Hosts and Geographic Distribution of Acanthocephalus
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Proc. Helminthol. Soc. Wash. 52(2), 1985, pp. 210-220 Hosts and Geographic Distribution of Acanthocephalus (Acanthocephala: Echinorhynchidae) from North American Freshwater Fishes, with a Discussion of Species Relationships OMAR M. AMIN Department of Biological Sciences, University of Wisconsin-Parkside, Box 2000, Kenosha, Wisconsin 53141 ABSTRACT: Populations of the three known species of the genus Acanthocephalus from North American fresh- water fishes are found in the Mississippi River drainage system, or waters previously connected to it, in the Mobile Bay drainage, Great Lakes, and in New England. Acanthocephalus dirus (Van Cleave, 1931) Van Cleave and Townsend, 1936 has the widest host and geographical distribution and exhibits the greatest morphological variability compared to A. tahlequahensis Oetinger and Buckner, 1976 and A. alabamensis Amin and Williams, 1983. The Wisconsin-Lake Michigan population of A. dirus was geographically isolated from Mississippi River A. dirus less than 15,000 years ago. The New England population of A. dirus may have originated in a like manner. Given sufficient time and continued isolation, the Wisconsin-Lake Michigan and New England pop- ulations may diverge enough to achieve a more distinct taxonomic status. The proposed associations provide satisfactory explanation of present geographical distribution, host relationships, and degree of morphological diversification. A critical appraisal of the taxonomic affinities smaller individuals of the two southern species, and associations within Acanthocephalus from A. tahlequahensis and A. alabamensis. It is herein North American freshwater fishes may only be proposed that the present distribution of the Wis- attained through detailed studies of intraspecific consin-Lake Michigan population of A. dirus, variability as well as ecological and host asso- and possibly that from New England, occurred ciations (Amin, 1975). The subject matter of this through dispersal from a Mississippi River source paper could not have been explored until after similar to A. dirus followed by geographical iso- the intraspecific variability of the most variable lation. species, Acanthocephalus dirus, had been stud- ied, the species redescribed (Amin, 1984), and 1. Geographical distribution the intra- and interspecific variability in all three Acanthocephalus dirus has the widest range of species analyzed (Amin and Huffman, 1984). geographical distribution in North America Additionally, knowledge of host and geograph- compared to the two other species. Practically ical distribution and geological history, made the all its Mississippi River populations studied and/ description of the intra- and interspecific rela- or reported in Amin (1984 and this paper) were tionships of this group of acanthocephalans pos- taken from tributaries of the Mississippi River sible. drainage system in 10 states of the United States. Only those collections from Ohio and Lake Erie Materials and Methods (Nos. 17-19, Fig. 1) are from waters not presently The characteristics of the three species of Acantho- connected to the Mississippi River but which cephalus discussed in this report are based on the stud- were connected via the Wabash River to glacial ies by Amin (1984) and Amin and Huffman (1984). Lake Maumee in the Erie