BULLETIN 445 APRIL 1973 Distribution of the Gcnus Ergasi in Several Gulf /s of Mexico Drainage Basins r :'+ AGRICULTURAL EXPERIMENT STATION AUBURN R. Dennis Rouse, Director UNIVERSITY Auburn, Alabama CONTENTS Page INTRODUCTION ----------------------BACKGROUND -- --- --- --- --- -- --- --- --- ---THE STUDY AREA AND FISH FAUNA-4 TAXONOMIC CONSIDERATIONS OF THE GENUS Ergasilus- - -3 4 4 8 METHODS AND MATERIALS---- LOCALITIES AND NUMBERS OF FISH MATERIALS-9 R E SU LTS --- -- - - -- - -- - - -- - -- - - -- - -- - - -PARASITE MATERIAL AND RESPECTIVE HOST INFORMATION ERGASILUS MATERIAL ACCUMULATED DURING STUDY FROM OUTSIDE OF STUDY AREA-27 -24 24 DISCU SSIO N ---------------------------------------- 29 HOST-PARASITE PARASITE-HOST RELATIONSHIPS -_________29 RELATIONSHIPS ANDDISTRIBUTION- -39 REVISED PARASITE-HOST CHECKLIST FOR THE GENUS ERGASILUS --------------------------- 61 SUMMARY AND CONCLUSIONS ---------------------- 64 ACKNOW LEDGMENTS-------------------------------66 A D DEND U M ----------------------------------------- 67 LITERATURE CITED ------------------------- =------69 FIRST PRINTING 3M, APRIL 1973 Distribution of the Genus Ergasilus in Several Gulf of Mexico Drainage Basins S. K. JOHNSON and W. A. ROGERS* INTRODUCTION IHE GENUS Ergasiluscomprises a group of parasitic crustaceans (copepods) that may be considered a threat to fish health. These parasites typically attach to the gill filaments of the fish host where they feed on body tissues. Irritation and tissue damage are apparent symptoms. Several epizootics caused by Ergasilus are noted in the literature. Ergasilus species may be considered a threat to a fish population when the sequence of natural events promotes the development of large numbers of the parasites. The effect is that of overburdening the hosts. The presence of few Ergasilus parasites typically has little harmful effect on the fish. The fundamental concepts of identity, host relationships, and distribution are necessary for the proper understanding of a group of parasites. The distribution of the genus Ergasilus in several Gulf of Mexico drainage basins is examined herein with emphasis on host relationships and distributional patterns. Combining the resulting data with the more reliable reports of the literature helps to establish more clearly the status of distribution of the Ergasilus species along with their preferred hosts. For this study gill material was examined from 6,965 fishes representing over 3,000 host localities. Fish specimens were obtained primarily from ichthyological museums. * Formerly Research Assistant, Department of Fisheries and Allied Aquacultures, now Fish Disease Specialist, Texas Agricultural Extension Service, Texas A&M University; Associate Professor, Department of Fisheries and Allied Aquacultures. 4 ALABAMA AGRICULTURAL EXPERIMENT STATION Background Wilson (99,100) made the most important contribution in establishing a taxonomy for the genus Ergasilus in North America. Several new species were described and comments were made on host preferences. Wilson's conclusions on host preferences, however, were frequently supported by little data. Following this work were several descriptions of new taxa but most of them were descriptions of previously described forms. In the 1960's, L. S. Roberts began revisory work that culminated in a general review of the genus (80). Most of Roberts' work was on material from the U.S. National Museum collection and material that R. V. Bangham had collected in his (19331955) survey work. Part of the latter material that Roberts had the opportunity to rework had by that time lost its locality data. Nevertheless, Roberts had provided valuable information for distributional work by revising that material on which he did have locality data. The Study Area and Fish Fauna The area selected for study includes several river drainages that flow southward into the Gulf of Mexico and that lie between 30-35 degrees N. latitude. The primary adjacent waters include the large Mississippi River drainage to the west and Atlantic Ocean drainages to the east. The drainage area studied lies within. Alabama, Mississippi, northwestern Florida, and western Georgia. If the size of the drainage area is considered it would be safe to say that the waters support a relatively diverse fish fauna (66). Nearly all of the North American freshwater fish families, with the exception of salmonids, umbrids, and percopsids, have representatives in the area. Smith-Vaniz (84) considered the zoogeography of the fish fauna of most of the area included in the present study and noted the Mobile Bay and Apalachicola Bay drainages to be centers of fish species differentiation. Taxonomic Considerations of the Genus Ergasilus There are 23 described Ergasilus species in North America that are regarded as valid species. Roberts' (80) review included 21 species but suggested that two of these, E. funduli Kroyer, 1863 and E. manicatus Wilson, 1911 may be the same. It appears that DISTRIBUTION OF THE GENUS ERGASILUS 5 the names represent indeed the same species with the name E. funduli having priority. Another species, included by Roberts (80) as E. mugilus Vogt, 1877 sensu Wilderman, 1968, unpublished, has been considered a synonym of E. versicolor (51). Since Roberts' (80) review, another taxonomic work (27) described a new species and transferred one North American Ergasilus species to a new genus. The new species, E. spathula Cressey, 1970, is herein considered a synonym of E. arthrosis Roberts, 1969. The described North American species of Ergasilus that are considered acceptable in this work are: E. arthrosisRoberts, 1969 (Syn. E. spathula Cressey, 1970); E. auritus Markevich, 1940; E. caeruleus Wilson, 1911; E. celestis Mueller, 1936 (Syn. E. osborni Tidd and Bangham, 1945); E. centrarchidarumWright, 1882 (Syn. E. nigritus Wilson, 1916); E. cerastes Roberts, 1969; E. chatauquaensis Fellows, 1887; E. clupeidarum Johnson and Rogers, 1972; E. cotti Kellicott, 1892; E. cyprinaceus Rogers, 1969; E. elongatus Wilson, 1916; E. felichthys (Pearse, 1947); E. funduli Kroyer, 1863 (Syn. E. manicatus Wilson, 1911); E. labracis Kroyer, 1863; E. lanceolatus Wilson, 1916; E. lizae Kroyer, 1863; E. luciopercarum Henderson, 1926 (Syn. E. confusus Bere, 1931; E. skrjabini Mueller, 1936); E. megaceros Wilson, 1916 (Syn. E. fragilis Mueller, 1936); E. nerkae Roberts, 1963; E. tenax Roberts, 1965; E. turgisus Fraser, 1920; E. versicolor Wilson, 1911 (Syn. E. elegans Wilson, 1916; and E. cyanopictus Carvalho, 1962); E. wareaglei Johnson, 1973. There is a group of names in the North American literature that has been given to Ergasilus species that parasitize coastal Mugil spp. The original descriptions of these species lacked the explicitness required for proper differentiation. Included in this group are E. nanus van Beneden, 1870, E. mugilus Vogt, 1877, and E. lizae Kroyer, 1863. The results of the present study will show there are probably three forms that will fit the descriptions of these vaguely defined species. One of the forms treated herein is reported as E. lizae following Roberts' (80) definition, and all three forms are depicted in Figure 1. Another point that had not been brought out in the literature was the close morphological similarity of E. arthrosis Roberts, 1969 to E. sieboldi Nordmann, 1832. E. sieboldi is considered to have an Eurasian distribution. The senior author had the opportunity to examine Asian and European material of this species 6 ALABAMA AGRICULTURAL EXPERIMENT STATION and found it to vary considerably in morphology for an Ergasilus species. Roberts (78) did not compare E. arthrosis to E. sieboldi 0.15mm , FIG. 1. Second antennae of three Ergasilus species: a-antenna from E. sp. Form A from gill rakers of Mugil cephalus from shore waters, Mobile Co., Ala.; b-antenna from E. Iizae Kroyer from gill filament of M. cephalus, same location; c-antenna of E. sp. Form B from gill arch of M. cephalus from Intracoastal Waterway, Baldwin, Co., Ala. DISTRIBUTION OF THE GENUS ERGASILUS 7 in his description. It is suspected that E. arthrosiswill eventually be shown to be part of a polytypic species of E. sieboldi or perhaps a member of a closely related species complex. As for now, E. arthrosis will be considered as a legitimate species. Roberts (78) described E. arthrosis from paratype material of E. versicolor Wilson, 1911. Roberts was able to show that Wilson had inadvertently based his description of E. versicolor on both holotype and paratype material and as a result characters of both animals were given in the description. Apparently misled by his own error, Wilson described a second species, E. elegans, that was actually the same as the E. versicolor holotype. Roberts (78) recognized the mistake, erected a new species, E. arthrosis, for the E. versicolor paratype material, and synonymized E. elegans with E. versicolor. It is of primary importance here to point out that after Wilson (99) and prior to Roberts (78) there was confusion regarding the taxonomy of these species and as a result species reported as E. versicolor and E. elegans must be regarded as questionable. A similar period of possible misinterpretation may be attributed to E. caeruleus Wilson, 1911. Mueller (68) synonymized E. skrjabini Mueller, 1936 and E. confusus Bere, 1931 with E. caeruleus. It was later shown by Roberts (76) that E. skrjabini and E. confusus were actually synonyms of E. luciopercarum Henderson, 1926. As above, species reported under these taxa must be considered with caution. Roberts (80) reported a species of Ergasilus as E. lizae (?) from Lepomis auritus, L. gibbosus, Perca flavescens, Esox niger, and Salmo salar. The report of this species is not well defined and it is assumed that this record will be given further attention by Roberts. Since there is uncertainty about the taxonomic status of the species that are close to E. lizae, this record will not be given further attention in the present work to prevent confusion. Thomsen (88) described a species of Ergasilus from South America as E. elongatus. This was shown to be a homonym by Yamaguti (106) and was given the new name of E. thomseni. Johnson and Rogers (51) later considered E. thomseni a junior synonym of E. felichthys (Pearse). METHODS AND MATERIALS Drainage basins selected for study are shown in Figure 2. Some of the larger basins were divided and the parts studied as 8 ALABAMA AGRICULTURAL EXPERIMENT STATION separate drainage units. An attempt was made to study samples of all representative fish species of each drainage unit. Gill material samples were obtained primarily from fish specimens stored in ichthyological museums supplemented with material from personal fish collections. Gill samples were removed from the right side of the fish specimens by removal of all right gill arches. The gill material was placed in 3 X 4 inch zip lip plastic bags partially filled with 5 per cent formalin. Collection data for the fish were recorded, the bag was numbered, and the gill material saved for future examination. Upon examination, each gill arch was thoroughly searched front and back and between the gill filaments with the aid of a dissecting microscope. When ergasilids were found, observations on location and pathology were noted and the specimens were removed and placed in glass vials filled with 5 per cent formalin. The 6,965 fish that were treated in this fashion are summarized on pages 9-28. Additional materials from outside of the primary study area were accumulated during the investigation and are also presented as part of these data. For counting and identification, the copepod specimens were studied as wet mounts in lactic acid or as permanent preparations mounted in Hoyer's medium or in glycerin gel by the method of Johnson (49). Phase microscopy proved particularly helpful and was used throughout the study. Host names and other taxa were reconciled with currently accepted nomenclature by following American Fisheries Society, Committee on Names of Fishes (1). The sequence of family presentation follows Greenwood, Rosen, Weitzman, and Myers (37), and with the exceptions of retaining Elassomidae and Eleotridae, the taxa are followed as well. Genera and species are arranged alphabetically. Most of the ergasilid material has been retained by the author but representative samples of representative species have been deposited in the following museums: British Museum, London; U.S.S.R. Zoological Institute, Leningrad; and U.S. National Museum Helminthological Collection, Beltsville. Localities and Numbers of Fish Materials Following the species name in this listing, numerical representations of data are presented as follows: number of specimens DISTRIBUTION OF THE GENUS ERGASILUS 9 FIG. 2. Drainages sampled: 1-Upper Pearl R.; 2-Lower Pearl R.; 3-Bay St. Louis; 4-Biloxi Bay; 5-Leaf R.; 6-Chickasawhay R.; 7-Pascagoula R.; 8-Escatawpa R.; 9-Mobile Bay; 10-Mobile-Tensaw R.; 11-Upper Tombigbee R.; 12-Lower Tombigbee R.; 13-Lower Alabama R.; 14--Upper Alabama R.; 15-Black Warrior R.; 16-Cahaba R.; 17-Upper Coosa R.; 18-Lower Coosa R.; 19-Upper Tallapoosa R.; 20-Lower Tallapoosa R.; 21-Perdido R.; 22-Escambia R.; 23-Blackwater R.; 24-Yellow R.; 25Ch~octawhiatch~ee R.; 26-Upper Chattahioochee R.; 27-Lower Chattahioochee R.; 28-Appalachicola R.; 29-Mississippi Sound. examined within a drainage, dash, number of localities examined within that drainage, and within parentheses, the number of the drainage as that shown on Figure 2. Lettering: TN - total number of specimens examined; TL - total number of localities examined; NP - number of specimens positive; LP - number of localities positive. ACIPENSERIDAE Scaphirhynchu8 platorynwhus. 1-1(13), TN-i, TL-1, NP-o, LP-O. 10 10 ALABAMA AGRICULTURAL EXPERIMENT STATION Aciperser oxyrhynchus. 1-1(1), 2-2(2), 1-1(13), TN-6, TL-6, NP-0, LP-0. 1-1(10), 1-1(11), POLYODONTIDAE Polyodon spathula. 1-1(1), 1-1(2), 1-1(12), 1-1(20), TN-4, TL-4, NP-i, LP-1. LEPISOSTEIDAE Lepisosteuas oculatus. 2-2(1), 1-1(4), 1-1(5), 1-1(6), 1-1 TN-14, TL-12, NP-0, LP-0. L. osseus. 1-1(8), 1-1(9), 1-1(11), 3-2(15), 2-1(18), TN-8, TL-6, NP-i, LP-1. A.MIIDAE Amia calva. 2-2(1), 1-1(3), 1-1(5), 1-1(8), 1-1(11), 3-2(15), 1-1(22), 1-1(25), 2-1(27), TN-13, TL-11, NP-0, LP-0. ELOPIDAE Elops saurus. 3-2(9), TN-3, TL-2, NP-0, LP-0. ANGUILLIDAE Anguilla rostrata. 1-1(1), 1-1(2), 2-2(3), 5-4(5), 3-3(6), 1-1(18), 1-1(20), 4-3(25), 1-1(27), TN-28, TL-25, NP-10, LP-8. CLUPEIDAE Alosa alabamae. 2-1(10), 2-1(16), TN-4, TL-2, NP-0, LP-0. A. chrysochioris. 1-1(1), 2-2(2), 2-2(5), 5-3(6), 2-2(9), 4-1(10), 4-2(11), 6-2(12), 1-1(15) 2-1(16), 2-1(18), 3-1(24), 3-1(25) , 1-1(27) , TN-38, TL-21, Brevoortia patronus. NP-4, LP-1. TN-5, 1-1(10), 3-1(23), 1-1(25), 7-4(1), 3-2(2), TL-3, NP-0, LP-0. Dorosoma cepedianum. 1-1(3), 2-2(5), 13-4(20), 3-3(22), 2-1(25), 2-1(26), 6-2(27), TN-54, TL-28, NP-i, LP-1. D. petenense. 4-3(1), 2-2(3), 2-1(8), 7-3(9), 1-1(10), 6-2(12), 3-1(15), 4-2(18), 5-2(20), 2-1(23), 3-1(25), 3-1(26), 6-2(27), TN-48, TL--22, NP-i, LP-1. DISTRIBUTION OF THE GENUS ERGASILUS 11 DISTRIBUTION OF THE GIENUS ERGASILUS I ENGRAULIDAE Anchoa mitchilli. 2-1(2), 5-3(3), 6-2(9), 4-1(10), 3-1(12), 2-1(28), TN-22, TL-9, NP-0, LP-0. A. hepsetus. 4-2(9), 7-2(29), TN-11, TL-4, NP-0, LP-0. HIODONTLDAE Hiodon tergisus. 5-1(1), 2-1(3), 2-1(6), 3-2(11), 3-1(12), 2-1(15), 3-2(16), TN-20, TL-9, NP-i, LP-1. ESOCIDAE Esox americanus. 1-1(1), 2-2(2), 7-4(3), 1-1(4), 5-5(5), 1-1(6), 6-4(7), 1-1(8), 1-1(9), 2-1(10), 4-3(11), 3-2(12), 1-1(13), 1-1(14), 3-2(15), 3-2(16), 3-2(18), 3-2(20), 2-1(21), 4-2(22), 1-1(23), 2-1(24), 5-3(25), 4-2(26), 4-3(27), 1-1(28), TN-71, TL-50, NP-3, LP-3. E. nicer. 2-2(1), 1-1(3), 3-2(5), 1-1(6), 2-2(7), 1-1(8), 1-1(25), 1-1(26), 2-2(27), TN-29, TL-26, NP-2, LP-2. CYPRINLDAE Cyprinus carpio. 4-2(27), TN-4, TL-2, NP-0, LP-0. Campostoma anomalum. 