huro), with Descriptions of Four New Forms. Misc. Pub!. Mus. Zool. Univ. Mich., 48:1-51, Pls. 1-6, Maps 1-2. Hubbs, Carl L. and Mott D. Cannon. 1935 The Darters of the Genera Hololepis and Villora. Misc_ Pub!. Mus. Zool. Univ. Mich., 30:1-93, Pls. 1-3. Hubbs, Carl L. and Karl F. Lagler. 1947 Fishes of the Great Lakes Region. Bull. Cranbrook Inst. Sci., 26:i-xi, 1-186, Colored pls. 26, Figs. 1-289, Map on end papers. Hubbs, Carl L. and Edward C. Raney 1944 Systematic Notes on North American Siluroid Fishes of the Genus Schilbeodes. Occ. Pap. Mus. Zool. Univ. Mich., 487: 1-36, Plate 1, Map 1. 1946 Endemic Fish Fauna of Lake Waccamaw, North Carolina. Misc. Pub. Mus. Zool. Univ. Mich., 65:1-30, Plate 1, Maps 1-2. 1947 Notropis alborus, a New Cyprinid Fish From North Caro- lina and Virginia. Occ. Pap. Mus. Zool. Univ. Mich., 498:1-17, Fig. 1, Plate 1, Map 1. Jordan, David Starr 1890 Report of Explorations Made During 1888 in the Allegheny Region of Virginia, North Carolina, and Tennessee, and in Western Indiana, with an Account of the Fishes Found in Each of the River Basins of Those Regions. Bull. U. S. Fish. Comm., 8(1888):97-173, Pls. 13-15. 1929 Manual of the Vertebrate Animals of the Northeastern United States Inclusive of Marine Species. 13th ed. World Book Co., N. Y.: i-xxxi, 1-446, Figs. 1-15, Map 1. Jordan, David S. and Barton W. Everrnann 1896 The Fishes of North and Middle America. . . Bull. U. S. Nat. Mus., 47, Pts. 1-4:1-3313, Figs. 1-958. Jordan, David S., Barton W. Evermann, and Howard W. Clark 1930 Check List of the Fishes and Fishlike Vertebrates of North and Middle America North of the Northern Boundary of Vene- zula and Columbia. Rept. U. S. Comm. Fish., 1928(2):1-670. Lachner, Ernest Albert 1947 Studies on the Biology of the Chubs (genus Nocomis, FRESHWATER FISHES 193

family Cyprinidae) of Northeastern United States. Cornell Ab- stracts of Theses, p. 207-210. Menzel, R. Winston 1945 The Catfish Industry of Virginia. Trans. Amer. Fish. Soc., 73(1943):364-372, Fig. 1. Miller, Robert R. 1946 Distributional Records for North American Fishes, with Nomenclatorial Notes on the Genus Psenes. Journ. Wash. Acad. Sciences, 36(6):206-212. Pearson, John C. 1941 The Young of Some Fishes Taken in Lower Chesapeake Bay, Virginia, with Special Reference to the Gray Sea Trout, Cynoscion regalis (Bloch). U. S. Fish and Wildlife Service, Fishery Bull. 36, Vol. 50:79-102, Figs. 1-26. Raney, Edward C. 1939 The Breeding Habits of the Silvery Minnow, Hybognathus regius Girard. Amer. Mid. Nat., 21(3):674-680, Figs. 1-3. 1940a The Breeding Behavior of the Common Shiner, Notropis cornutus Zoologica 25(1):1-14, Pls. 1-2. 1940b Nests Under the Water. Bull. N. Y. Zool. Soc., 43(4): 127-135, Figs. sev. 1940c Reproductive Activities of a Hybrid Minnow, Notropis cornzaus x Notropis rubellus. Zoologica 25(3): 361-367. 1941 Records of the Brook Lamprey, Lampetra aepyptera ( Abbott), From the Atlantic Drainage of North Carolina and Virginia. Journ. Elisha Mitchell Sci. Soc., 57(2):318-320. 1947a Subspecies and Breeding Behavior of the Cyprinid Fish Notropis procne (Cope). Copeia, No. 2:103-109. 1947b Nocomis Nests Used by Other Breeding Cyprinid Fishes in Virginia. Zoologica, 32(3):125-132, Figs. 1-2. Raney, Edward C. and Carl L. Hubbs. 1948 Hadropterus notogrammus, a New Percid Fish from Mary- land, Virginia, and West Virginia. Occ. Pap. Mus. Zool. Univ. Mich. 512:1-26, Plates 2, Map 1. Raney, Edward C. and Ernest A. Lachner. 1943 Age and Growth of Johnny Darters, Boleosoma nigrum olmstedi (Storer) and Boleosoma longimanum (Jordan). Amer. Midl. Nat., 29(1):229-238, Figs. 1-5. 1946a Age, Growth, and Habits of the Hogsucker, Hypenteliurn nigricans (Le Sueur), in New York. Amer. Mid. Natl., 36(1): 76-86, Figs. 1-4. 1946b Age and Growth of the Rustyside Sucker, Thoburnia rhothoeca (Thoburn). Amer. Midl. Nat., 36(3):675-681, Figs. 1-2. 194 THE JAMES RIVER BASIN

Raney, Edward C. and Ernest A. Lachner. 1946c Thoburnia hctmiltoni, a New Sucker from the Upper Roanoke River System in Virginia. Copeia, p. 218-226, Pl. 1. 1947 Hypentelium roanokense, a New Catostomid Fish from the Roanoke River in Virginia. Amer. Mus. Novitates, No. 1333: 1-15, Figs. 1-3. Raney, Edward C. and Royal D. Suttkus. 1948 The Subspecies of the Shielded Darter, Hadropterus pelta- tus. Abstracts 28th Ann. Meeting Amer. Soc. Ichth. and Herp. New Orleans, p. 2. Richmond, Neil D. 1940 Nesting of the Sunfish, Lepomis auritus (Linnaeus), in Tidal Waters. Zoologica, 25(3):329-330, Figs. 1-2. Schultz, Leonard P. 1939 The Fresh-water Fishes of Virginia. Explorations and Field-work of the Smithsonian Institution in 1938, p. 55-58, Figs. 53-57. Ithaca, New York