basin during the late Sources of each of the host and geographic distribu- tional records used in the map (Fig. 1) and Table 1 are Woodfordian Substage of the Wisconsinan gla- footnoted in the legend to the map and given the same ciation about 14,000 B.P. (before the present) numbers. (Hough, 1958;Goldthwaitetal., 1965;Fullerton, 1980). The two northern populations of A. dirus Results and Discussion add three more states to that species: New Three species of Acanthocephalus (A. dirus, A. Hampshire, Massachusetts, and Michigan. The tahlequahensis, A. alabamensis) are so far known Michigan material was obtained by Muzzall (pers. from North American freshwater fishes. These comm., 1982) from the Rogue River, a tributary appear to fall into two presumably natural groups of the Grand River which flows into Lake Mich- distinguished primarily by size. Acanthocephalus igan. dirus, includes larger forms compared to the Compared to A. dirus, the distribution of the 210 Copyright © 2011, The Helminthological Society of Washington 211 • A. diruSi MISSISSIPPI R. A A. dirUS; WIS.- L. MICHIGAN • A., tahlequanensis A A- alobamensis © Acanthocephalus sp. Figure 1. Map of eastern United States showing the present distribution of Acanthocephalus spp. from freshwater fish in the Mississippi River drainage system, Great Lakes and the St. Lawrence watershed as well as from New England, Nova Scotia, and Alabama. Numbers refer to the following collections: 1: Arnold et al. (1967,1968); 2: Van Cleave (1931) (specimens from Ictiobus sp. in USNM Helm. Coll. No. 37600); 3: Oetinger and Nickol (1981) (host species undisclosed); 4: Bangham and Venard (1942); 5: Ewell (1953); 6: Gleason (pers. comm., 1982), McDonough and Gleason (1981), Talton and Gleason (1978); 7: Van Cleave and Townsend (1936); 8: Page (1974); 9: White (1974, pers. comm., 1981), White and Harley (1973, 1974); 10: Combs et al. (1977); 11: Huffman (pers. comm., 1981); 12: Gash et al. (1972), R. Gash (pers. comm., 1982); 13: Camp (1977, pers. comm., 1981), Camp and Huizinga (1980); 14: Seidenberg (1973); 15: Schmidt et al. (1974); 16: Amin (this paper); 17: Muzzall and Rabalais (1975a, b), Muzzall (pers. comm., 1982); 18: Bangham (1972); 19: Venderland (1968); 20: Collins and Dechtiar (1974); 21: Dechtiar and Berst (1978), Dechtiar (pers. comm., 1983); 22: Frechette et al. (1978); 23: Wiles (1975); 24: Bullock (1962, 1963, pers. comm., 1982-1984); 25: Amin (1975); 26: Amin (1977a, unpubl.), Amin and Burrows (1977); 27: Amin (1977b); 28: same as for E. salmonis in Amin (1978a); 29: Oetinger and Buckner (1976); 30: Amin and Williams (1983), Williams and Rogers (1982), Williams (1974a, b); 31: Burns (1970, 1971); 32: Skelly (pers. comm., 1982); 33: Muzzall (pers. comm., 1982). All speci- mens from Canada (20-23) are regarded as Acanthocephalus sp. until further study. Copyright © 2011, The Helminthological Society of Washington 212 PROCEEDINGS OF THE HELMINTHOLOGICAL SOCIETY Table 1. Comparative distribution of adult Acanthocephalus spp. in North American freshwater fish hosts. Species of Acanthocephalus and host records (reference numbers correspond to those in map, Fig. 1) Acanthocephalus dims Acantho- Acantho- rpnhnhis renhnhis ttrnnthn- Mississippi Wisconsin-Lake New tahle- alaba- cephalus Fish host* River Michigan England quahensis rnensis sp.f Amiidae Amia calva* (16, 18) Anguillidae Anguilla rostrata 24 Catostomidae Catostomus commersoni* (9, 16, 17) (25,26,27,33) (24) 20,21 Hypentelium etowanum 30 H. nigricans* (6, 17) 21 Ictiobus sp.