10-4(11), 7-3(12), 3-1(13), 3-1(14), 6-2(15), 3-1(16), 2-1(17), 4-2(18), 3-1(19), 3-1(20), 2-1(22), 1-1(27), TN-47, TL-19, NP-i, LP-1. Ericymba buccata. 3-1(1), 12-5(2), 3-1(4), 1-1(5), 6-4(6), 21-8(7), 5-2(11), 3-1(12), 3-1(13), 3-1(14), 5-2(16), 3-1(18), 7-2(20), 9-3(22), 3-1(24), 9-3(25), , TN-lOS, TL-40, NP-0, LP-0. 3-1(26) , TN-3, TL-1, NP-0, LP-0. Hemitremia flammea. Hybognathus hayi. 6-2(27) 5-2(1), 3-1(17), 4-2(11), 4-2(15), 2-1(20), TN-iS, TL-7, NP-0, LP-0. H. nuchalis. 2-1(1), 1-1(5), 4-1(6), 4-1(10), 6-2(11), TN-39, TL-15, NP-2, LP-2. 4-2(16), Hybopsis aestivalis. 8-2(5), 3-1(6), 9-3(11), 3-1(12), 1-1(19) , 2-1(20), 9-3 (22) , TN-39, TL-14, NP-0, LP-0. H. amplops. 3-2(1), 2-1(3), 8-1(4), 8-2(5), 3-1(6), 4-2(7), 3-1(11), 3-1(12), 3-1(14), 3-2(15), 1-1(16), TN-36, TL-15, NP-2, LP-1. H. harperi. 3-2(2),4-2(5), TN-7, TL-4, NP-0, LP-0. 12 12 ALABAMA AGRICULTURAL EXPERIMENT STATION H. storeriana. 1-1(2), 6-3(5), 10-3(6), 6-3(10), 6-3(11), 8-3(12), 5-2(15), 3-1(16), 3-1(21), TN-48, TL-20, NP-4, LP-3. H. lineapunctata. 5-2(18), TN-5, TL-2, NP-0, LP-0. Hybopsis sp. cf. winchelli 4-2(21), 4-2(22), 2-1(24), 9-3(25), 3-1(26), 2-1(27), TN-24, TL-10, NP-0, LP-0. Nocomis leptocephalus. 6-3(1), 2-2(2), 3-1(5), 2-1(6), 11-5(11), 4-2(12), 3-1(13), 4-2(14), 3-1(15), 5-2(16), 2-1(19), 2-1(22), 5-2(26), TN-52, TL-24, NP-3, LP-1. Notemigonus crysoleucas. 2-1(1), 1-1(6), 1-1(7), 2-1(10), 9-3(11), 2-2(12), 3-1(13), 2-1(14), 7-3(15), 3-1(16), 2-2(17), 1-1(18), 3-1(19), 5-3(20), 5-2(22), 1-1(23), 9-4(25), 2-1(26), 2-2(27), TN-73, TL-39, NP-i, LP-1. asperifrons. 3-1(15), 2-1(16), 3-1(18), 1-1(19), TN-9, TL-4, NP-0, LP-0. N. atherinoides. 6-3(1), 11-4(5), 8-2(6),4-2(7), 3-1(12), 3-1(15), 1-1(16), 1-1(18), 2-1(19), TN-39, TL-16, NP-0, LP-0. N. baileyi. 5-1(5), 12-4(11), 3-2(12), 4-1(13), 3-1(14), 3-1(15), 7-3(16), 3-1(18), 15-2(20), 3-1(22), 6-2(27), TN-64, TL-19, NP-5, LP-4. N. bellus. 9-3(11), 4-2(12), 3-1(13), 3-1(14), 6-2(16), 2-1(18), 3-1(19), 15-2(20), TN-45, TL-13, NP-3, LP-1. N. caeruleus. 2-1(17), 1-1(18), TN-3, TL-2, NP-0, callistius. 4-1(14), 8-3(15), 3-1(16), 3-1(17), 2-1(18), 5-3(3)) 6-4(5), Notropis 3-1(19), 5-2(26), N. LP-0. TN-23, TL-8, NP-0, LP-0. N. callitaerda. 2-1(26), 4-1(27), TN-6, TL-2, NP-0, LP-0. N. chalybaeus. 3-1(3), 4-2(4), 3-1(10), 2-1(12), 4-1(22), TN-21, TL-8, NP-0, LP-0. N. chrosomus. 4-1(15), 6-2(16), 3-1(17), 3-1(18), TN-16, TL-5, NP-0, LP-0. N. chrysocephalus. 1-1(1) , 27-11(2), 2-2(6), 6-3(7), 12-4(11), 7-3(12), 4-1(13), 4-1(14), 6-3(15), 6-2(16), 1-1(17), 3-1(18) , 4-1(19) , TN-83, TL-34, NP-4, LP-3. N. cummingsae. 4-1(27), TN-4, TL-1, NP-0, LP-0. N. emilae. 1-1(1), 1-1(3), 2-2(6), 3-1(7), 5-2(10), 5-2(11), 8-3(12), 3-1(13), 3-1(14), 7-3(15), 3-1(16), 7-3(20), 2-1(22), LP-3. N. euryzonus. 3-1(23) , 3-1(27), TN-56, TL-24, NP-4, 7-2(27), TN-7, TL-2, NP-0, LP-0. DISTRIBUTION OF THE GENUS ERGASILUS 13 DISTRIBUTION OF THE G;ENUS ERGASILUS 1 N. gibbsi. 3-1(20), TN-3, TL-1, NP-0, N. hypselopterus. 6-2(9), 6-2(10), 2-1(12), 3-1(13), 9-3(21), 8-4(22), 5-2(23), 6-2(24), 10-3(25), 7-1(27), TN-62, TL-21, NP-0, LP-0. N. hypsilepis. 7-2(26), 4-1(27), TN-li, TL-3, NP-i, LP-1. N. lirus. 2-1(14), 2-1(18), TN-4, TL-2, NP-0, N. longirostris. 3-1(1), 4-2(2), 4-2(3), 2-1(4), 13-5(5), 18-7(6), 49-14(7), 2-1(8), 6-2(21), 10-3(22), 6-2(23), 6-2(24), 9-3(25), 3-1(26), 6-2(27), TN-141, TL-48, NP-3, LP-1. N. edwardraneyi. 6-2(11), 6-2(12), 3-1(14), 7-2(15), 4-1(16), 6-2(20), TN-32, TL-10, NP-i, LP-1. N. manculatus. 3-1(2), 3-2(10), 2-1(15), 1-1(23), 5-2(25), 1-1(27), TN-15, TL-8, NP-0, LP-0. N. petersoni. 3-1(10), 6-2(23), 2-2(24), 3-1(25), 3-1(28), TN-17, TL-7, NP-0, LP-0. N. roseipinnis. 6-2(1), 14-6(2), 3-1(3), 7-2(4), 39-14(5), 30-10(6), 37-12(7), 6-2(8), 2-1(9), 2-1(10), TN-146, TL-51, NP-5, LP-2. N. shumardi. 6-2(10), 6-2(11), 3-1(15), TN-iS, TL-5, NP-0, LP-0. N. signipinnis. 3-1(2), 3-1(3), 8-3(4), 7-3(5), 12-4(7), 9-3(8), 3-1(9), 3-1(10), 4-2(11), 1-1(13), 6-2(21), 6-2(22), 6-2(23), 6-2(24), 6-2(25), TN-83, TL-30, NP-0, LP-0. LP-0. LP-0. N. stilbius. 7-3(11), 5-2(12), 9-3(15), 3-1(16), 3-1(17), 2-1(18), 3-1(19), N. texanus. 2-2(1), 11-4(2), 5-2(3), 5-2(4), 33-14(5), 14-6(6), 24-8(7), 5-3(8), 3-1(10), 9-3(11), 10-4(12), 3-1(13), 3-1(14), 3-1(15), 3-1(16), 5-2(20), 6-2(21), 7-2(22), 6-2(23), 3-1(24), 11-4(25), 1-1(20), TN-33, TL-13, NP-0, LP-0. 3-1(26), 7-2(27), TN-181, TL-69, NP-iS, LP-9. N. trichroistius. N. urartosco pus. 3-1(14), 3-1(16), 2-1(17), 7-3(18), TN-iS, TL-6, NP-0, LP-0. LP-0. N. venustus. 3-1(1), 2-1(16), 3-1(20), TN-5, TL-2, NP-0, 27-9(2), 8-3(3), 6-2(4), 13-5(5), 27-11(6), 24-11(7), 2-1(8), 3-1(10), 9-3(11), 6-2(12), 3-1(13), 3-1(14), 9-3(15), 3-1(16), 2-1(17), 3-1(18), 3-1(19), 5-3(20), 3-1(21), 9-3(22), 4-2(23), 5-2(24), 14 ALABAMA AGRICULTURAL EXPERIMENT STATION 11-4(25), 3-1(26), 3-1(27), TN-197, TL-75, NP-7, LP-3. N. volucellus. 3-1(2), 6-2(11), 4-2(12), 3-1(13), 3-1(14), 2-1(15), 3-1(16), 3-1(17), 3-1(20), TN-30, TL-11, NP-0, LP-0. N. welaka. 3-1(2), 7-4(7), TN-10, TL-5, NP-0, LP-0. N. xaenocephalus. 3-1(16), 3-1(17), 3-1(18), 3-1(20), TN-12, TL-4, NP-0, LP-0. N. zonistius. 5-2(26), 4-2(27), TN-9, TL-4, NP-1, LP-1. Notropis sp. cf. bellus. 3-1(15), 2-1(26), 2-1(27), TN-7, TL-3, NP-0, LP-0. Notropis sp. cf. longirostris. 11-4(11), 6-3(12), 3-1(13), 3-1(14), 4-2(15), 4-2(16), 2-1(19), 6-2(20), 2-1(22), TN-41, TL-17, NP-0, LP-0. Phenacobius catostomus. 3-1(16), 3-1(18), 1-1(19), TN-7, TL-3, NP-0, LP-0. Pimephales notatus. 2-2(1), 1-1(5), 1-1(6), 9-3(11), 4-2(12), 3-1(13), 3-1(14), 5-2(15), 1-1(16), 3-1(21), TN-32, TL-15, NP-0, LP-0. P. promelas. 3-1(21), TN-3, TL-1, NP-0, LP-0. P. vigilax. 3-2(1), 15-6(5), 8-2(6), 3-1(7), 9-3(11), 3-1(12), 2-1(13), 3-1(14), 3-1(15), 3-1(16), 2-1(18), 6-2(19), 3-1(20), TN-63, TL-23, NP-4, LP-2. Rhinichthys atratulus. 5-2(17), 3-1(18), TN-8, TL-3, NP-0, LP-0. Semotilus atromaculatus. 4-2(1), 1-1(6), 12-5(11), 1-1(12), 3-1(13), 3-1(14), 9-3(15), 3-1(16), 6-2(17), 6-3(18), 1-1(19), 6-2(20), 6-2(22), TN-61, TL-25, NP-8, LP-4. CATOSTOMIDAE Carpiodes sp. 2-1(5), 6-2(6), 2-1(16), 1-1(22), 6-2(25), TN-17, TL-7, NP-0, LP-0. C. cyprinus. 2-1(10), 2-2(11), TN-4, TL-3, NP-0, LP-0. C. velifer. 2-1(1), 2-1(6), 12-4(11), 1-1(14), 2-2(15), 6-3(16), 1-1(20), 3-1(22), TN-29, TL-14, NP-0, LP-0. C. elongatus. 3-1(1), 1-1(11), 1-1(16), TN-5, TL-3, NP-0, LP-0. Erimyzon oblongus. 2-1(1), 1-1(6), 1-1(8), 5-3(11), 5-3(15), 2-1(18), 2-1(20), 1-1(26), TN-19, TL-12, NP-2, LP-2. E. sucetta. 3-2(1), 5-5(2), 6-3(3), 1-1(4), 1-1(5), 3-3(6), 4-3(7), 2-1(10), 6-5(11), 4-3(12), 1-1(15), 1-1(16), 3-2(20), 1-1(22), 11-5(25), 2-1(27), 1-1(28), DISTRIBUTION OF THE GENUS ERGASI'LUS 15, DISTRIBUTION OF THE GENUS ERGASILUS 1 TN-55, TL-39, NP-5, LP-4. E. tenuis. 4-1(2), 5-3(3), 1-1(4), 9-6(5), 1-1(6), 15-8(7), 9-4(8), 7-3(9), 5-2(10), 3-2(11), 7-3(12), 3-1(13), 2-1(14), 4-2(16), 8-3(21), 3-1(23), 6-2(24), 4-2(25), TN-96, TL-46, NP-4, LP-4. Hypentelium etowanum. 9-6(11), 4-2(12), 3-1(13), 10-4(15), 5-2(16), 4-2(17), 3-1(18), 3-2(19), 1-1(20), TN-42, TL-21, NP-i, LP-1. H. nigricans. 11-6(2), 4-2(3), 1-1(4), 6-5(5), 8-7(6), 26-15(7), TN-56, TL-36, NP-3, LP-3. Ictiobus bubalus. 1-1(2), 3-1(6), 5-3(11), 1-1(16), 2-2(18), TN-12, TL-8, NP-i, LP-1. Minytema melanops. 1-1(1), 1-1(2), 1-1(3), 1-1(4), 9-7(5), 1-1(6), 1-1(8), 7-3(11), 1-1(14), 2-2(15), 1-1(16), 1-1(17), 2-2(18), 2-1(19), 2-2(20), 1-1(21), 2-1(22), 4-2(23), 3-1(24), 4-2(25), 2-1(26), 9-2(27), TN-58, TL-36, NP-10, LP-6. Moxostoma carinatun. 1-1(2), 1-1(6), 1-1(15), 2-1(16), TN-5, TL-4, NP-0, LP-0. M. duquesnei. 3-2(15), 1-1(16), 2-2(17), 3-2(18), 1-1(19), TN-10, TL-8, NP-0, LP-0. M. erythrurum. 2-1(11), 1-1(13), 2-1(14), 3-2(15), 1-1(16), 1-1(17), 1-1(19), 1-1(20), TN-12, TL-9, NP-0, LP-0. M. lachneri. 1-1(26), 1-1(27), TN-2, TL-2, LP-0. M. poecilurum. 2-2(1), 2-1(2), 4-3(3), 2-1(4), 12-9(5), 3-3(7), 4-2(8), 2-2(11), 2-1(12), 1-1(13), 2-1(14), 5-4(15), 5-3(16), 2-2(18), 2-2(19), 1-1(20), 3-2(21), NP-0, 2-2(22), 1-1(23), LP-3. 5-2(24), 6-4(25)) TN-68, TL-49, NP-4, Moxostoma sp. cf. poecilurum. 1-1(26), 7-3(27), TN-8, TL-4, NP-7, LP-3. ICTALURIDAE Ictalurus brunneus. 3-1(26), 3-1(27), TN-6, TL-2, NP-2, LP-l. I. catus. 1-1(27), TN-i, TL-i, NP-0, LP-0. 6-2(18), I. furcatus. 1-1(3), 3-1(9), 2-1(10), 2-1(11), 1-1(15), TN-iS, TL-7, NP-4, LP-3. I. melas. 2-1(1), 1-1(2), 1-1(3), 2-1(5), 1-1(6), 3-3(11), 1-1(12), 2-1(13), 3-1(15), 3-1(16), 1-1(17), 1-1(19), 1-1(20), 3-2(25), TN-25, TL-17, NP-i, LP-1. 16 ALABAMA AGRICULTURAL EXPERIMENT STATION I. natalis. 1-1(1), 1-1(2), 3-2(3), 1-1(4), 8-3(5), 2-2(7), 3-2(11), 1-1(12), 1-1(13), 2-2(15), 3-1(16), 1-1(17), 1-1(18), 3-2(19), 1-1(20), 3-1(21), 1-2(23), 2-2(24), 8-5(25), 1-1(26), 1-1(27), 1-1(28), TN-49, TL-35, NP-3, LP-3. I. nebulosus. 1-1(2), 1-1(4), 3-2(19), 5-3(25), 2-1(26), 4-2(27), TN-16, TL-10, NP-7, LP-6. I. platycephalus. 2-1(26), TN-2, TL-1. I. punctatus. 1-1(1), 2-1(2),5-3(3), 1-1(4), 5-3(5), 6-5(6), 2-2(7), 6-3(10), 9-3(11), 5-3(12), 1-1(13), 1-1(14), 8-4(15), 7-3(16), 6-3(18), 1-1(19), 2-2(20), 1-1(22), 1-1(23), 5-2(24), 1-1(25), 3-1(26), 2-2(27), TN-81, TL-48, NP-15, LP-10. Noturus sp. 8-6(5), TN-8, TL-6, NP-0, LP-0. N. funebris. 3-2(2), 2-1(3), 1-1(4), 1-1(5), 1-1(6), 10-5(7), 2-2(8), 3-1(9), 1-1(10), 6-3(11), 4-2(12), 3-1(13), 3-1(14), 6-2(15), 4-3(16), 2-2(19), 1-1(21), 2-1(22), 3-1(23), 3-1(25), TN-61, TL-33, NP-0, LP-0. N. gyrinus. 2-1(3), 1-1(4), 5-3(5), 3-1(6), 2-1(7), 4-2(8), 7-6(11), 1-1(12), 2-1(14), 8-3(15), 3-1(20), 3-2(25), TN-41, TL-23, NP-0, LP-0. N. leptacanthus. 2-2(2), 9-3(3), 3-2(4), 8-3(6), 9-5(7), 4-2(8), 8-3(11), 2-1(12), 2-1(13), 4-2(15), 5-3(16), 2-1(20), 1-1(21), 3-1(22), 2-1(23), 6-2(24), 4-2(25), 3-1(26), 9-3(27), TN-86, TL-39, NP-0, LP-0. N. miurus. 7-5(2), 1-1(2), TN-8, TL-6, NP-0, LP-0. N. munitus. 9-3(11), 3-1(12), 1-1(16), TN-13, TL-5, NP-0, LP-0. N. nocturnus. 2-1(1), 6-3(4), 10-5(6), 5-3(7), 2-1(13), 2-2(16), TN-27, TL-15, NP-1, LP-1. Pylodictisolivaris. 1-1(1), 5-2(4), 3-3(5), 2-1(6), 1-1(7), 1-1(8), 1-1(11), 1-1(12), 5-2(15), 1-1(16), 2-1(18), 3-2(19), TN-26, TL-17, NP-4, LP-2. ARIIDAE Bagre marinus. 1-1(3), 2-2(9), 2-1(29), TN-5, TL-4, NP-6, LP-5. Arius felis. 2-1(3), 6-2(9), 3-1(22), 6-2(29), TN-17, TL-6, NP-0, LP-0. AMBLYOPSIDAE Typhlichthys subterraneus. see text, NP-0, LP-0. DISTRIBUTION OF THE GENUS ERGASILUS 17 DISTRIBUTION OF THE G;ENUS ERGASI LUS 1 APHREDODERIDAE Aphredoderus sayanus. 5-2(1), 6-2(2), 7-5(3), 8-8(5), 8-3(6), 5-5(7), 3-2(8), 3-1(10), 7-3(11), 2-1(13), 2-1(14), 5-2(15), 1-1(16), 2-1(20), 4-2(22), 5-2(23), 2-2(24), 5-3(25), 1-1(27), TN-81, TL-47, NP-5, LP-5. BELONLDAE Strongylura marina. 1-1(2), 2-1(3), 1-1(6), 1-1(7), 6-2(9), 1-1(15), 1-1(20), 4-2(22), 2-1(25), 1-1(28), 2-1(29), TN-22, TL-13, NP-0, LP-0. CYPRINODONTIDAE Cyprinodon variegatus. 3-1(7), 12-4(9), 4-1(10), 5-1(21), 21-7(22), 1-1(28), TN-46, TL-15, NP-8, LP-4. Fundulus chrysotus. 4-2(3), 2-1(10), 2-1(21), 4-2(23), 7-2(25), 3-1(28), TN-22, TL-9, NP-0, LP-0. F. cingulatus. 3-1(9), 2-1(10), 3-1(23), 2-1(24), 11-4(25), TN-21, TL-8, NP-0, LP-0. F. confluentus. 3-1(9), 5-2(10), 6-2(21), TN-14, TL-5, NP-0, LP-0. F. grandis. 3-2(3), 9-3(7), 4-2(9), 1-1(10), 3-1(21), 3-1(22), 2-1(28), 2-1(29), TN-27, TL-12, NP-7, LP-6. F. notatus. 6-3(11), TN-6, TL-3, NP-0, LP-0. F. notti. 1-1(2), 2-1(3), 5-2(4), 3-1(5), 3-2(6), 20-8(7), 2-1(8), 3-1(9), 3-1(10), 6-2(11), 6-2(12), 3-1(13), 1-1(14), 3-1(15), 1-1(16), 3-1(21), 3-1(22), 6-2(23), 1-1(27) , TN-81, TL-34, NP-0, LP-0. F. 3-2(24), 3-1(25), olivaceus. 6-5(2), 15-6(3), 6-2(4), 44-19(5), 13-6(6), 59-21(7), 3-1(8), 3-1(10), 11-5(11), 10-4(12), 3-1(13), 3-1(14), 7-3(15), 9-3(16), 2-1(17), 3-1(18), 1-1(20), 1-1(21), 6-2(22), 3-1(23), 6-2(24), 3-1(25), 7-2(27), TN-224, TL-90, NP-2, LP-2. F. pulvereus. 6-1(3), 2-1(7), TN-8, TL-2, NP-i, LP-1. F. similis. 1-1(3), 2-1(7), 7-2(9), 9-3(23), 4-1(28), 5-2(29), TN-28, TL-10, NP-8, LP-5. 3-1(26) , TN-22, TL-9, NP-0, LP-0. 3-2(19), 2-1(20),1 F. stellifer. 2-1(13), 4-1(14), 2-1(16), 3-1(17), 3-1(18), Leptolucania ommata. 1-1(22), 8-3(23), 2-1(25), TN-il, TL-5, NP-0, LP-0. 1-1(25), 6-2(28), Adinia xenica. Lucania parva. 3-2 (4), 5-1(9 ), 6-2(10) ,3-1(22 ), 6-2 (23), TN-30, TL-11, NP-4, LP-3. TN-7, TL-4. 2-1(9), 3-2(22), 2-1(29), 18 ALABAMA AGRICULTURAL EXPERIMENT STATION 18 ALABAMA AGRICULTURLEPRMN TTO POECILIDAE Gambusia affinis. 2-1(1), 2-1(2), 13-5(3), 15-6(5), 3-2(6), 18-9(7), 3-1(8), 5-2(9), 6-3(10), 9-3(11), 6-2(12), 3-1(13), 3-1(14), 12-4(15), 6-2(16), 5-2(17), 3-1(18), 4-2(20), 1-1(21), 8-3(22), 3-1(23), 3-1(24), 7-3(25), 3-1(26), 3-1(27), 2-2(28), TN-148, TL-61, NP-i, LP-1. Hetcrardria formosa. 9-3(3), 3-1(4), 5-2(25), 3-1(28), TN-20, TL-7, NP-O, LP-0. Poecilia latipinna. 1-1(3), 3-1(4), 6-2(7), 8-3(9), 3-1(10), 6-2(22), 4-1(28), 3-1(29), TN-34, TL-12, NP-6, LP-5. ATHERINIDAE Labidesthes sicculus. 3-1(1), 5-2(2), 5-3(3), 5-2(4), 5-2(5), 3-1(6), 16-7(7), 3-2(8), 3-2(10), 6-2(11), 3-1(12), 3-1(13), 7-3(15), 2-1(16), 2-1(20), 5-2(21), 4-1(22), 4-1(23), 3-1(24), 10-4(25), 6-2(27), TN-103, TL-42, NP-i, LP-1. Menidia audens. 2-1(2), 4-2(3), 1-1(9), TN-7, TL-4, NP-0, LP-0. M. beryllina. 3-1(3), 3-1(7), 6-2(9), 6-2(10), 4-1(22), 6-2(23), 3-1(25), 4-1(28), TN-35, TL-11, NP-8, LP-3. COTTIDAE Cottus carolinae. 2-1(12), 3-1(13), 3-1(14), 3-1(15), 3-1(16), 6-2(17), 3-1(18), 4-2(19), 1-1(20), TN-28, TL-11, NP-0, LP-0. C. pygmaeus. 3-1(17), TN-3, TL-1, NP-0, LP-0. SYNGNATHLDAE Syngnathus loucisianae. 4-2(4), TN-4, TL-2, NP-0, LP-0. Synguathus sp. 1-1(7), 1-1(28), 2-1(29), TN-4, TL-3, NP-0, LP-0. PERCJCHTHYIDAE Mo'rone chrysops. 1-1(15), 3-2(18), 2-1(19), 2-1(27), TN-8, TL-5, NP-i ,LP-1. M. mississippiensis. 1-1(3), 1-1(7), 1-1(10), TN-3, TL-3, NP-2, LP-2. M. saxatilis. 3-2(10), TN-3, TL-2, NP-i, LP-1. DISTRIBUTION OF THE GENUS DISTRIBUTION OF THE G'ENUS ERGASILUS ERGASILUS 19 1 ELASSOMJDAE Elassoma evergladei. 3-1(9), 2-1(13), 1-1(21), 1-1(24), 2-1(25), TN-9, TL-5, NP-0, LP-0. E. okefenokee. 6-2(25), TN-6, TL-2, NP-0, LP-0. E. zonatum. 3-1(1), 2-1(2), 6-4(3), 4-2(4), 6-4(5), 1-1(6), 10-5(7), 8-4(8), 2-1(10), 6-2(11), 6-2(12), 2-1(13), 1-1(14), 5-2(15), 3-1(16), 3-1(20), 5-2(21), 3-1(22), 2-1(23), 1-1(24), 5-2(25), 3-1(27), TN-87, TL-41, NP-0, LP-0. CENTRARCHIDAE Ambloplitis rupestris. 3-1(1), 2-2(2), 3-3(3), 2-2(4), 1-1(5), 5-4(6), 2-2(7), 4-2(8), 5-3(11),2-1(13), 1-1(14), 3-2(15), 5-2(16), 1-1(19), 1-1(20), 1-1(21), 3-2(22), 1-1(23), 3-1(24), 5-2(25), TN-53, TL-35, NP-0, LP-0. 1-1(6), Centrarchus macropterus. 2-1(1), 1-1(2), 2-1(7), 2-2(11), 1-1(13), 3-1(15), 1-1(20), 3-1(22), 7-4(25), 3-1(27), TN-31, TL-19, NP-4, LP-4. Enneacanthus gloriosus. 3-1(10), 1-1(21), 3-1(23), 3-1(25), TN-10, TL-4, NP-0, LP-0. Lepomis auritus. 1-1(1), 3-3(5), 7-6(7), 4-2(19), 1-1(20), 3-1(26), 7-2(27), 2-1(28), TN-28, TL-17, NP-5, LP-2. L. cyanellus. 5-5(1), 3-3(2), 3-2(3), 1-1(4), 19-9(5), 1-1(6), 6-3(7), 4-3(11), 1-1(12), 2-1(13), 3-1(14), 10-4(15), 2-1(16), 6-3(17), 3-2(18), 1-1(19), 1-1(22), 5-4(5), TN-79, TL-45, NP-6, LP-4. 3-1(25) , L. gulosus. 1-1(1), 2-2(2), 11-6(3), 1-1(4), 3-1(26),12-1(27), 4-3(6), 2-7(7), 2-1(8), 2-1(9), 2-1(10), 5-4(11), 2-1(13), 2-1(14), 6-3(15), 1-1(17), 1-1(18), 3-2(19), 2-2(20), 1-1(21), 2-1(22), 2-1(23), 2-2(24), 13-4(25), 2-1(26), 1-1(27) , 1-1(28) , TN-85, TL-60, NP-9, LP-8. L. humilis. 3-1(11), 12-10(5),- 1-1(14), 1-1(15), TN-5, TL-3, NP-0, LP-0. L. macrochirus. 4-3(1), 12-7(2), 14-8(3), 6-2(4), 21-12(5), 8-5(6), 16-12(7), 4-2(8), 11-3(9), 3-1(10), 12-6(11), 3-1(12), 3-1(13), 3-1(14), 11-4(15), 4-2(16), 6-2(17), 3-1(19), 2-1(20), 5-2(21), 4-1(22), 4-2(23), 5-2(24), 15-3(25), 2-1(26), 4-2(27), 1-1(28), TN-186, TL-88, NP-34, LP-17. L. marginatus. 1-1(3), 1-1(4), 6-2(5), 2-2(7), 1-1(8), 2-1(9), 2-1(12), 3-1(13), 2-1(15), 3-1(21), 1-1(22), 20 20 ALABAMA AGRICULTURAL EXPERIMENT STATION 4-2(23), 1-1(24), 3-1(25), 2-1(27), TN-34, TL-18, NP-7, LP-3. L. megalotis. 3-2(1), 4-3(2), 14-7(3), 6-2(4), 33-16(5), 7-4(6), 23-13(7), 4-3(8), 9-3(11), 3-2(12), 3-1(13), 2-1(14), 2-1(15), 6-3(16), 2-1(17), 4-2(18), 2-1(19), 2-2(20), 3-2(21), 5-2(22), 3-2(23), 4-3(24), 6-3(25), 1-1(26), 2-1(27), 2-1(28), TN-155, TL-82, NP-iS, LP-10. L. microlophus. 