* (2) Minyatrema melanops 10 Moxostoma duquesnei 6 M. erythrurum* (6, 10, 32) M. macrolepidotum 32 M. poecilurum 30 Centrarchidae Ambloplites rupestris 16, 17 Lepomis cyanellus* (3, 16, 17) (25, 27) L. gibbosus* (16, 17) 24 L. macrochirus* (1,7, 16, 17) (25) L. megalotis 6 1 L. microlophus 1 Microptems coosae 30 M. salmoides* (7) (25) Pomoxis nigromaculatus* (16) Clupeidae Alosa pseudoharengus 26 20,21 Cottidae Cottus bairdf* (5, 15) C. cognatus* (26) 21 Cyprinidae Campostoma anomalum 17 Carassius auratus* (17) Cyprinus carpio* (16,17) (26) 30 Hybopsis sp. 14 Nocomis asper 29 Notemigonus crysoleucas* (17) (25) 24 Notropix cornutus* (11) N. crysocephalus 17 N. hudsonius 17 N. pilsbryi 29 N. spllopterus 17 N. umbratilis* (17) Pimephalus notatus 14 P. promelas* (25) Semotilus atromaculatus* (6, 16, 13, 17) (25) 24 S. margarita 25 Cyprinodontidae Fundulus diaphanns 23 Esocidae Esox sp. (24) Copyright © 2011, The Helminthological Society of Washington OF WASHINGTON, VOLUME 52, NUMBER 2, JULY 1985 213 Table 1. Continued. Species of Acanthocephalus and host records (reference numbers correspond to those in map, Fig. 1) Acanthocephalus dims Acantho- Acantho- rcnhnliis rcnhnhi*; Acantho- Mississippi Wisconsin-Lake New tahle- alaba- cephalus Fish host* River Michigan England quahensis mensis sp-t E. arnericamts* (11) (24) E. lucius* (16) E. niger 24 Gadidae Lota lota 26 Microgadus tomcod 24 Gasterosteidae Culaea inconstans 25 Ictaluridae Ictalurus melas* 16 (25) I. natalis* (12, 16) I. nebulosus (24) 22 I. punctatus 7, 16 Osmeridae Osmerus mordax* (26) 20 Percidae Etheostoma helium 6 E. blennoides 6, 17 E. coeruleum 6 E. punctulaturn* (29) E. spectabile 29 E. squamiceps 6,8 E. stigmaeum 6 31 E. whipplei 31 E. zonale 6 Perca flavescens* (16) (24) 20 Percina caprodes 6 Stizostedion vitreum 16 Salmonidae Coregonus clupeaformis 20 C. hoyi 26 Onchorhynchus kisutch* (26) O. nerka 20 O. tshawytscha* (26, 28) Salmo gairdnerf* (25, 26, 33) (24) S. irideus* (5) S. trutta 26, 33 24 Salvelinus fontinalis* 33 (24) S. fontinalis x 51. namaycush 20,21 S. namaycush* (26) Sciaenidae Aplodinotus grunniens* (2,4,7, 19) Umbridae Umbra limi* (16) No. of host species (families) 46(11) 22(10) 14(9) 4(2) 6(4) 11(9) * Fishes marked with asterisks are those from which gravid worms were reported from populations/species with parenthesized reference numbers. t All specimens from Canada (20-23) are regarded as Acanthocephalus sp. pending examination. Copyright © 2011, The Helminthological Society of Washington 214 PROCEEDINGS OF THE HELMINTHOLOGICAL SOCIETY Table 2. Geographical variation in the number of proboscis hooks per row in Acanthocephalus dims* No. hooks/row (mean) TV Species Population Males Females A. dirus Mississippi (Van Cleave) 11-13 (?) 11-13 (?) (MISS) Mississippi (type, Amin) 11-12(11.5)3 11-13(12.3)5 Kentucky (Gleason, White) 7-12 (9.5)21 8-14(10.9)59 West Virginia (Huffman) 6-11 (8.6)7 8-11 (9.1)15 Illinois (Camp, Seidenberg, 9-12(10.5)25 9-13(10.3)53 Schmidt) Ohio (Muzzall) 7-11 (9.3)18 8-11 (9.2)17 Wisconsin (Amin) 7-12(10.3)39 9-13(10.8)83 All A. dirus (MISS) 6-13 (9.9)113 8-14(10.6)232 A. dirus Wisconsin (Amin) 8-11 (9.3)142 8-11 (9.6)206 (WI-LM) A. dirus New England (Bullock) 7-10 (8.3) 167 7-10 (8.9) 130 (N-ENG) * Listed in order of northward distribution.