2-1(1), 2-2(3), 5-3(5), 1-1(7), 5-1(9), 4-2(10), 3-3(11), 1-1(12), 2-2(14), 5-4(15), 2-1(16), 1-1(17), 1-1(18), 2-1(20), 2-2(22), 5-2(23), 11-2(25), 1-1(26), 7-3(27), TN-62, TL-34, NP-13, LP-10. L. punctatus. 2-2(2), 3-2(3), 4-3(4), 7-6(5), 2-1(6), 5-3(7), 5-2(8), 4-1(10), 4-2(11), 4-3(12), 2-1(13), 3-1(14), 3-2(15), 1-1(16), 3-2(20), 2-1(21), 5-2(23), 3-2(24), 9-3(25), 3-1(26), 3-1(27), 7-2(28), TN-84, TL-44, NP-22, LP-17. Micropterus coosae. 6-4(15), 2-1(16), 2-2(17), 1-1(18), 2-2(19), 1-1(20), 1-1(26), TN-iS, TL-12, NP-0, LP-0. M. punctulatus. 3-1(1), 6-5(2), 7-4(3), 3-1(4), 15-11(5), 15-8(6), 16-11(7), 1-1(8), 3-1(11), 8-4(12), 2-1(13), 5-3(15), 6-2(16), 2-1(17), 1-1(18), 4-2(19), 4-2(20), 2-1(21), 4-2(23), 4-3(24), 6-3(25), TN-117, TL-68, NP-10, LP-8. M. salmoides. 1-1(1), 2-2(2), 5-4(3), 1-1(4), 8-8(5), 3-2(6), 3-2(7), 5-3(8), 6-1(9), 5-2(10), 7-5(11), 3-2(12), 2-1(13), 4-2(14), 5-3(15), 3-1(16), 2-1(17), 9-3(18), 1-1(19), 3-3(20), 5-2(21), 7-5(22), 6-2(23), 1-1(24), 6-2(25), 1-1(26) , 5-2(27), TN-109, TL-63, NP-28, LP-16. Pomoxis annularis. 4-3(1), 1-1(2), 3-3(5), 4-1(9), 5-2(11), 1-1(13), 2-1(14) 3-1(15), 1-1(17), 1-1(19), 2-2(20), 4-1(25), TN-31, TL-18, NP-8, LP-6. P. nigromaculatus. 2-1(1), 2-2(5), 1-1(6), 4-2(10), 4-2(11), 2-1(14), 4-2(15), 2-1(16), 1-1(17), 2-1(19), 4-3(20), 3-1(22), 1-1(25), 5-2(26), 1-1(27), TN-38, TL-22, NP-6, LP-6. PERCIDAE Ammocrypta asprella. 6-2(11), NP-0, LP-0. 3-1(16), TN-9, TL-3, A. beani. 1-1(1), 4-2(2), 3-2(3), 3-1(4), 1-1(S), 12-6(6), 17-9(7), 5-3(11), 6-2(12), 1-1(14), 2-1(15), 2-1(16), 2-1(20), 6-2(21), 9-3(22), 6-2(23), 6-2(24), 7-3(25), DISTRIBUTION OF THE GENUS ERGASILUS 21 DISTRIBUTION OF THE GENUS ERGASILUS 2 TN-93, TL-43, NP-0, LP-0. A. vivax. 2-1(1), 1-1(2), 1-1(4), 2-2(5),,4-2(6), 8-3(11), 6-2(12), 3-1(13), 3-1(14), 1-1(15), 1-1(16), 3-1(20), TN-35, TL-17, NP-0, LP-0. TL-4, Etheostoma chiorosomum. 2-1(1), 4-3(5), NP-0, LP-0. F. coosae. 6-2(17), 5-2(18), TN-i1, TL-4, NP-0, LP-0. F. davisoni. 2-1(22), 1-1(23), 5-2(26), TN-8, TL-4, NP-0, LP-0. F.ditrema. 2-1(17), TN-2, TL-1, NP-0, LP-0. E. edwini. 1-1(21), 1-1(22), 3-1(23), 5-2(24), 5-2(25), 6-2(27), TN-21, TL-9, NP-0, LP-0. E. fusiforme. 2-1(3), 1-1(4), 1-1(8), 2-1(10), 7-3(12), 1-1(22), 1-1(23), 3-1(25), 3-1(26), TN-21, TL-11, NP-0, LP-0. E. 7-3(11), TN-7, TL-3, NP-0, LP-0. E. jordani. 1-1(14), 8-3(15), 3-1(16), 3-1(17), 2-1(19), TN-17, TL-7, NP-0, LP-0. F. nigrum. 9-3(11), 2-1(12), 3-1(13), 2-1(14), 1-1(15), 3-1(20), TN-20, TL-8, NP-0, LP-0. E. nuchale. 2-1(15), TN-2, TL-1, NP-0, LP-0. E. okaloosae. 6-2(27), TN-6, TL-2, NP-0, LP-0. F.parvipinne. 1-1(6), 8-3(11), 5-2(12), 1-1(13), 3-2(14), 6-2(15), 3-1(20), 2-1(22), 2-1(25), 1-1(27), TN-32, TL-15, NP-0, LP-0. E. proeliare. 2-1(1), 1-1(10), 11-4(11), 6-2(12), 2-1(13), 2-1(14) , 2-1(15) , 1-1(16), TN-27, TL-12, NP-0, LP-0. E. rupestre. 9-3(11), 3-1(12), 3-1(13), 3-1(14), 6-2(15), TN-6, histrio. 6-2(16), 3-1(20), TN-33, TL-11, NP-0, LP-0. E. stigmaeurn. 1-1(1), 4-1(2), 3-1(3), 6-3(5), 2-1(6), 24-14(7), 1-1(8), 9-4(11), 6-2(12), 3-1(13), 6-2(15), 2-1(16), 3-1(17), 2-1(18), TN-76, TL-37, NP-0, E. swami. 3-2(2), 5-2(3), 6-2(4), 15-10(5), 1-1(6), 16-8(7), 5-2(8), 1-1(10), 7-4(11), 4-2(12), 1-1(13), 2-1(14), 6-2(15), 6-2(16), 1-1(20), 6-2(22), 2-1(23), LP-0. 2-1(20), 1-1(21), 1-1(23), 2-2(24), 6-2(25), 2-1(27), TN-97, TL-49, NP-0, LP-0. E. whippici. 1-1(1), 1-1(2), 1-1(6), 9-3(11), 8-3(12), 2-1(13), 3-1(14), 6-2(15), 3-1(16), 3-1(18), 2-1(20), TN-39, TL-16, NP-0, LP-0. E. zonale. 1-1(1), 3-1(2), 5-2(3), 1-1(5), 2-1(6), 5-2(7), 22 ALABAMA AGRICULTURAL EXPERIMENT STATION 22 ACLABAMA AGRICULTURLEPRMN TTO TN-17, TL-8, NP-0, LP-0. E. zoniferum. 1-1(12), 3-1(14), TN-4, TL-2, NP-0, LP-0. E. (Ulocentra) spl. 12-4(11), 5-2(12), 3-1(15), 6-2(16), 4-2(22), 6-2(23), 6-2(24), 6-2(25), TN-48, TL-17, NP-O, LP-0. E. (Ulocentra) sp. 15-2(15), TN-iS, TL-2, NP-0, LP-0. E. (Ulocentra) sp. 6-2(19), TN-6, TL-2, NP-0, LP-0. Perca flavescens. 3-1(26), 2-1(27), TN-5, TL-2, NP-0, LP-0. Percina caprodes. 5-2(1), 2-1(3), 1-1(4), 4-2(5), 6-4(6), 3-2(7), 3-1(8), 1-1(11), 11-4(15), 2-1(16), 2-1(17), 3-1(18), 3-1(19), 1-1(20), 1-1(22), 1-1(25), TN-49, TL-25 NP-i, LP-1. P. copelandi. 3-1(15), 1-1(16), TN-4, TL-2, NP-0, LP-0. P. lenticula. 3-1(1), 1-1(6), 2-2(16), TN-6, TL-4, NP-0, LP-0. P. maculata. 1-1(11), 1-1(12), 2-1(15), 1-1(16), TN-5, TL-4, NP-0, LP-0. P. nigrofasciata.11-7(2), 5-2(3), 6-2(4), 14-8(5), 15-6(6), 26-13(7), 6-2(8), 2-1(9), 2-1(10), 12-4(11), 3-2(12), 3-1(13), 11-4(15), 3-1(16), 4-2(17), 4-3(18), 12-4(25), 2-1(26), 3-1(27), TN-144, TL-65, NP-4, LP-3. P. palmaris. 3-1(17), 8-3(19), TN-il, TL-4, NP-0, LP-0. P. sciera. 3-3(7), 9-3(11), TN-12, TL-6, NP-0, LP-0. P. shumardi. 5-2(11), 1-1(13), 3-1(15), 1-1(16), TN-10, TL-5, NP-0, LP-0. P. uranidea. 1-1(1), 2-1(5), 1-1(6), 7-3(11), 1-1(12), 2-1(13 ), 2-1(22), TN-16, TL-9, NP-0, LP-0. P. (Alvordius) sp. 3-1(19), TN-3, TL-1, NP-0, LP-0. Stizostedion vitreum. 1-1 (1), 1-1(2), 2-1 (15 ), TN-5, TL-4, NP-0, LP-0. CARANGIDAE Caranx hippos. 1-1 (18), 1-1(7), 2-1(9), Oligoplites saurus. 3-1(4), 3-2(9), TN-3, TL-2, NP-i, LP-1. TN-6, TL-3, NP-0, LP-0. SPARIDAE Archosargus probatocephalus. 1-1(3), 1-1(23) , 4-2(22 ), 1-1(29), TN-7, TL-6, NP-0, 3-2(4), LP-0. 1-1(9) , Lagodon rhornboides. 1-1(3), 1-1(7), 2-1(9), 1-1(21), 2-1(25), 4-1(28), TN-iS, TL-8, NP-i, LP-1. DISTRIBUTION OF THE GENUS ERGASILUS 23 DISTRIBUTION OF THE GENUS ERGASILUS 2 SCIAENIDAE Aplodinotus grunniens. 1-1(1), 1-1(2), 4-3(11), 2-2(12), 4-2(16), 1-1(18), 2-2(20), TN-15, TL-12, NP-0, LP-0. Bairdiella chrysura. 2-1(9), 1-1(29), TN-3, TL-2, NP-0, LP-0. Gynoscion arenarius. 2-1(9), 1-1(22), TN-3, TL-2, NP-0, LP-0. C. nebulosus. 1-1(3), 1-1(21), 3-1(29), TN-5, TL-3, NP-0, LP-0. Leiostomus xanthurus. 2-1(2), 3-1(3), 3-1(7), 5-1(10), 1-1(21), 1-1(23), 1-1(25), 4-1(28), TN-20, TL-8,-NP-7, LP-4. Micropogon undulatus. 4-2(3), 4-1(10), 2-1(22), 1-1(28), TN-i1, TL-5, NP-0, LP-0. Pogonias cromis. 2-1(3), 2-2(9), TN-4, TL-3, NP-i, LP-1. Sciaenops ocellata. 1-1(3), 1-1(9), TN-2, TL-2, NP-0, LP-0. MUGILIDAE Mugil ceplhalus. 1-1(2), 12-5(3), 1-1(6), 2-2(7), 6-2(9), 4-2(10), 7-2(21), 5-1(22), 1-1(25), 4-1(28), TN-43, TL-18, NP-23, LP-13. M. curema. 3-1(4), 5-3(9), 2-1(29), TN-i0, TL-5, NP-4, LP-4. ELEOTRIDAE Dormitator maculatus. 2-1(3), 1-1(10), 1-1(23), TN-4, TL-3, NP-0, LP-0. Eleotris pisonis. 1-1(4), 1-1(23), 1-1(25), TN-3, TL-3, NP-0, LP-0. GOBIIDAE Gobionellus boleosoma. 3-1(9), 3-1(28), TN-6, TL-2, NP-0, LP-0. C. hastatus. 2-1(21), TN-2, TL-i, NP-0, LP-0. G. shufeldti. 1-1(9), 4-3(10), TN-5, TL-4, NP-0, LP-0. G. stigmaticus. 2-1(7), 3-1(28), TN-5, TL-2, NP-0, LP-0. Gobiosoma bosci. 2-1(9), 2-1(23), 2-2(28), 8-2(29), TN-14, TL-6, NP-3, LP-1. Microgobius gulosus. 1-1(9), 1-1(28), 3-1(29), TN-5, TL-3, NP-0, LP-0. 24 ALABAMA AGRICULTURAL EXPERIMENT STATION 24 ALABAMA AGRICULTUA XEIMN TTO BOTHIDAE Paralichthyslethostigma. 2-2(3), TN-2, TL-2, NP-O, LP-0. SOLEIDAE Archirus lineatus. 3-2(2), 1-1(4), 2-1(5), 2-2(6), 1-1(29), TN-9, TL-7, NP-3, LP-2. Trirectes maculatus. 1-1(3), 1-1(4), 2-1(9), 1-1(25), TN-5, TL-4, NP-O, LP-0. RESULTS Parasite Material and Respective Host The following listings give the parasite and the hosts from which the parasite species were recovered. The parentheses following the hosts include an initial number, which represents the number of parasite specimens recovered from a locality, and a following number, which represents the drainage basin of the collection. This latter number corresponds to the number given for the drainage basin in Figure 2. Information Ergasilus arthrosis Ictalurus furcatus (4-9), (5-18); I. punctatus (12-16), (112), (1-12), (3-12), (2-7), (1-1), (2-20), (26-18), (2-18); Fylodictis olivaris (19-18), (16-20); Aphredoderus sayanus (13); Labidesthes sicculus (1-10); Morone mississippiensis (153), (12-7); M. saxatilis (4-10); Lepomis gulosus (1-23), (410); L. macrochirus (16-23), (1-15), (34-9 ), (2-9 ), (4-22); L. marginatus (28-23 ), (7-21); L. megalotis (5-12); L. microlophus (2-12), (5-23), (9-23), (1-10), (7-7), (1-3); L. punctatus (8-23), (15-7), (8-7); Micropterus punctulatus (5-12), (2-12 ), (2-11), (1-15); M. salmoides (31-23 ), (9-23 ), (511), (8-12), (1-10), (7-9); Pomoxis nigrornaculatus (1-10), (2-10); Percina caprodes (1-7); Caranx hippos (1-7); Achirus lineatus (2-10), (5-3). Ergasilus caeruleus Aphredoderus sayanus (12-5 ), (11-7 ), (2-1), (11-1); Lepomis auritus (4-27), (10-27); L. (2-6); L. gulosus (6-5), (1-5), (8-2), (3-2); L. macrochirus (1-14), (5-12), (1-11), (2-22), (8-6), (7-5), (6-5), (1-5), (3-5), (2-27), (2-27); L. marginatus (3-14); L. megalotis cyanellus (3-12), (2-5), DISTRIBUTION OF THE GENUS ERGASILUS 25 DISTRIBUTION OF THE GENUS ERGASILUS 2 (1-5 ), (1-5 ), (6-5), (3-5), (16-7), (1-6 ), (6-20); L. microlophus (1-5), (1-5); L. punctatus (4-20), (2-11), (1-12), (19-16), (4-26), (3-5), (2-5), (3-5), (8-8), (5-7), (7-3), (2-4), (61-27); Pomoxis annularis (15-11), (8-11), (1-14), (23-20), (7-20), (4-5); P. nigromaculatus (8-20), (4-15), (1-11); Percina nigrofasciata (1-5), (6-5), (3-5). Ergasilus celestis Anguilla rostrata (4-3), (29-5), (3-5 ),(24-1), (7-20), (1-18), (1-25). Ergasilus centrarchidarum Morone chrysops (1-19); Centrarchus macropterus (1-27), (4-5), (1-1), (1-27); Lepomis cyanellus (2-5); L. gulosus (1-15); L. macrochirus (6-25), (2-27); L. punctatus (2-25); Micropterus punctulatus (1-19), (5-19), (2-5), (1-5); M. salmoides (7-25), (15-25), (1-11), (5-11), (1-11), (3-5), (22-5), (1-5), (1-27), (4-20), (6-20); Pomoxis annularis (1-20); P. nigromaculatus (2-20), (1-26). Ergasilus cerastes Ictalurus furcatus (1-3); I. punctatus (50-10). Ergasilus clupeidarum (1-8), Alosa chrysochioris (16-9); Dorosoma cepedianum (4-20); (1-27). D. petenense Ergasilus cyprinaceus Campostoma anomalum H. (1-11); winchelli (2-5); Nocomis leptocephalus Hybopsis storeriana (1-5); (2-20), (21-5); Notemigonus crysoleucas (1-5); Notropis baileyi (2-11), (212), (1-20), (3-20); N. chrysocephalus (5-5), (1-5), (7-5); N. longirostris (8-5); N. hypsilepis (1-27); N. roseipinnis (26), (1-5); N. texanus (1-5), (1-6), (1-20); N. venustus (1-5); N. zonistius (2-26); Pimephales vigilax (7-5); Semotilus atromaculactus (11-20), (6-20), (1-1), (1-6); (5-1), (3-2); I. rwbulosus (1-25), (1-22); Noturus nocturnus (1-1); Fundulus olivaceus (1-15), (1-6). Ictalurus natalis 26 ALABAMA AGRICULTURAL EXPERIMENT STATION 26 ALABMA AGRICULTUA XEIMN TTO Ergasilus elongatus Polyodon spathula (14-20). Ergasilus felichthys Bagre marinus (1-9), (1-9), (2-29), (1-29), (4-3). Ergasilus funduli Adinia xenica (2-9), (5-9); Cyprinodon variegatus (2-7), (12-21); Fundulus grandis (1-9), (1-22), (1-7), (1-7), (228); F. pulvereus (1-3); F. similis (1-22), (4-22), (2-29), (1-9), (1-9), (2-28); Lucania parva (1-23), (2-25), (2-22); Gambusia affinis (1-22); Poecilia latipinna (4-9), (1-9), (17), (1-7), (4-22); Menidia beryllina (6-9), (35-7), (4-3); Gobiosoma bosci (12-29). Ergasilus lanceolatus Hiodon tergisus (1-1). Ergasilus lizae Cyprinodon variegatus (2-22), (1-22); Fundulus grandis (1-10); Lepomis macrochirus (3-9); Leiostomus xanthurus (5-7), (1-21), (15-3), (11-10); Lagodon rhomboides (1-7); Mugil cephalus (1-3), (5-9), (1-25), (1-22), (1-7), (3-3); Mugil curema (1-29), (1-4), (5-9). Ergasilus megaceros Dorosoma cepedianum (1-20); Esox americanus (11-5), (7-12), (7-14); E. niger (1-5); Hybognathus nuchalis (1-6), (1-12); Hybopsis storeriana (1-12), (4-5); Pimephales notatus (4-20); P. vigilax (4-13); Notropis bellus (6-20); N. edwardraneyi (2-12); N. emilae (2-12), (9-11), (3-7); N. roseipinnis (2-5); N. shumardi (8-11); N. texanus (5-5), (7-5), (3-4), (2-1), (5-22), (2-20), (2-14); N. venustus (2-15), (2-4), (3-5); Erimyzon oblongus (50-1), (9-26); E. sucetta (200+ -7), (1-5); Moxostoma sp. of. poecilurum (2-27); Ictalurus melas (25-20). DISTRIBUTION OF THE GENUS ERGASILUS 27 DISTRIBUTION OF THE GENUS ERGASILUS 2 Ergasilus tenax Esox niger (1-20); Lepomis gulosus (2-5), (1-5); L. macrochirus (9-5), (2-5), (5-5); L. megalotis (4-5), (62-5), (65), (3-6); L. microlophus (1-5), (13-5); L. (8-5), (1-6); Pomoxis annularis (4-11), (3-14); P. (6-10), (2-20). punctatus nigromaculatus Ergasilus versicolor Lepisosteus osseus (1-18); Erimyzon oblongus (1-27); E. sucetta (1-7), (5-25); E. tenuis (18-5), (5-25); bubalus (1-11); Minytream melanops (2-26), (4-22), (2-5), (28-26), (100's-27), (25-27); Moxostoma sp. cf. poecilurum (30-26), (30-27); Moxostoma poecilurum (2-8), (3-4), (422); Ictalurus natalis (7-25); I. nebulosus (21-25), (37-26), (10-25), (7-27); Mugil cephalus (6-22), (14-3), (1-9), (8-7), (7-7), (11-3), (6-3), (5-3), (8-3), (1-2); Pogonias cromis (1-9). Ictiobus Ergasilus wareaglei Hypentelium etowanum (5-16); H. nigricans (14-5), (8-5), (2-6). Ergasilus sp. cf. lizae (Form A) Mugil cephalus (8-3), (4-9), (1-25), (3-7); Mugil curema (2-9), (1-9). Ergasilus sp. cf. lizae (Form B) Mugil cephalus (1-9). Ergosilus Material Accumulated during Study from Outside of Study Area After the host names below, information is abbreviated as: (Number of parasite specimens - drainage state - county.) Ergasilus qrthrosis caeruleus Morone chrysops (6-Mississippi, Miss.-Warren). Ergasilus Lepomis cyanellus (1-Big Black, Miss.-Hinds); L. macro- 28 ALABAMA AGRICULTURAL EXPERIMENT STATION chirus (1-Tennessee, Miss.-Alcorn); L. punctatus (4-Atlantic, Fla.-Palm Beach). Ergasilus centrarchidarum Lepomis auritus (1-U. Chattahoochee, Ala.-Lee); Micropterus salmoides (4-Apalachicola, Fla.-Jackson). Ergasilus cerastes Ictalurus catus (25-Ashley, S.C.-Charleston). Ergasilus clupeidarum I. nebulosus (20-Potomac, Unk.). Ergasilus cyprinaceus Notropis atherinoides (1-Mississippi, Miss.-Cahoma). Ergasilus funduli Fundulus heteroclitis (4-Sapelo Is., Ga.-Mclntosh); Mugil cephalus (4-Sapelo Is., Ga.-Mclntosh). Ergasilus labracis Morone saxatilis (Many-Cooper, S.C.-Berkeley). Ergasilus lizae Mugil cephalus (6-Sapelo Is., Ga.-Mclntosh). Ergasilus megaceros Cycle ptus elorngatus (2-Mississippi, Miss.-Warren) ; Erimyzon sucetta (6-Big Black, Miss.-Hinds). Ergasilus versicolor Erimyzon sucetta (3-Ochiockonee, Fla.-Leon) ; Minytrema melanops (1-Flint, Ga.-Crisp) ; Mugil cephalus (3-Sapelo Is., Ga.-Mclntosh). Ergasilus sp. cf. lizae Form B Mugil cephalus (6-Sapelo Is., Ga.-Mclntosh). DISTRIBUTION OF THE GENUS ERGASILUS DISTRIBUTION OF THE GENUS ERGASILUS 29 29 DISCUSSION Host-Parasite Relationships Acipenseridae: specimens examined-7; infested-0; localities-7. No Ergasilus species have been recorded from this family in North America and none were recorded from the specimens examined in this work. Polyodontidae: specimens examined-4; infested-1; localities -4. Specimens of Ergasilus elongatus were taken from the gill rakers of a specimen of Polyodon spathula which had been collected on the Tallapoosa River. This species of Ergasilus was recovered from P. spathula only in this study and is the only species that has been reported in the literature from this host, Table 1. Lepisosteidae: specimens examined-22; infested-i; localities -18. Of the 22 gars examined, only 1 was found positive for Ergasilus. A single longnose gar from the lower Coosa Drainage was infested with a single E. versicolor. Previous works have recorded what represents two species of Ergasilus from gars, Table 1. E. versicolor was one of these. The other, E. caeruleus,was reported from this family on the basis of a single specimen from a longnose gar and a questionable inclusion in a host list in the same work (100). E. versicolor may be considered the only ergasilid that has been firmly established as a gar parasite. Amiidae: specimens examined-13; infested-O0; localities-11. No records of Ergasilus have been reported in the literature and none were recovered from the material examined in this work. The lack of records of Ergasilus species from Amia calva does not reflect limited published examinations of this fish for parasites but rather a true absence of parasitism by Ergasilus. Future findings of Ergasilus on A. calva will probably be the result of infestations by the Ergasilus species that are not very selective in their choice of hosts. Elopidae: specimens examined-3; infested-O0; localities-2. Three specimens of Elops saurus were negative. The only report of Ergasilus on a North American elopid was made by Causey (22). He reported one specimen of Ergasilus under the name E. mugilus from E. saurus. 30 ALABAMA AGRICULTURAL EXPERIMENT STATION Anguillidae: specimens examined-28; infested-10; localities25. More than one-third of the localities examined produced specimens of E. celestis from Anguilla rostrata. Previous records were also of this species with the exceptions of an unidentified species record and one E. caeruleus record, Table 1. From this information it can be assumed that E. celestis will be the species of Ergasilus to be expected as a parasite of Anguilla rostrata. Clupeidae: specimens examined-149; infested-6; localities76. Previously reported Ergasilus species from North American clupeids are represented by two lots of material. Both of these were initially reported on by Wilson (100). One lot, represented by one specimen, was identified as E. versicolor, but Roberts (78) reexamined this material and designated it as E. arthrosis Roberts. The other material reported by Wilson (100) was described as a new species, E. lanceolatus Wilson. Neither E. lanceolatus nor E. arthrosis were recovered from the clupeid material reported on herein. However, a new species, E. clupeidarum Johnson and Rogers, has been recovered and described (51). The new species was collected from all the clupeid species included in this study and also from Alosa aestivalis from the Atlantic coastal drainage. E. clupeidarum was not found on fishes other than clupeids, and the frequency with which it was recovered suggests that it is the species most likely to be encountered on clupeids. A single specimen of E. megaceros was found loose in the opercular cavity of a specimen of Dorosoma cepedianum. Eight host specimens were collected simultaneously but seven were negative. Engraulidae: specimens examined-33; infested-0; localities13. No ergasilids were obtained from the anchovies examined in this study and no reports in the literature record ergasilids from engraulids. Hiodontidae: specimens examined-20; infested-i1; localities9. One specimen of E. lanceolatus was taken from one of several specimens of Hiodon tergisus that had been collected in the upper Pearl River. This is the first report of E. lanceolatus since the DISTRIBUTION OF THE GENUS ERGASILUS 31 original description by Wilson in 1916. Wilson's material for the original description was taken from the gizzard shad (Dorosoma cepedianum). Esocidae: specimens examined-100; infested-5; localities-76. Ergasilus megaceros was found from four localities as parasites on specimens of Esox americanus and E. niger. One specimen of Ergasilus tenax was found on a specimen of E. niger. Voth and Larson (97) reported an ergasilid from Esox spp. from the Goose River, North Dakota, as Ergasilus sp., but Dr. O. R. Larson in a personal communication related that the material from the study was not retained. Cyprinidae: specimens examined-2,116; infested-73; localities -815. E. cyprinaceus and E. megaceros were the only species that were found on cyprinids in this study. Pages 25-26 show that the two parasites are generally distributed in the various cyprinid genera. Previous reports are somewhat diverse in regard to recorded species of Ergasilus infesting cyprinids. From the literature summary in Table 1 it was noted that much of the material reported by R. V. Bangham was recognized as E. nerkae by Roberts (76). It is possible that other specimens listed by Bangham and Adams (12) were also E. nerkae. Mueller (68) expressed doubt to his own identification of E. versicolor. Voth and Larson (97) suggested that Hoffman (45) may have misidentified the E. caeruleus therein. Furthermore, there is a doubt of the Voth and Larson (97) identification of E. confusus Bere that was reported from Pimephales promelas. The identification is considered doubtful because other specimens reported as E. confusus in the same study were actually representative of a new species, E. wareaglei Johnson, 1972. By combining host record data that have a high degree of certainty as to correct identification with those accumulated in this study, the Ergasilus species that infest cyprinids may be narrowed to three. These are E. cyprinaceus, E. megaceros, and E. nerkae. E. nerkae appears to be limited to more northern temperate regions in North America because it was not taken in this study. Catostomidae: specimens examined-498; infested-37; localities 301. In this study, E. megaceros, E. versicolor, and E. wareaglei 32 ALABAMA AGRICULTURAL EXPERIMENT STATION were the only species collected from catostomids. These three parasites were found on numerous samples of fishes. In the literature the records of catostomid infestation by Ergasilus species are dominated by reports of E. caeruleus, Table 1. However, most of the reports were by Bangham (6-9,11,12) and are all to some degree questionable in regard to correct identification. Roberts (76,80) reexamined some of Bangham's material and found it to be of different species than that reported as E. caeruleus. E. caeruleus was collected 49 times in the present study and was never found on catostomids. If it be true that E. caeruleus records are erroneous, then it appears that E. megaceros, E. nerkae, E. versicolor, and E. wareaglei are the primary ergasilid parasites of North American Catostomidae. Ictaluridae: specimens examined-465; infested-37; localities -265. E. arthrosis, E. versicolor, E. cerastes, E. cyprinaceus, and E. megaceros were encountered, in order of decreasing frequency, from ictalurid hosts. The literature summary shows E. versicolor predominating in past records, Table 1. However, the knowledge of this species has undergone a period of taxonomic confusion and, as explained above, a portion of these literature records probably represent E. arthrosis. The data from this study reported only one collection of E. megaceros from an ictalurid, but the copepods were plentiful on this sample. Add the fact that the specimens from which Wilson originally described E. megaceros came from a channel catfish and it must be assumed that the parasite can be expected on other ictalurids. Prior to this work, E. cyprinaceus was recorded only once from ictalurids, in this case the yellow bullhead (Ictalurus natalis). In the study area bullheads and one madtom were found burdened by this parasite. This information substantiates the association between E. cyprinaceus and ictalurids and suggests a subfamilial preference for the smaller ictalurids by the parasite. In estuarine areas the ictalurids were found infested with E. cerastes. The three Ictalurus spp. that provided E. cerastes specimens were all captured in river mouths. Ariidae: specimens examined-22; infested-6; localities-l10. Pearse (72) reported a copepod from the gills of the gafftopsail catfish (Bagre marinus) and the sea catfish (Arius felis) from DISTRIBUTION OF THE GENUS ERGASILUS 33 Beaufort, North Carolina. Pearse gave the new name Macrobrachinus felichthys to the animal. This copepod is actually an Ergasilus species and Johnson and Rogers (51) have placed it in the genus Ergasilus as E. felichthys (Pearse, 1945). The species is exceptionally close to E. cerastes Roberts, 1969, but has not been combined with the latter since there are differences in morphological detail between the two forms. E. felichthys was found on several samples of Bagre marinus gills in the course of the study. The parasite appeared to be quite common on this host. Although Pearse had obtained one specimen of E. felichthys from A. felis none were obtained from that host in this study. Amblyopsidae: specimens examined-7; infestedO0; localities4. Typhlichthys subterraneus specimens were examined from material from four localities. All specimens proved negative. The collections of the fishes were considered part of the Tennessee River system, but as Smith-Vaniz (84) pointed out, the underground habitat of Typhlichthys allows it to not be dependent on surface drainage systems for distribution. The host specimens recorded in this report were captured near the Coosa system. Aphredoderidae: specimens examined-81; infested-5; localities -47. The pirate perch (Aphredoderus sayanus) has only once been noted as a host for Ergasilus. The species was E. caeruleus and was recorded by Bangham and Venard (14) from Reelfoot Lake, Tennessee. Herein, E. caeruleus and E. arthrosis are reported from this host from four and one localities, respectively. The fact that the E. arthrosisrecord was based on one individual parasite as opposed to numerous parasites per host for E. caeruleus leads one to conclude that E. caeruleus is the Ergasilus species usually expected to be encountered on A. sayanus. Belonidae: specimens examined-22; infested-O; localities 13. Specimens of Strongylura marina represented this family in the study. No Ergasilus specimens were found on the fishes examined as part of the study, but Cressey and Collette (27) did report three Ergasilus species from North America. The first was reported as E. spathula n. sp., but it is actually a synonym of E. arthrosis Roberts. The second species was considered unde- 34 ALABAMA AGRICULTURAL EXPERIMENT STATION scribed, reported as Ergasilus sp. A, and was suggested to be the same as E. lizae Kroyer sensu Thomsen (88). This latter name has been synonymized with E. versicolor Wilson by Johnson and Rogers (51). The third species was considered to be the same as E. tenax Roberts but the description of the animal by Cressey and Collette made it obvious to the authors that it was actually a new species. Summarizing, E. arthrosis and an unnamed species close to E. tenax may be expected to occur on North American belonids. Another species, possibly E. versicolor, has been reported from North American belonids but the collection was based on only one specimen. Cyprinodontidae: specimens examined-547; infested-30; localities-217. The results of this study as well as that of the literature show this group to be dominated by E. funduli and E. lizae as prospective parasites, Table 1. Both of these species are found on cyprinodontids in brackish water areas, especially tide pools. Inlandranging cyprinodontids were found free of ergasilids with the exception of Fundulus olivaceus, specimens of which harbored single E. cyprinaceus specimens at two localities. Poeciliidae: specimens examined-202; infested-7; localities80. Ergasilids have been recorded from poecilids three times in the North American literature, twice as E. sp. from Gambusia affinis and Poecilia latipinna and once as E. manicatus from G. holbrooki. All of these collections were made in the vicinity of Englewood, Florida by Bere (18) and Bangham (8). Ergasilus funduli has been recorded from five localities herein, once from G. affinis and five times from P. latipinna. Since E. manicatus is a synonym of E. funduli, the latter is the only species of Ergasilus that has been found on poecilids. Atherinidae: specimens examined-145; infested-9; localities57. E. funduli has been reported from Menidia menidia from Massachusetts (99,102,80) and New Brunswick (16). Within the study area, this species was obtained from three localities on Menidia beryllina. This information should justify the supposition that E. funduli is the common Ergasilus species of coastal atherinids. The inland atherinid, Labidesthes sicculus, was found to host DISTRIBUTION OF THE GENUS ERGASILUS 35 a single specimen of E. arthrosis from one locality, Table 1. The importance of this recovery is probably minor. Cottidae: specimens examined-31; infested-0; localities 12. The only species of Ergasilus that has been found to infest cottids is E. cotti Kellicott, 1892. The parasite has been recovered only once since the original description and this was from a percid, Etheostoma caeruleum (91). No cottids or percids were found infested with E. cotti in the present study. Unless the parasite is particularly rare in the Southeast, it is likely that it is absent from this region. Syngnathidae: specimens examined-8; infested-0; localities5. The specimens of pipefishes that were examined in the study area proved free of ergasilids. This negative result is likewise found in the literature. Percichthyidae: specimens examined-14; infested 4; localities10. E. arthrosis was found on three samples within the study area and E. centrarchidarumon one from the Atlantic coast. E. labracis has been noted as common on the striped bass. Reports of E. labracis dominate the literature records of ergasilids found on percichthyids, but all of these records were from Atlantic coastal localities, Table 1. It seems quite possible that the mainland distribution of E. labracis is localized on the Atlantic coast with other species parasitizing sea basses in Gulf drainages. E. arthrosis, E. centrarchidarum, E. caeruleus, and E. versicolor have been taken from gills of fishes from Gulf of Mexico drainage basins. Due to the confusion in the literature concerning E. versicolor, it is strongly suspected that the reports of E. versicolor (24) were based on material that was actually E. arthrosis. Specimens of E. centrarchidarumand E. caeruleus collected from sea basses number three each including material from this study. In contrast, E. arthrosis was found in relative abundance on the gills. Elassomidae: specimens examined-102; infested-0; localities -48. No ergasilids were reported from this family herein, and none have been recorded in previous works. Centrarchidae: specimens examined-1,122; infested-167; localities-632. 36 ALABAMA AGRICULTURAL EXPERIMENT STATION The results of this study demonstrate four species of Ergasilus from centrarchids. These are E. arthrosis, E. caeruleus, E. centrarchidarum,and E. tenax. The literature with slight exception contains records of the same species but is dominated by reports of E. caeruleus and E. centrarchidarum,Table 1. Some of the reports of E. versicolor from centrarchids were corrected to E. arthrosis by Roberts (78). The other reports of E. versicolor from centrarchids, notably those of Causey (24), may also have been E. arthrosis for reasons previously mentioned. E. tenax has been reported only from Pomoxis spp. but was found more frequently on Lepomis spp. than Pomoxis spp. in this study. Specimens of Ambloplites rupestris provided no Ergasilus spp. in this study even though specimens from 35 localities were examined. Previous works have reported numerous cases of E. centrarchidarumand E. caeruleus from A. rupestris. Centrarchus macropterus specimens were found to host only E. centrarchidarum in this study. Previously, one specimen of C. macropterus was reported as being parasitized by E. caeruleus (14). Species of the genus Lepomis were found to host E. arthrosis, E. caeruleus, E. centrarchidarum,and E. tenax from 20, 43, 5, and 13 localities, respectively. Infestations by E. arthrosis were prevalent in the lower parts of the drainages. E. caeruleus was generally distributed and E. centrarchidarumand E. tenax were prevalent in the upper reaches of the drainages. E. tenax were present in the upper portions as well. Species of the genus Micropterus hosted E. centrarchidarum from 15 localities and E. arthrosisfrom 10. The apparent absence of E. caeruleus from Micropterus spp. in the study area overwhelms the senior author's confidence in the literature concerning the Centrarchidae. An objective attempt to discredit the portion of literature that reports E. caeruleus from centrarchicds was met with some success: Smith's (83) spine-seta count on the species he reported as E. caeruleus fits E. arthrosis much better than E. caeruleus. Becker, et al. (15) reported that copepod identifications were made under a dissecting microscope, a questionable procedure in the senior author's estimation. The material reported as E. caeruleus in Bangham's (4-11) works have been considered unreliable earlier. Ergasilid reports from Pomoxis in the literature include reliable records of E. caeruleus, E. centrarchidarum,and E. tenax, Table DISTRIBUTION OF THE GENUS ERGASILUS 37 1. The same copepods were collected in this study from nine, three, and four localities, respectively. In addition, specimens of E. arthrosis were collected from two localities. Larger numbers of E. caeruleus and E. tenax rather than E. arthrosis and E. centrarchidarumwere obtained per fish. This result was compatible with the literature. Percidae: specimens examined 921; infested-5; localities-419. Although a large number of localities were examined for Ergasilus species on percids, only four localities provided parasites. Three localities provided specimens of E. caeruleus from Percina nigrofasciata and one locality supplied a single specimen of E. arthrosis from P. caprodes. Most of the specimens of Ergasilus that have been taken from percids and reported on in previous work came from the larger perches - Stizostedion canadense, S. vitreum, and Perca flavescens. Although specimens of S. vitreum and P. flavescens were examined in the present work, the number of localities checked was small. E. luciopercarum has been established by Roberts (80) as the prevalent species on the larger perches. Roberts (80) noted that a synonym of this species (E. confusus Bere, 1931) was reported from other hosts besides percids by Bere (17) and considered the records as probable misidentifications. That E. luciopercarumwas not found on fishes examined in this study suggests that the parasite may not occur here. Also, that no specimens of this parasite have been taken below 41 ° N. latitude adds substance to the assumption. E. cotti has been reported twice in the literature and one of these was from Etheostoma caeruleum (92). The latter record is unique for the genus Etheostoma. It is apparent from the data presented herein that ergasilids are not common on the small percids. Those ergasilids that do occur occasionally as parasites are likely to be the ones that show a similar affinity to the larger percids. Carangidae: specimens examined-9; infested-l; localities-5. One specimen of Caranx hippos was found to host a single specimen of E. arthrosis. This is the first record of an Ergasilus from a carangid from North American waters. Sparidae: specimens examined-22; infested-l; localities-14. Specimens of the sheepshead (Archosargus probatocephalus) 38 ALABAMA AGRICULTURAL EXPERIMENT STATION and the pinfish (Lagodon rhomboides) were examined. One specimen of E. lizae was taken from a pinfish at the mouth of the Pascagoula River. This represents the first report of an Ergasilus from a North American sparid. Sciaenidae: specimens examined-63; infested-8; localities 37. Two species of Ergasilus were taken from two sciaenid species in the present study. One was E. lizae from the spot (Leiostomus xanthurus) from several localities. Causey (23) reported E. lizae and E. mugilus from the spot and says of E. mugilus: "This appears to be the most common ergasilid on the Gulf Coast." It is possible that the species of ergasilid that Causey referred to as E. mugilus (see section on "Taxonomic Considerations") is the same as that reported as E. lizae herein. Causey (21) also reported E. nanus and E. mugilus from the croaker, Micropogon undulatus. In the present work, one specimen of E. versicolor was taken from a black drum (Pogonias cromis) but the relative importance of this find is questionable. The only other sciaenid infestation that the authors are aware of is a freshwater drum (Aplodinotus grunniens) that was collected from the Tennessee River drainage by the junior author and was seen to be heavily infested with an undetermined Ergasilus. Mugilidae: specimens examined-53; infested-27; localities23. Three nominal species of Ergasilus have been reported from this family in the North American literature, E. lizae, E. mugilus, and E. nanus, Table 1. As discussed earlier these three species are poorly defined. One of the species of Ergasilus that was found on Mugil spp. in this work has been called E. lizae following Roberts' (80) designation. Two of the other forms that were obtained from mugilids have been partially figured along with E. lizae, Figure 1, but given no names. No species were given the identification of E. nanus or E. mugilus. Because of the ambiguity involved in previous reports of this species group, the literature records are not considered at this time. The species of Ergasilus that were recovered within the study area were E. versicolor, E. lizae, E. sp. Form A, and E. sp. Form B. The latter two species were collected only from Mugil spp. and are now considered to be restricted as parasites to the family. Within the marine or brackish water environment, E. versicolor DISTRIBUTION OF THE GENUS ERGASILUS 39 and E. lizae were found on other hosts, but primarily on Mugil spp. Collections of E. funduli were examined from Mugil cephalus from Atlantic coastal waters but the infestations were considered light. The association probably was the result of close association of the young mugilids with a dense cyprinodontid population. Gobiidae and Eleotridae: specimens examined-37; infested-l; localities-18: specimens examined-7; infestedO0; localities6. The literature reports only one Ergasilus species from a gobiid in North America. The parasite-host relationship between E. auritus and Gillichthys mirablis was reported by Roberts (80) from specimens collected in southern California waters. Both animals may be considered species that range only in Pacific coastal drainages. In this study E. funduli was the only species obtained from gobiids. Numerous specimens of E. funduli were taken from Gobiosoma bosci from one locality. Negative findings of other Ergasilus species and numerous copepods per fish suggests E. funduli is the important ergasilid parasite of gobiids on the Gulf Coast. Eleotrids all proved negative for ergasilid parasites. Bothidae and Soleidae: specimens examined-2; infested-0; localities-2: specimens examined 14; infested-5; localities-11. No Ergasilus species have been reported from these two families in the North American literature. In this study, specimens of E. arthrosis were recovered from specimens of Achirus lineatus from two of seven localities. This establishes A. lineatus as an acceptable host for E. arthrosis. No Ergasilus specimens were obtained from the other flatfishes examined. Parasite-Host Relationships and Distribution Ergasilus arthrosis Roberts, 1969. The literature reports E. arthrosis from ictalurids, a clupeid, and centrarchids. The results of this study record this parasite from ictalurids, an aphredoderid, an atherinid, percichthyids, centrarchids, percids, a carangid, and a soleid. Roberts (80) considers E. arthrosis the common species on ictalurids with occurrences on other families occasional events. The copepod was found commonly on ictalurids in this 40 ALABAMA AGRICULTURAL EXPERIMENT STATION study and this supports Roberts' assumption that the parasite is common on catfishes. However, centrarchids and percichthyids appear to be just as susceptible to infestation by E. arthrosis. Adequate numbers of Achirus lineatus were not examined but the frequency with which the parasite was encountered on this host (2 of 7 localities) and the intensity of infestation suggest that E. arthrosismay be found commonly on this host too. The miscellaneous collections from the other host families serves to demonstrate the wide range of suitable hosts for this parasite species. Although E. arthrosis appears widely distributed on Figure 3, most collections were taken from the lower portions of the drainages. Those collections from the upper drainage areas were primarily from ictalurids and from rivers and reservoirs. Outside of the study area E. arthrosis has been documented in several FIG. 3. Distribution of E. arthrosis (squares) and E. clupeidarum (dots). DISTRIBUTION OF THE GENUS ERGASILUS 41 localities in the Mississippi River drainage basin and in two collections from coastal areas of Louisiana and Costa Rica. Ergasilus caeruleus Wilson, 1911. This species name is particularly abundant in survey records. One (103) reports the parasite from a lamprey. The species was reported most frequently by Bangham, but since Roberts (76,80) has found so many discrepancies in Bangham's works, it is suspected that the majority of them are erroneous. The same must be said for other reports, with the exceptions of Wilson (99,100), Tedla and Fernando (85), and Roberts (76), since there had been taxonomic confusion regarding E. caeruleus prior to Roberts (76). E. caeruleus was found on Aphredoderus sayanus, Lepomis spp., Pomoxis spp., and Percina nigrofasciata in this study. The frequency of occurrence and intensity of infestation show that these species are suitable and common hosts for E. caeruleus. Furthermore, E. caeruleus was recovered from each of these host groups from more localities than any other Ergasilus sp. The distribution presented itself as a random pattern within the study area, Figure 4. No specimens, however, were taken in brackish or salt waters. Comments on continental distributions could be biased because of numerous ambiguous literature records, but there are enough reliable reports available to allow for generalization. It can be said with certainty that E. caeruleus may be expected to be a common ergasilid within the Mississippi River drainage, east of it in the United States, and in eastern Canada. Further, well documented reports will probably extend the range westward, complementing that of the natural range of the Centrarchidae. Ergasilus celestis Mueller, 1936. Lota lota and Anguilla rostrata have been reported as hosts for this species, from three and two localities respectively. The results presented herein firmly establish A. rostrata as a preferred host for this species. It is possible that E. celestis is restricted to these two hosts. As can be seen from Figure 5, E. celestis is more or less randomly distributed on A. rostrata in the study area. Outside the study area, E. celestis has been collected from single localities in New York, Massachusetts, and Ontario. Ergasilus centrarchidarum Wright, 1882. Previous reports record E. centrarchidarumfrom catostomids, centrarchids, percichthyids, and percids. Centrarchids and a percichthyid are re- 42 ALABAMA AGRICULTURAL EXPERIMENT STATION 42 ALBAMA ARICUILTUA XEIMN TTO FIG. 4. (ring). Distribution of E. coeruleus (squares), E. lonceolafus (dot), and E. elongotus ported as hosts herein. It was suspected that the catostomid parasites were actually E. wareaglei since the two have very similar morphology and the percid records were erroneous. Tedla and Fernando (85) checked fishes of four species from one Ontario locality and found three of them, Lepomis microlophus, Ambloplites rupestris, and Micropterus salmoides, infested with F. centrarchidarum. The heavier infestations were on the latter two species. Since the parasites were larger on A. rupestris than M. salmoides, A. rupestris was considered the preferred host for F. Roberts (80) studied Wilson's (99,100) material, part of Bangham's collections, and material sent to him as gifts, and proposed that Micropterus and Am-bloplites are the preferred hosts of E. centrarchidarum. From the results presented here it will be noted that no A. rupestris were found posi- centrarchidarum. DISTRIBUTION OF THE GENUS ERGASILUS 43 43 DISTRIBUTION OF THE GENUS ERGASILUS FIG. 5. Distribution of E. celestis (squares) and E. cerastes (dots). tive for this parasite and Centrarchus macropterus, Micropterus spp., and Lepomis spp. were suitable hosts for E. centrarchidarum. The one specimen taken from Morone chrysops serves to substantiate this fish as an occasional host, for Wilson (100) had reported a similar symbiosis. The distribution of E. centrarchidarumin the study area reveals that the species is distributed primarily in the upper portions of the drainages. No collections were taken in areas of tidal influence. In North America E. centrarchidarum has been recorded from throughout the Eastern United States and Eastern Canada. Ergasilus cerastes Roberts, 1969. Roberts (79) described this species from an Ictalurus species from a Washington, D.C. fishmarket. Roberts correctly postulated that the species of Ergasilus 44 ALABAMA AGRICULTURAL EXPERIMENT STATION that Mueller (68) had reported as E. elegans (= E. versicolor) was also E. cerastes. E. cerastes is reported from four localities in this paper, all on Ictalurus spp. Apparently this species is restricted to Ictalurus in its host selection. The distribution of the parasite is peculiar in that it has only been taken near or in estuaries, Figure 5. Ergasilus clupeidarum Johnson and Rogers, 1972. This species has been taken only from clupeids and it would be expected that future collections of this parasite be collected from these hosts. The distribution may be expected to be generalized since specimens were taken from far inland localities as well as Mobile Bay, Figure 3. FIG. 6. Distribution of E. centrarchidarum (squares) and E. felichthys (dots). DISTRIBUTION OF THE GENUS ERGASILUS DISTRIBUTION OF THE GENUS ERGASILUS 45 45 FIG. 7. Distribution of E. cyprinaceus (squares) and E. funduli (dots). Ergasilus cyprinaceus Rogers, 1969. Rogers (81) reported this species from four genera of cyprinids and Roberts (80) added another cyprinid genus to the list. Also, Roberts (80) reported one specimen from a stickleback (Culaea inconstans) and three specimens from Ictalurus natalis that were collected by Bangham (9) and reported as E. versicolor. Herein, the parasite was collected from seven cyprinid genera, ictalurids, and one species of Cyprinodontidae. The cyprinodontid Fundulus olivaceus has a peculiadistribution for a member of a secondary division (70) fish group, being found in strictly fresh-water habitats. This ecological peculiarity may account in part for its selection as a host. The ictalurid records add importance to the collection by Bangham (9) from I. natalis. The occurrence of the parasites on the bullheads was considered frequent enough to establish these fish as pre- 46 ALABAMA AGRICULTURAL EXPERIMENT STATION ferred hosts. The one specimen from the Noturus nocturnus specimen was not considered as important because more madtoms were examined than bullheads. The distribution of E. cyprinaceus, Figure 7, may be considered as scattered within the study area with no collections recorded near or within the estuaries. Continental records from outside the study area have been reported from single localities in Mississippi (this work), North Dakota (80), and Florida (80). Ergasilus elongatus Wilson, 1916. This species has been found only as a parasite of the paddlefish, Polyodon spathula, Table 1. As mentioned in the section on host-parasite relationships, this parasite is probably restricted as a parasite to this host species. E. elongatus has now been recorded from the Mississippi River and Tallapoosa River-Mobile Bay drainage systems. Ergasilus felichthys (Pearse, 1947). As mentioned under Ariidae above, this species appears to be restricted in host preference to ariid catfishes and especially to Bagre marinus in North America. The distribution of this parasite reflects that of the host family by being confined to brackish or coastal areas, Figure 6. Ergasilus funduli Kroyer, 1863. Individual data on locality of host capture was not as complete as one would prefer. Nevertheless, it is adequate enough to indicate that E. funduli has tide pools and brackish lagoons as its habitat. The host species on which E. funduli were found were cyprinodontids, poecilids, an atherinid, a mugilid, and a gobid, and all were collected from or in the proximity of the same habitat type, Figure 7. It is likely that cyprinodontids and poecilids are the preferred hosts and the other species become infested when sharing a closed habitat with these fishes. The literature reports this species from cyprinodontids, an atherinid, gasterosteids, a poecilid, and an osmerid. These collections were made on the coasts of Louisiana, Florida, North Carolina, Massachusetts, and New Brunswick (55,18,72,99,102,80). Ergasilus lanceolatus Wilson, 1916. This species was found on the mooneye (Hiodon tergisus) from the Pearl River, Figure 4, in this study. Previously, E. lanceolatus was known only from the original description (100) which reported only Dorosoma cepedianum as the host. Little can be said about distribution and host affinities with such limited data as the basis of discussion. DISTRIBUTION OF THE GENUS ERGASILUS 47 Ergasilus lizae Kroyer, 1863. Samples of this species were collected from numerous localities along the cost from cyprinodontids, mugilids, a sparid, and a sciaenid, Figure 9. Previous works report this parasite several times from Mugil spp. and once from a sciaenid. The sciaenid, Leiostomus xanthurus, was also the species that is reported as a host herein. Roberts (80) suggests correctly that E. lizae is probably a cosmopolitan parasite of Mugil spp. Future taxonomic work will probably reveal that E. lizae has been described under several names. Two names, E. nanus van Beneden, 1870 and E. mugilus Vogt, 1877, are two fine prospects. Reports of ergasilids under these taxa have been reported by Wilson (99), Causey (21,22) and Pearse (75). Wilson deposited material of E. mugilus, but those catalogued at the U.S. National Museum contain free living copepods (Dr. R. Cressey, USNM, personal communication). The materials collected by Causey and Pearse were not available for examination, but it is suspected that the identifications were erroneous. The cyprinodontids from which E. lizae have been reported are probably the victims of confined habitat and close association with young Mugil spp. in coastal situations. The converse was noted in the earlier discussion of E. funduli. Kelley and Allison (53) reported E. lizae from centrarchids from Lake Shelby, Baldwin County, Alabama. Roberts (78) reported that this material was actually E. arthrosis. He based this assertion on the examination of some of the Kelley and Allison material. Recently, the senior author had the opportunity to examine a larger portion of the material and found that E. lizae was indeed present in the material. However, the species was present in very small numbers as compared to E. arthrosis,the true causative agent of the fish mortality that was reported by Kelley and Allison (53). Ergasilus megaceros Wilson, 1916. E. megaceros was found to be rather evenly distributed in the study area, Figure 8. The parasite was found on several species of cyprinids and catostomids, two esocids, one ictalurid, and one clupeid. The importance of the latter collection appears minor since only one specimen was found unattached in the opercular cavity of one of eight fish from the locality. The data from the other collections indicate that the parasite is adapted to the families Cyprinidae, Cato- 48 48 ALABAMA ALABAMA AGRICULTURAL EXPERIMENT STATION AGRICULTURAL EXPERIMENT STATION FIG. 8. (ring). Distribution of E. megaceros (squares), E. sp. Form A (dots), and E. sp. Form B stomidae, Esocidae, and Ictaluridae. The reports of E. megaceros in the literature, Table 1 do not alter this hypothesis. The distribution of E. megaceros has been established in the Mississippi River basin (100), the study area, south Florida (8), the Atlantic coastal drainages (80), and the Great Lakes drainages (67). Future collection will probably extend the range throughout most of North America. Ergasilus tenax Roberts, 1965. This species was found in two drainage basins, Figure 9. The fish hosts included representatives from two centrarchid genera, Lepomis and Pomoxis, and a single collection from an esocid, Esox niger. Importance cannot be attributed to the latter collection since only one parasite specimen was obtained. The literature, Table 1, includes reports only from Pomoxis DISTRIBUTION OF THE GENUS E!RGASILUS 49 DISTRIBUTION OF THE G;ENUS EiRGASILUS 4 annularis for this species. It is assumed, therefore, that this species is selective for Lepomis and Pomoxis species as fish hosts. As mentioned previously, this species was placed in a new genus by Cressey and Collette (27), but the species probably should be retained in Ergasilus. The genus was established on the basis that the first antenna possessed only five segments as opposed to six for Ergasilus. However, at least one Ergasilus, E. kandti van Douwe, 1921, has been seen (British Museum No. 1950.7.29.17) to possess only five first antennal segments. The two species that were assigned to this new genus also had two segmented first endopods. This character is peculiar to a group of Ergasilus species of this hemisphere and, especially, North America. FIG. 9. Distribution of E. fenax (squares) and E. Iizae (dots). 50 ALABAMA AGRICULTURAL EXPERIMENT STATION Ergasilus versicolor Wilson, 1911. In this study, E. versicolor has been taken from catostomids, ictalurids, a lepisosteid, a mugilid, and a sciaenid. The one collection of E. versicolor from the lepisosteid is considered of importance since the relationship between the parasite and lepisosteid has been reported many times in the literature. The importance of the percichthyid record is uncertain. The distribution of E. versicolor, Figure 10, is peculiar for the genus. It has an inland to coastal range in the study area and those that were found in saline waters were, with the exception of the percichthyid, on Mugil cephalus. Nevertheless, all material was considered E. versicolor by Johnson and Rogers (51) and herein for the reasons of morphological conformity. There is a slight difference in the morphology of some of the Mugil infesting copepods, but there is so much overlap between characters that a detailed allomeric study will have to be undertaken to separate the groups. Even then, this will not justify separate species status. Although the literature has reported E. versicolor from occasional centrarchids, cyprinids, and percichthyids, Table 1, the centrarchid and percichthyid material may have been E. arthrosis, and the cyprinid report was erroneous. E. versicolor, then, appears to be selective for species of the families Lepisosteidae, Catostomidae, Ictaluridae, and Mugilidae. The continental distribution of E. versicolor must be derived with scrutiny since the literature of this species is characterized by nomenclatural weaknesses. Reliable records and the data present can establish the distribution in coastal waters on the Atlantic side of North America at least up to Georgia and in inland waters in the Mississippi River basin and eastward. Ergasilus wareaglei Johnson, 1973. Collections of this species were made from four localities, Figure 10. E. wareaglei was found on Hypentelium nigricans from three localities and on H. etowanum from one locality. As mentioned above, this species was reported from Catostomus commersoni from the Mississippi River drainage by Voth and Larson (97) as E. confusus. Apparently this species is specific for fishes of the family Catostomidae. Ergasilus spp. Forms A and B. These species were taken from the gill rakers and arches of Mugil cephalus and Mugil curema from several coastal localities, Figure 8. The species were noted to DISTRIBUTION ISTRBUTIN OF OF THE GENUS EIRASLUS THE GENUS ERGASILUS 51 5 FIG. 10. Distribution of E. versicolor (squares) and E. wareaglei (dats). be morphologically very close to E. lizae with the exception of the second antennae. In one species, E. sp. Form A, the length of the third antennial segment was relatively longer than the same of E. lizae, and the terminal (fourth) segment was observed to be relatively shorter, Figure 1. The ratio (fourth segment 0.450.5 X the length of third) is comparable to the specimens that Roberts (78,80) reported for specimens of E. lizae from Chile and Israel. Roberts (78,80) reported other E. lizae from Puerto Rico and Georgia with antennal characteristic comparable to those of the specimens of E. lizae that have been listed as such herein, Figure 1. Both E. lizae and E. sp. Form A have been taken simultaneously from one fish. The usual habitat (gill rakers) and distinct morphology of the antenna provide the basis for separation of this form from E. lizae in this work. 52 ALABAMA AGRICULTURAL EXPERIMENT STATION The other species, E. sp. Form B, similarly may be separated from E. lizae. This species frequents the gill arches rather than the rakers or gill filaments and has segmental features of the antenna that appear stouter than E. lizae or Form A, Figure 1. Form A and Form B have only been found on gill rakers and arches of Mugil spp. in coastal waters. The senior author has seen only Form B outside of Gulf waters, but it is likely that Form A is more widespread as well. Ergasilus auritus Markevich, 1940; Ergasilus chatauquaensis Fellows, 1877; Ergasilus cotti Kellicott, 1892; Ergasilus luciopercarum Henderson, 1926; Ergasilus nerkae Roberts, 1963; Ergasilus turgisus Fraser, 1920. These species were not found in the present study area; and with the exception of E. chatauquaensis, it is probable that they do not occur in the Southeastern United States. E. auritus has been reported (59,76,41) from British Columbia, Newfoundland, and Labrador from a stickleback (Gasterosteus aculeatus) and a salmon (Oncorhynchus nerka). Another collections from southern California was reported by Roberts (80) from a goby (Gillichthys mirabilis). It is probable that this species' distribution in North America is confined to Pacific Ocean drainages. Hanek and Trelfall (39) recorded in figures this species from Newfoundland, but the figures most likely depicted something other than E. auritus. E. chatauquaensis has been reported only as part of plankton tow (29,99,104). E. cotti has been reported from a sculpin and a darter from the vicinity of Westerville, Ohio. (52,92). E. luciopercarum,with the exception of one specimen collected from Ambloplites rupestris, has only been reported from Perca and Stizostedion, Table 1. The species has not been taken below 410 N. latitude and west of 91 ° W. longitude in North America. This species has been recovered in plankton tow in a collection from Newfoundland (28). E. nerkae has been found to infest salmonids, catostomids, and cyprinids in drainages that empty into the Pacific Ocean in Washington and British Columbia (76,80). One report (80) from a "cisco?" from Woodruff, Wisconsin is the only record of E. nerkae in waters of the eastern North American drainage. Could this have been an error or an introduction? DISTRIBUTION OF THE GENUS ERGASILUS 53 DISTRIBUTION OF THE GENUS RGSI LUS 5 E. turgisus has been reported from a salmon, a stickleback, and a surfperch from Pacific drainages (35,25,20,76,80). Ergasilus labracis Kroyer, 1863. This species occurs in the Southeastern United States but is considered to have a localized distribution. F. labracis has only been noted as a parasite of the striped bass (Morone saxatilis) on the Atlantic coast, Table 1. It is apparent that this ergasilid is economically important since the striped bass is a prized sport fish, and the infestations by F. labracis were always noted in high numbers per fish. That this parasite was not recorded from the study area indicates that the species perhaps has not established here, and it would be wise for fishery biologists to take every precaution that this species is not inadvertently introduced into the Gulf of Mexico. TABLE 1. SUMMARY OF LITERATURE REPORTING Ergasilus spp. Host and Parasite Polyodontidae Polydon spathula E. elongatus E. elongatus E. elongatus Lepisosteidae Lepisosteus osseus E. versicol'r----E. caerulewr---L. platostomus E. versicolor'---L. spatula E. caeruleus.vrior---Anguillidae Anguilla rostrata E. caeruleus---E. celestis-----E. sp.------- -Cyprinidae Acrocheilus alutaceus E. caruleus Campostoma anomalur E. cyprinaceus------Gila atraria E. nerkae'----------- Locality Reference ------- Ill. ---L -- a. -- Iow a-- -Wilson, 1916 _Causey, 1957 Wilson, 1916 Ioa __Io a--------------- . Iowa ---------- ------------ ilson, W W ilson, Wilson, 1916 1916 1916 Iowa-------------- Iowa ------------ Wilson, _La.-------------- 1916 Cansey, 1957 _Tenn-------------Mass.------------ Bangham and Venard, 1942 N.Y.-------------Newfoundland---- Roberts, Mueller, 1969a 1936a Labr.---------- _Hanek Hanek and Threif all, 1970 and Threlfall, 1970 SB.C. _Ala. _Baugham and Adams, 1954 -. Rogers, 1969 _Bangliam, 1951 (Cont.) 54 ALABAMA AGRICULTURAL EXPERIMENT STATION TABLE 1. (Cont.) __Bangham and Adams, 1954 _. Bangham and Adams, 1954 Hybopsis plumbea E. nerkae2 Mylocheilus caurinus E. nerkae2 Nocomis leptocephalus E. cyprinaceus Notemigonus crysoleucas E. megaceros E. sp.- _B.C.--B.C.-.Ala.___ _Fla.--- Rogers, 1969 -Me. Wis.__ .Ala.___ _N.Y.__ _.Ala.___ _N.D_. N.D._. - .Bangham, 1941a -. Meyer, 1954 _.Fischthal, 1952 Rogers, 1969 -Mueller, 1940 _Rogers, 1969 __Voth and Larson, 1968 -- offman, 1953 H __Voth and Larson, 1968 -- oberts, 1970 R __Bangham and Adams, 1954 Notropis baileyi E. cyprinaceus Notropis crysocephalus E. versicolor Notropis venustus E. cyprinaceus Pimephales promelas E. confusus E. ceeruleu E. sp.- Pimephales sp. E. cyprinaceus Ptychocheilus oregonensis E. nerkaeY Rhinichthys osculus E. caeruleus Richardsonius balteatus E. nerkae2------------Semotilus atromaculatus E. caeruleus E. .- :lB.C. -Wy. I~Ohio Ohio __Bangham, 1951 Bangham and Adams, 1954 sMich. lorOnt. ---------E. sp --------------cyprinaceus--------.corporalis -Hoffman, 1953 -Rogers, 1969 _Voth and Larson, 1968 Semotilus E. ;Wis _Mueller, 1936a, 1936b -Mueller, 1940 - -- F sp .-- - - - - - - - - - - - . megaceros.--------- ILabr. _B.C.-- Catostomidae Catostomus catostomus 2 E.' centrarchidarum'_--_ E. caeruleus----------E. E. caeruleus--------- __Bangham, 1941a, 1955 __Threlfall and Hanek, 1970a Bangham and Adams, 1954 Bangham, 1941b E. nerkae -----------E. versicolor ----------- Out. Catostomus commersoni E. caeruleus----------- Lab.. E. caeruleus__----_caeruleus----------_ .Wilson, 1924 Bangham, 1941a, 1955 -Hoffman, 1953 Bangham, 1946 _Threlfall and Hamek, 1970a (Cont.) DISTRIBUTION OF THE GENUS ERGASILUS 55 DISTRIBUTION E. E. E. E. E. E. E. E. OF THE GENUS ERGASILUS TABLE 1. 55 (Cont.) _Voth and Larson, 1968 _Roberts, 1970 .Hunter, 1942 _Voth and Larson, 1968 .Baugham, 1946 _Fischthal, ]1950 Sinderman, 1953 _Baugham, 1951 Baugham and Adams, 1954 .Bangham, 1941c _Baugham, 1941b wareaglei 4 megacerossp ."--- - --sp ."--- - - - sp ."--- --- sp ."--- -- -- ND.__ .Mass._ Conn._ versicolor-- _Mass._ Catostomus fecundus nerkaei--------- Catostomus macrocheilus E . nerkaeY-------------Erimyzon sucetta E. E. caeruleus ------------__ caeruleus.-----------centrarchidarum---- E. E. megaceros ----------- Baugham, 1941b _Baugham, 1941c _Wilson, 1924 .Baugham, 1946 Ictiobus cyprinellus E. versicolo------------Moxostoma valenciennesi E. caeruleus------------Moxostoma macrolepidotum E. nerkaee -------------Ictaluridae Ictalurus furcats E. versicolor___ Ictalurus natalis E. arthrosis__-E. elegans____ E. versioon__ E. versicolar___ E. versicolor___ E. versicolor___ E. cyprinaceus .Roberts, 1970 _La. _ Unk ----------- . -Tenn _ ------------ - - - -Ill. - - ------ - - - -I d.- ------ ---- --- _Causey, 1957 -Roberts, .Wilson, 1969a Bangham and Venard, 1942 .Wilson, 1911 O hio ------------Fl a..-- -- - -- - - - -W K an . -is. .Harms, 1911 1959, 1960 Bangham, 1941b .Bangham, 1941c Ictalurus melas E. versicolcn E. versicolor__ E. arthrosis--_ 5 E. versicolor__ E. versicolor__ E. versicolor__ __ ----------Iow a ---- -N .----- .D -T -----------enn. - -- _Roberts, 1969a -Wilson, 1916 _Voth and Larson, 1968 -Roberts, 1969a _Bangham and Venard, 1942 -Harms, 1959, 1960 _Bangham, 1941b _Voth and Larson, 1968 ----------U nk .----- --. Kan .----Ohio-- -- -- - -- - --. - -E . sp.-------Ictalurus nebulos ___ -E. arthrosis - -- --- -- o .? --------.B a -- .C -E. arthrosis 7 Iow ----------- a ._-_ E. arthrosis E. versicolor-_ -- --- - -- -- -- --- --- is.--- --- W E. versicolor__ -E. versicolor__ -- --- -- -- -- -O .E. versicolor__ -- - ----------- - -- -- -- -- F la. E. versicolor__ ----- M ass.E. versicolor__ -- ----- N .D .-- Roberts, 1969a -Wilson, 1916 Roberts, 1969a 1911 1946 .I d. O h io -Wilson, n t. --- -- Bangham, Bangham, 1955 _Bangham, 1941b Mueller, 1936b Sinderman, 1953 (cont.) 56 ALABAMA 56 Ictalurus punctatus E. arthrosis--------E. arthrosis -------E. caeruleus-------E. ele gans--------_ E. megaceros------E. arthrosis7 -------7 E. arthrosis ALABAMA AGRICULTURAL EXPERIMENT STATION TABLE 1. (Gont.) AGRICULTURAL EXPERIMENT STATION L. LErie--. --- Ohio-----Tenn.----- - - Fla.- - - - - - ----------- ______Roberts, 1969a -.---Roberts, 1969a and Venard, 1942 _ - ____Bangham E. versicolo------E. versicolos-------E. versicolor-------E. versicolor-------E. versicolor-------E . sp . -- -- -- - --- -Ictalurus sp. E. cerastes--------E. elegans---------Noturus gyrinus E. caeruleus - ----E. versicolor-------Noturus E. versicolor-------_ ----- Iowa----Iow ------ a---------- ---______Wilson, _Bangham, 1941c __ ------hio----O ------ O hio---------- Tenn. ---T ex.- Iowa----- K ------an. ----L. ----- Erie-------. ----- 1916 ----Wilson, 1911 ______Wilson, 1916 _____Tidd, 1931 _____Bangham, 1941h ._____Bangham and Venard, 1942 ._____Harms, 1959, 1960 _____Bangham and Hunter, 1939 Lawrence and Murphy, 1967 ----- --- -----Unknownla .----- ----Roberts, ----- -----Mueller, Bangham, 1969b 1936b 1941b ---O -- hio----L. ------ ----- . Erie- -L .____Bangham and Hunter, 1939 augham and Hunter, 1939 .____B .____.Causey, 1957 flavus Erie--- Pylodictus olivaris E. versicolor -_-- -- a.---L ---- Cyprinodontidae Cyprinodom variegatus E. funduli'--------Floridichthys carpio E . lizae--------- --E . sp . -- - - - - -- - - -Fundulus chrysotus E. caeruleus-------Fundulus con fluentus E. f unduli---------Fundulus grandis E. lizae-- ---- ----Fundulus heteroclitus E. funduli ---------E. manicatus------_ ------.C N .----- .____Pearse, 1947 - F la .--- - - - - --- la .-- - - - - --F --- la .-- - - - - -.F -- --L 1936 ._____Bere, ._____Baugham, 1941c ._____Baugham, ._____Kr~yer, 1941c 1863 - -- a .- - - - -- --la .-F ------ ._____Bere, 1936 ----M ass.-----Wilson, 1932 Roberts, 1970 -----M ass.----- ----- Fundulus majalis E. funduli --------- -----N .C .---- Pearse, 1947 _______ 1936 ._____Bere, 1936 ._____Bere, ._____Baugham, 1941c Pearse, 1947 (Cont.) Fundulus similis E . lizae---------Jordanella --- la .--- -F - - --la .- - - - -F E. manicatus--------F ---E sp .. ------------ -- la .-- floridae Lucania parva E. funduli---------- ----NC. ---- DISTRIBUTION DISTRIBUTION OF THE GENUS ERGASILUS TABLE OF THE GENUS ERGASILUS 57 57 1. (Cont.) Percichthyidae Morone saxatilis E. labracisE. labracis ---E. labracisE. labracis- M. americanus E. confusus--E. labracis E. labracis---- -- --- d.---- ---M --Pa. - - - - - - -- ass . ----M __Wash., _ a. - - - - - - - -V M ass . ----- D.C.- .Kr~yer, 1863 _Leidy, 1887 -Wilson, 1911 .Wilson, 1911 -Wilson, 1932 _.Tedla and Fernando, 1969b -- eyer, 1954 M -Wilson, 1916 -Wilson, 1916 S_Causey, 1957 .Causey, 1957 -Wilson, 1911 ---- --E . sp.-M. chrysops E. cacruleus -E. centrarchidarum E. versicolor M. mississippiensis E. versicolor Centrarchidae Ambloplites rupestris E. caeruleus E. caeruleus -E. cacruleus--- ----M e .--- - - ---- --. .Y .------N Iowa---- --- Iowa La.--------- _La.. .__Mich. i.-cpesWis. -Wis. E. E. E. E. E. E. E. e. E. E. cacruleus caeruleus centrarchidarum centrarchidarumcentrarchidarum centrarchidarum centrarchidarum centrarchiaru centrarchidarum centrarchidarum -Ont. --;.__-------N.Y.----- d.--I ---Mich_. .- Mich.-Que.---- -Ot. L Erie Ot. --. ?-__Wis.--. 1924 __.Pearse, Bgham, 1946 Bu__. __. Fischthal, 1950, 1952, 1947 __.Bangham, 1955 __.Mueller, 1940 -- ilson, W __Tidd, 1911 1931 --.-- W is.---Unk.----- N .Y.--- Wilson, 1924 __ 1924 __.Pearce, -Roberts, 1970 __Tedla and Fernando, 1969a _Bangham and Hunter, 1939 Wright, 1882 -_Bere, 1931 _Bangham, 1946 .Roberts, 1970 __.Hunnien, 1936 _Baugham and Venard, 1942 .Wright, 1882 _Meyer, 1954 Wilson, 1916 ---Wilson, 1916 Bangham, 1941b Bailey, 1966 1963 Le.oicnus E. cele s E. ---lc ier crum----- -- ----------- --- Tenn.__ ___ E. caeruleus O --- nt.? M .-- e.- -,i/Ohio E. centrarchidarum----- -- - Iowa___ E. a t rssp ----------Lepomis cganeolus Iowa--E. caeruleus------------ ----Okla.-E. .McDaniel and _McDaniel, caeruleus-------_____ ---------- E. caeruleus E. caeruleus---- ------ .__Iowa__. W --- is. -Wis.- -Wilson, 1916 ..Bangham, 1946 ..Fischthal, 1950, 1947 Bangham, 1941b (Cont.) 58 ALABAMA 58 E. E. E. E. ALABAMA AGRICULTURAL EXPERIMENT TABLE 1. AGRICULTURAL EXPERIMENT STATION STATION (Cont.) _______Bangham, 1955 _______ and Fernando, 1969a Tedla Tedla and Fernando, 1969 Wright, 1882 ------------- Meyer, 1954 ___ Wilson, 1916 caeruleus-Ont. caeruleus centrarchidarum centrarchidarum - -Ont. Ont. Ont.? M e.----- E. sp. ----- -- Lepomis gulosus E. caeruleus-Iowa E. caeruleus____a. caeruleus.____Tenn. E. E. ------------Causey, ---- E. caeruleus____ Tenn. E. centrarchidarum_ E. centrarchidarum E. liae E. versicolor-La sp. ------- - -d. Iowa ------W .------ ilson, 1911 1957 Bangham, 1941c Venard, 1941 Bangham and Venard, 1942 Ala.___ Tex.----- ------- Wilson, 1916 ---. Kelly and Allison, 1962 ---- awrence and Murphy, 1967 L ---Roberts, 1969a ---- mith, 1949 S 1931 Lepomis macrochirus E. arthrosis_____a. E. caeruleus-N.J. -d._ E. ceruleus E. caeruleus-Ohio E. caeruleusOi__o E. caeruleus-Fla. E. caeruleus-Wis. E. caeruleus-Wis. E. caeruleus-Wis. E. caeruleus-Wis. IowaE. centrarchidarum E. centrarchidarum Ga. E. centrarchidarumTenn. E. centrarchidarumd. E. centrarchidarum E. centrarchidarumOhio E. lizae -------Ala.------------- Okla._ E. versicolor E. versicolorLa. E. sp.-Calif.- ._-____ Wilson, 1911 _______Tidd, _______Bangham, 1941b _______Bangham, 1941c _______Bangham, 1946 ---- nthony, 1963 A ______Fischthal, 1950, 1952, 1947 -------- 1970 Roberts, ------- Wilson, 1916 _______Bangham, 1941b ------- Roberts, 1970 .___Bangham and Venard, 1942 ------- Wilson, 1911 __ Bangham and Hunter, 1937 K ---- elly and Allison, 1962 McDaniel, 1963 _______ Causey, 1957 ._____ -- -- E . sp.--------------- 1953 _______Haderlie, _______ Lawrence and Murphy, 1967 ---- cDaniel, 1963 M .______.Baugham, 1941c -------. Kelly and Allison, 1962 _______.Bangham, 1941c --- T ex.-- Lepomis megalotis E. versicolor------------ Okla.-Lepomis microlophus E. caeruleus.____________Fla.___ E . lizaee ---------------- Ala.--Lepomis punctatus E. caeruleus.___-------Fla.___ Micropterus dolomieui E. E. caeruleus--------------La. caeruleus------------- Ohio-- -------Causey, 1957 E. caeruleus_____________W is.-. E. E. E. E. caeruleus.-----------. caeruleus_____________Ont. __ Ohio-caeruleus---caeruleus------------- Tenn.3 B.C.-Wis._. ----- Banighami, 1933 Bangham, 1946 -------- ____Bangham and Adams, 1954 _____Bangham, 1955 Bangham, 19411), 1947, 1952 E. centrarchzdarum E. centrarchidarum-____--Iowa-- ------ Tenn. _ ----Venard, --------- 1940 Bangham and Venard, 1942 Bangham, 1946 W ---- ilson, 1916 (Cont.) DISTRIBUTION ERGASILUS DISTRIBUTION OF THE GENUS ERGASILUS OF THE GENUS 59 59 TABLE 1. (Cant.) centrarchidarum -Ohio--centrarchidarum a.-- -_ centrarchidarum --Fla.---centrarchidarum ---. -- Fla.---centrarchidarum_---_- __.M ass. -_ centrarchidarum -Ohio--centrarchidarum -_.Conn. __ centrarchidarum L. Erie_ centrarchidarum Iowa centrarchidarum -Iowa.-sp .-- - - - - - - - - - - - - - - --. Calif . _ sp . -- - - - - - - - - - - - - -- -Tex.--'sp .-- - - - - - - - - - - - - - - -N.Y.--sp .- - - - - - - - - - - - --- - W is.---Pomoxis annularis E. caeruleus---------E. ceruleus---E. centrarchidarum ------- a--Iow E. sp. -- - - ------- --Tex. E. caeruleus ----- Wis E. caeruleus A rk.--------E. centrarchidarum E. centrarchidarumE. centrarchidarum. -Ohio ----E. centrarchidarum-----Ohio E. centrarchidarum- ------- M ass. E. centrarchidarum- ------O E. centrarchidarum- ------E. confusus ------------- Wis.---E. sp."-- - - - - - - - -- M e.--- ---- --OhioE. s -- - - - - -- - - - - - ---p E. sp --- - - - -- - - - - - - -. ---- N .Y.- -Micropterus punctulat-us E. caeruleus ---- -Ohio E. caeruleus -----A rk.-------Ohio E. centrarchidarum_ E. versicolor_______ -- a.--L -_--Iowa--- E. E. E. E. E. E. E. E. E. E. E. E. E. E. __G ----1931 hRobe___________ rts, 1970 _Tidd, ------Bangham, 1941c, ------- 1936b Mueller, 1938 _ ___________Sinderman, 1953 Ba___________Bgham, 1933, 1941b lHunt___________ er, 1940 1970___________Rbrs, Wilso___________ on, 1916 _______ ___Haderlie, 1953 .--------- ---Roers __Bangham and Hunter, 1939 ._unin Lawrence and Murphy, 1967 ,13 1936___________ ------- Gilbert, 1905 Marshall and -Ohio- - -------Becker, et Wilson, 1916___________W aghm 1941b___________Bnhm Wilson,1916___________W ----Lawrence _.___.Fischthal, and Murphy, 1967 1947, 1952 al, 1966 -- nt.--- oWils___________ on, 1911 Smith___________ , 1949 _Bangha___________ m, 1933 _ ___________Bangham, 1941b ------------ 1970 Roberts, _________Tedla and Fernando, 1969a ____________1931 Bere, -----Meyer, 1954 L. Erie_ ------Hunnien, 1936 ------- Bangham and Hunter, 1939 Bgam 1926___________Bagh - Banham 1933___________Bag .___________Becker, et al, 1966 Bgham 1933___________Banh - -- - -----Causey, 1957 Micropterus salmoide$ ------_.Bangham, 1941 E. caeruleus_ -- ---------1949 Smith, E. caeruleus_ Mueller, 1936b ------- N .Y.--E. caeruleus_ ___________Bangham, 1933, 1941b -----Ohio E. caeruleus_ .___________Becker, et al, 1966 E. caeruleus_ -Ark.------------A E. caeruleus_ ------wis.--is-------W nthony, 1963 Fischthal, 1950, 1952, 1947 E. caeruleus_ ------- Wis.--and Allison, 1962 ----- _ __________Kelly E. lizae _____ - la. -A T ex.-------- oberts, 1965 R E. tenax____. -----Roberts, 1970 Tex.--------E. tenax_____ F la .- ------- - - ------Wilson --- ------- Pomoxis nigromaculatus E. caeruleus E. caeruleus__ ------- a--Iow E. caeruleus E. caeruleus E. sp.------------ -- L a. ------------N .J.---- - -- ------.D - Wis.--W is.-----N .--- -----------Causey, 1957, .______Smith, 1949 ------Bangham, 1946 916___________W ------Voth and Larson, ______Fischthal, 1950, 1952, 1947 1968 (Cont.) 60 ALABAMA 60 Percidae Etheostoma caeruleum E. cotti-Etheostoma nigrum E. sp.- ALABAMA AGRICULTURAL EXPERIMENT STATION TABLE AGRICULTURAL EXPERIMENT STATION 1. (Cont.) Ohio--- ________Tidd, 1931 E. sp.- Perca flavescens E. ceeruleus -- - - --- .D.--N ---- .D .--N - -. -- _____Voth and Larson, 1968 ___N___ _Hoffman, 1953 Tidd, 19________31 --------Pearse, 1924 E. E. E. E. E. E. E. E.sp.E. sp.- ceeruleus 2 luciopercarum' caeruleus centrarchidarum luciopercarum 4 luciopercarur 4 luciopercarum 4 - Ohio Wis.- - - ---- is.---W ---- nt. --O ---- nt.?--O ---- is .---W ----Ont. - ----- nt.---O ---- is .-W ____Bangham, 1946 ________Bangham, 1955 Wright, 1882 ---_______.Bere, 1931 Tedla and Fernando, 1969 Telda and Fernando, 1970 .---Marshall and Gilbert, 1905 Voth and Larson, 1968 Stizostedion canadense E. caeruleusE. caeruleusE. caeruleus E. caeruleusE. centrarchidarum Stizostedion vitreum E. caeruleus E. luciopercarum E. caeruleus 12 E. luciopercarum2 E. luciopercarum1 E. centrarchidarum E. centrarchidarum 4 E. luciopercarum E. luciopercarum.----E . luciopercarum-----Pikeperches E. luciopercarum----Mugilidae -Iowa ---. Erie_ L. ---- a --Iow L. Erie__. ---- is.---W ---- is.---W ---O nt.------- Ohio--- --- ________Tidd, ---Wilson, 1916 1931 Bangham and Hunter, 1939 ---------. _____ Wilson, 1916 Tidd, 1931 Bangham and Hunter, 1939 Baugham, 1946 Bangham, 1955 N .Y . Erie__. --L. L. Erie__. Mueller, 1940 Baugham and Hunter, 1939 --- d .- - - -I W ---- is.------- is.---W --Penn.------ ue.- -Q Bangham and Hunter, 1939 -------. Wilson, 1911 Bere, 1931 Roberts, 1970 ---Roberts, 1970 Henderson, 1926 Mugil cephalus E. lizae-------------E. lizae-------------E. lizac-------------E. lizae E . lizae-------------- E. lizae----- -------- E. mugilus----------E. mugilus----------E. mugi lus----------E . mugilus------ ---E . mugilus ----------- -------- ----- ----ex.---T --Tex.------ .C.---N ----la. ---F ----la. ---F --- a. ---G ---- N .C.- ----- Causey, 1953b Pearse, 1952a Tex.---- --- Fla. ---- Pearse, 1947, 1948 Pearse, 1952b Bere, 1936 Roberts, 1970 Wilson, 1911 Causey, 1953b Pearse, 1952b -Causey, --- is.---M Mexico__. - Causey, 1955 1955 Mugil curema E . lizaee-------------E. lizae La-_Fla.. Kroyer, 186314 Pearse, 1952h (Cont.) DISTRIBUTION OF THE GENUS ERGASILUS TABLE - 61 1. (Cont.) Causey, --. 1953a Causey, 1957 E. lizae ---------------------- ere, 1936 Fla.-----------------B E. mugilus------------ La. E. mugilus------------- - La. E. nanus----------------------------- La-- - - Causey, 1953a 'Reexamined, name corrected to E. versicolor from E. elegans by Roberts, (1970). 2 Reexamined (at least in part), name corrected to E. nerkae from E. caeruleus by Roberts (1963, 1970). (at least in part), name corrected to E. centrarchidarumfrom E. caeruleus by Roberts (1970). 4 Originally reported as E. coufusus. (at least in part), name corrected to E. nerkae from E. sp. by Roberts (1963, 1970). 6Reexamined, name corrected to E. cyprinaceus from E. versicolor by Roberts (1970). name corrected to E. arthrosis from E. versicolor by Roberts (1969a). 8Reexamined, name corrected to E. versicolor from E. elegans by Roberts (1969a). Reexamined (at least in part), name changed to E. manicatus from E. by Roberts (1970). 10Reexamined (in part), name changed to E. arthrosis from E. versicolor by Roberts (1969a). 11Reexamined, name corrected to E. centrarchidarumfrom E. nigratus by Roberts, (1970). 'Reexamined (at least in part), name changed to E. luciopercarum from E. caeruleus by Roberts (1970). 13Reexamined, name changed to E. caeruleus from E. centrarchidarum by Roberts (1970). 14 Host reported as Mugil liza. 'Reexamined 'Reexamined 'Reexamined, funduli REVISED PARASITE-HOST CHECKLIST FOR THE GENUS ERGASILUS Host parasite relationships that were reported from material of this study are noted with a single asterisk (*) following the host name. Relationships that were reported from material of this study and represent new records are noted with a double asterisk (* ) Host names followed by (1) represent associations where only a single parasite has been recovered from the respective host, either in this study or the literature. Host names followed by (?) are host-parasite relationships that have been recorded in the literature but are considered questionable for reasons of doubt on correct parasite identification. Ergasilus arthrosis Roberts Alosa chrysochioris (1) Ictalurus I. I. I. I. ° Pylodictis olivaris44 Aphredoderus sayanus0 '* (1) Labidesthes sicculus0 (1) Morone misrissippiensis* 44 M. saxitalis 0 Lepomis gulosus4 Strongylura marina fJurcatus ° melas natalis nebulosus0 punctatus 62 62 ALABAMA AGRICULTURAL EXPERIMENT STATION L. macrochirus* L. marginatus** L. megalotis* 0 L. microlophus L. punctatus*0 Micropterus punctulatus* M. salmoides* Pomoxis nigromacuiatus* Percirna caprodes* (1) Caranx hippos** (1) Achirus iineatus* * Ergasilus centrarchidarum Wright Osmerus mordax Catostomus catostomus (?) Microgadus tomcod (? ) Morcme chrysops0 Amloplites rupestris 0 Centrarchus macropterus Lepomis auritus L. cyanellus* L. gibbosus L. guiosus* L. rnacrochirus* L. punctatus*0 Micropterus dolomieui M. punctulatus* M. saimoides* Pomoxis annuiaris* P. nigromacuiatus* Stizostedion canadense S. vitreum Ergasilus cerastes Roberts Erimyzon sucetta (?) Ergasilus auritus Markevich Giilichthys Onchorhynchus nerka Gasterorteus aculeatus mirabilis Ergasilus caeruleus Wilson Petromyzon marinus Lepisosteus osseus (1) L. spathula Anguilia rostrata (?) Coregonus artedi (?) Coregonus clupeaformis (?) (?) (?) Catostomus catostomus (?) Acrochelis alutaceus Ictalurus catus* I. f urcatus'* Catostomus commersoni (?) Moxostoma valenciennesi (?) Erimyzon sucetta (?) Noturus gyrinus N. f lavus Aphredoderus sayanus* Percopsis omiscomaycus (?) Moron chrysops Ambloplites rupestris Centrarchus macropterus 0 Lepomis auritus * L. cyanellus* L. gibbosus L. gulosus* L. humilus L. macrochirus* L. marginatus** L. megalotis* L. microlophus* L. punctatus* Micropterus dolomieui M. punctulatus M. salmoides Pamoxis annularis* P. nigromaculatus* Perca f lavescens fasciata** Percina nigro Stizostedion canadense S. vitreum S. glaucum Ergasilus celestis Mueller 0 I. nebulosuso I. punctatus ° Ergasilus chatauquaensis Fellows No hosts Ergasilus clupeidarum Johnson and Rogers (?) Fundulus chrysotus Lota iota (?) Alosa aestivalis** A. chrysochloris* Dorosoma cepedianum* D. petenense*0 Etheostoma caeruieus Cottus bairdi Ergasilus cotti Kellicott Ergasilus cyprinaceus Rogers Campostoma anomaium* Hybopsis storeriana* H. winchelli* Nocomis leptocephaius* Notemigonus crysoieucas** Notropis baiieyi* N. chrysocephalus* N. iongirostris** N. hypsilepis* N. roseipinnis* N. texanus*4* N. venustus** N. zonistius 0 Semotilus atromacuiatus Ictalurus nataiis* I. nebulosusO * 0 Noturus nocturnus 0 olivaceus* Fundulus Culaea inconstans (1) Polyodon spathuia* Anguilla rostrata Lota Iota Ergasilus elongatus Wilson DISTRIBUTION OF THE GENUS ERGASILUS 63 DISTRIBUTION OF THE GENUS ERGASILUS Ergasilus feliclithys- 63 Arias felis (1) Bagre marinus* (Pearse) Ergasilus funduli Kroyer E. sucetta* Moxostoma sp. cf. poecilurum Ictalurus melas* I. punctatus Ergasilus nerkae Roberts Osmerus mordax Adinia xenica** Cyprinodon variegatus* Fundulus grandis* F. heteroclitus* F. majalis F. pulvereus* F. similis* Jordanella Gila atraria Hybopsis plumbea Mylocheilus caurinus Ptychocheilus oregonensis Richardsonius balteatus Catostomus catostomus C. Lucania parva* Gambusia af f inis* Poecilia latipinna** Menidia beryllina** Gasterosteus aculeatus (1) G. wheatlandi (1) Ergasilus labracis Kr~yer floridae C. macrocheilus Oncorhynchus nerka Prosopium williamsoni Salmo gairdneri ErgasiLus tenax Roberts fecundus Morone saxatilis* Ergasilus lanceolatus Wilson Dorosoma cepedianum Hiodon tergisus** (1) Esox niger** (1) Lepomis gulosus** L. macrochirus* L. megalotis* L. microlophus*4 L. punctatus**0 Pomoxis annularis* P. nigromaculatus* Ergasilus turgidus Fraser Ergasilus lizae Kr~yer Cyprinodon variegatus* Floridicthys carpio Fundulus grandis* F. similis* Lepomis macrochirus Lagodon rhomboides** Leiostomus xanthurus* Chaetodipterus Oncorhynchus nerka Gasterosteus Cy-matogaster aggregatus Ergasilus versicolor Wilson aculeatus Mugil cephalus* M. curema* faber Lepisosteus osseus* L. platostomus L. spatula Ergasilus luciopercarum Henderson Ambloplites rupestris (1) Perca Stizostedion vitreum Ergasilus megaceros Wilson flavescens Dorosorna cepedianum* * (1) Esox americanus* Esox niger Hybognathus nuchalis** Hybopsis storeriana * Notemigonus crysoleucas Notropis bellus**0 N. edwardraneyi** N. emilae*0 N. roseipinnis* N. shumardi** N. texanus*4 N. venustus*0 Pimephales promelas** P. vigilax* 0 Semotilus corporalis Catostomus commersoni Erimyzon oblongus** Notropis chrysocephalus Catostomus commersoni Erimyzon oblongus* E. sucetta** E. tenuis* Ictiobus bubalus** I. cyprinellus Minytrema melanops**0 Moxostoma poecilurum Ictalurus furcatus I. melas 0 I. rtatalis I. nebulosus" I. punctatus Pylodictis olivaris Morone chrysops (?) Lepomis gulosus (?) Micropterus punctulatus (?) M. salmoides M. interrupta (?) Mugil cephalus** Pogonias cromis**0 (1) Ergasilus wareaglei Johnson 0 (?) Catostomus commersoni * 64 ALABAMA AGRICULTURAL EXPERIMENT STATION 64 Hypenteliumr etowanum H. nigricans* Ergasilus Form A ALABAMA AGRICULTURLEPRMN M. curema Ergasilus Form B TTO Mugil cephalus**4 Mugil cephalus0 Host-parasite associations from the literature that are not applicable to acknowledged species: Ergasilus confusus Bere Coregonus clupeaformis Coregonus sp. Salvelinus namaycush Pimephales promelas Morone americana Ambloplites rupestris Micropterus dolomieui Perca flavescens Sitzostedion vitreum I. natalis Ambloplites repestris Ergasilus mugilus Vogt Elops saurus (1) Micropogon undalatus Leiostomus xanthurus Mugil cephalus Ergasilus nanus Van Beneden Trachinotus carolinus Micropogon uncdatus Mugil curema Ergasilus elegans Wilson Ictalurus melas Ergasilus skrjabini Mueller Morone chrysops Percopsis omiscomaycus SUMMARY AND CONCLUSIONS This study provides a basis for opinions that hitherto had backing. It had been the consensus of most workers that ergasilids exhibited "loose" specificity. Although the specificities of some Ergasilus species have broad ranges, some of the species have narrower preferentialism than most workers had previously surmised. Other species, such as E. arthrosis, have had their ranges of preferred hosts extended by the present study. Perhaps as important as determining the positive host affinities was demonstrating absence of various Ergasilus species on certain host groups. Some of the Ergasilus species were shown to have very narrow specificity. E. elongatus is strictly specific to the paddlefish (Polyodon spathula), and E. celestis is strictly specific to Anguilla rostrata in the Southeast but has been shown in other work to accept the burbot (Lota iota) as a host within its range. That E. celestis infests these two unrelated fishes and not others is indeed a curiosity. Those species that were shown to infest hosts within one main genus are E. cerastes, E. felichthys, F. sp. Form little A, and E. sp. Form B. Other species apparently restrict their parasite activity to families of hosts. E. clupeidarum and E. wareaglei were noted as examples of infesting fishes within a family. E. centrarchidarum was shown to infest one family and at least occasional members DISTRIBUTION OF THE GENUS ERGASILUS 65 of others. Although there is little information to draw on, E. lanceolatus might have similar distribution. The species that were recorded frequently from two or more families represent the remainder of the species in this study. Those species were E. lizae, E. caeruleus, E. cyprinaceus, E. megaceros, E. versicolor, and E. arthrosis. Even though they each had a relatively broad spectrum of host affinities, a limited number of fish were acceptable. It was hoped that the study of specificity would contribute information of a practical nature. Smith's (83) report of a fish epizootic in a reservoir at New Brunswick, New Jersey showed E. caeruleus causing mortality of its preferred hosts but not other species. Kelly and Allison (53) concluded that a centrarchid epizootic that occurred in a brackish water lake in Alabama could be attributed to E. lizae, an ergasilid of mullets. It has been shown herein that E. lizae was actually a common parasite of centrarchids in coastal areas (E. arthrosis). It is our conviction that ergasilids will only cause epizootics of their preferred hosts when conditions favor rapid expansion of the parasite population. The possible exceptions to this assertion are cases where the unusual host is introduced into the population and must undergo a period of acclimatization. There is much unknown about the ecological requirements of the species of Ergasilus. Post-hatching phases in the genus Ergasilus spend a large part of their lives free-living, and no doubt each has adapted to a preferred habitat. Where the hosts and species were obtained in this study provides limited reliable information on the specific habitat requirements since the hosts are not stationary. Some generalizations, however, may be made based on the distribution patterns. The species may be grouped as follows: Coastal - E. felichthys, E. funduli, E. lizae, E. spp. cf. lizae. Estuarine - E. cerastes. Coastal and inland - E. clupeidarum, E. versicolor. Estuarine and inland - E. arthrosis. General inland - E. celestis, E. caeruleus, E. megaceros, E. tenax. Inland - E. centrarchidarum, E. cyprinaceus, E. wareaglei. It is likely that salinity affects the distribution of a number of these species but temperature should not be excluded as a possibility. The distributions of preferred hosts certainly influence the parasites' distributions. 66 ALABAMA AGRICULTURAL EXPERIMENT STATION The interdrainage distributions of the recorded species were shown to be generalized within the study area. Further study will be required to establish objectively the continental distributions for each species. It is apparent, however, that some species range on the Pacific Northwest isolated from the other species of the genus. The literature indicates that there is a zonal pattern of distribution for E. luciopercarum for even though preferred hosts range further southward the species has not been collected below a certain latitude. One species, E. labracis, is apparently restricted in continental distribution to the east coast of North America. Strong endemism has been demonstrated by only one species, E. cotti, but future investigation will probably extend its range. Dispersal of Ergasilus has likely been facilitated by migrations of hosts. No resting stage is known for Ergasilus, and although movement by wind and in association with piscivorous birds or other animals are possibilities, an acceptable host must be sought shortly following displacement. Whatever the case, the parasites appear to be generally distributed where their preferred hosts are successfully established. Incidence, intensity, and other numerical factors were not considered statistically since a certain amount of bias was introduced with incomplete and otherwise inadequate sampling. Generalized statements may be made from listed data from some fish groups. Ergasilids were very common on Bagre marinus, Anguilla rostrata, and the mugilids; they commonly occurred on centrarchids, brackish cyprinodontids and atherinids, catostomids, ictalurids, and percichthyids. The frequency of parasitism by Ergasilus was considered to be low for percids, cottids, and inland cyprinodontids. ACKNOWLEDGMENTS Appreciation is expressed to Dr. Roger Cressey, U.S. National Museum, Washington; Dr. A. Gusev, Zoological Institute, Academy of Sciences, Leningrad; and Mr. W. A. Smith and Dr. K. G. McKenzie, British Museum, London, for loaning type material of various Ergasilus species. Dr. J. W. Cliburn, University of Southern Mississippi; Mr. B. E, Gandy, Mississippi State Wildlife Museum; Dr. H. T. Boschung, Jr., University of Alabama; and Dr. J. S. Ramsey, Auburn University, were kind enough to allow us to obtain host materials DISTRIBUTION OF THE GENUS ERGASILUS 67 from the ichthyological collections of their respective institutions. Without their help this study would have been impossible. Dr. Ramsey aided in providing information on fish distributions and occasional host identifications. Collectors of museum fish specimens were not recorded for acknowledgment for the list would be too extensive. However, several materials of unpublished master's theses added significantly to the materials that were available for this study. These were by Caldwell (19), Gilbert (36), King (54), Martin (61), Mettee (64), and Williams (98). Collections by the late Miss Fannye A. Cook that formed the basis for her book (26) on Mississippi fishes provided much material. The authors are particularly thankful to those that aided in personal collections during the study. Those men were: E. Williams, M. Rawson, S. Chein, J. Sullivan, R. Phelps, J. Ramsey, and W. Armbrester, all of Auburn University. ADDENDUM Since this was written, another Ergasilus species, E. rhinos, has been described* from the nasal fossae of centrarchid fishes of the Black River, North Carolina. The hosts were Centrarchus macropterus,flier; Lepomis gibbosus, pumpkinseed; and Lepomis auritus, redbreast. In the same paper, the authors recorded host relationships for E. megaceros. E. megaceros was noted to be found on the gills of Notemigonus crysoleucas, golden shiner, and Ictalurus natalis, yellow catfish. * BuRRs, K. W. AND G. C. MILLER. 1972. Ergasilus rhinos sp. n. (Copepoda: Cyclopoida) from the nasal fossae of three centrarchid fishes of North Carolina. J. Parasit. 58:600-604. DISTRIBUTION OF THE GENUS ERGASILUS 69 LITERATURE CITED (1) AMERICAN FISHERIES SOCIETY, Committee on Names of Fishes. 1970. A List of Common and Scientific Names of Fishes from the United States and Canada. Parasites of Eastern Wisconsin Fishes. Trans. Parasites Other Than Cestodes in Black (2) ANTHONY, J. D. 1963. BANGHAM, R. V. 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J. 72 ALABAMA AGRICULTURAL EXPERIMENT STATION (57) (58) (59) (60) (61) (62) (63) (64) (65) (66) (67) (68) (69) (70) (71) (72) (73) (74) (75) J. 1889. Parasites of the Striped Bass. Proc. Acad. Nat. Sci. Philadelphia, 1888:125, 166-167. MARGOLIS, L. 1965. Parasites of an Auxiliary Source of Information about the Biology of Pacific Salmons (genus Oncorhynchus). J. Fish. Res. Bd. Canad., 22:1,387-1,395. MARKEVICH, A. P. 1940. New Freshwater Copepoda Parasitica of the Family Ergasilidae. Pratsi Naukovo-doslidnii Inst. Biol. KDU, 4:107-121 (In Ukrainian, summaries in Russian and English). MARSHALL, W. S. AND N. C. GILBERT. 1905. Rep. U.S. Bur. Fish., 1904:513-522. MARTIN, BILLY J. 1963. The Fishes of the Tallahala Creek Drainage System (Jasper, Jones, and Perry counties, Miss.). Unpublished Master's Thesis. Univ. of Sou. Miss. McDANIEL, J. S. 1963. Parasites from the Genus Lepomis (Centrarchidae) in Lake Texoma, Oklahoma. Trans. Amer. Microsc. Soc., 82:423-425. AND H. H. BAILEY. 1966. 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Interscience, New York. 1,104 p. ------------------------ AGRICULTURAL EXPERIMENT STATION SYSTEM OF ALABAMA'S LAND-GRANT UNIVERSITY NN'ith an agricultural research unit in every major soil area, Auburn Uniersitx serves the needs of field crop, livestock, forestrx , and horticultural producers in each regioii in Alabama. Ex ry citizen of the State has a stake in this research program, since any advantage from newv and more econo umical way s of p)roducing and handling farm prolucts dircctlx Ieneflts the consuring public.2 0 iQ 0 00 s ® ° 0 14 4 H i Research Unit Identification * i. 2. 3. 4. 5. 6 7 8 9. 10 11. 12. 13 14. 15. 16. 17. 18. 19. 20. 21. Main Agricultural Experiment Station, Auburn Tennessee Valley Substation, Belle Mina. Sand Mountain Substation, Crossville. North Alabama Horticulture Substation, Cullman. Upper Coastal Plain Substation, Winfield. Forestry Unit, Fayette County. Thorsby Foundation Seed Stocks Farm, Thorsby. Chilton Area Horticulture Substation, Clanton. Forestry Unit, Coosa County. Piedmont Substation, Camp Hill. Plant Breeding Unit, Tallassee. Forestry Unit, Autauga County. Prnttville Experiment Field, Prattville. Block Belt Substation, Marion Junction, Tuskegee Experiment Field, Tuskegee. Lower Coastal Plain Substation, Camden. Forestry Unit, Barbour County. Monroeville Experiment Field, Monroeville. Wiregrass Substation, Headland. Brewton Experiment Field, Brewton. Ornamental Horticulture Field Station, Spring Hill. Gulf Coast Substation, Fairhope. i