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Proceedings of the Helminthological Society of Washington PROCEEDINGS OF THE HELMINTHOLOGICAL SOCIETY OF WASHINGTON VOLUME 45 JANUARY 1978 NUMBER 1 Systematic Status of Proteocephalid Cestodes from Reptiles and Amphibians in North America with Descriptions of Three New Species1 DANIEL R. BROOKS2 Parasitology Section, Gulf Coast Research. Laboratory, Ocean Springs, Mississippi 39564 ABSTRACT: Based on collections of cestodes from reptiles and amphibians primarily in the southeast- ern United States, three new species of Proteocephalus are described and supplemental data for 16 ad- ditional species are presented. Proteocephalus amphiumicola sp. n. from Amphiuma means in south- ern Mississippi and southern Alabama most closely resembles P. amphiumae and P. alternans from which it differs in scolex morphology and number of uterine branches. It is unique in possessing spi- raling rather than sinuous lateral longitudinal osmoregulatory ducts and has a vaginal "sphincter" similar to that in P. alternans. Proteocephalus aberrans sp. n. from Siren lacertina in northern Florida is unique among known members of the genus because it lacks lateral uterine branches. Proteoceph- alus variabilis sp. n. from Natrix cyclopion cyclopion in southern Louisiana and N. rhombifer in northern Louisiana resembles P. perspicua in size of scolex and number of uterine branches, but dif- fers from it in average number of testes, position of genital pore and shape of ovary in gravid pro- glottids; it resembles P. agkistrodontis in the last three characteristics. New hosts for P. perspicua include N. c. cyclopion, N. c. floridense, and N. fasciata confluens; it also parasitized N. rhombifer and N. sipedon. The report of P. perspicua in northern and southern Florida, northern and southern Louisiana, and southern Mississippi represents new locality records. Proteocephalus faranciae, known previously from Texas in Farancia abacura, infected F. abacura in northern Florida and N. c. flori- dense in southern Florida. Natrix c. cijclopion and N. c. floridense represent new hosts for P. grandis, and both northern and southern Florida and Louisiana are new localities. Agkistrodon piscivorus hosted P. marenzelleri in northern and southern Louisiana and P. agkistrodontis in northern Louisi- ana, all new localities. Proteocephalus sireni, known previously from Siren intermedia in southern Illi- nois, was collected from the same host in southern Louisiana. Proteocephalus testudo from Trionyx spiniferm in Nebraska and Indiana and Chrysemys scripta elegans in Illinois, the latter two being new localities, more closely resembles P. australis from a teleost fish than any species from amphibians or reptiles. Circumstantial evidence suggests that P. magnus, P. olor, P. saphenus, and P. gracilis, all from aquatic frogs of the genus Rana, are geographical variants of a single species. The reports also includes results of examining type-material and deposited specimens of P. agkistrodontis, P. perspicua, P. marenzelleri, P. grandis, P. olor, P. magnus, P. saphenus, P. cryptobranchi, P. filaroides, P. loenn- bergii, P. amphiumae, and P. alternans. Since, on the basis of the new material, no distinctions exist among the genera Ophiotaenia, Batrachotaenia, and Testudotaenia from herptiles and Proteocephalus from teleost fishes other than host type, Proteocephalus is considered the senior synonym of all four, and appropriate new combinations are made. Analysis of the interspecific relationships of the 18 spe- cies parasitizing amphibians and snakes in North America reveals distinct groupings according to host category. A monophyletic origin of the species parasitizing caudate amphibians is postulated based on their morphological and geographical homogeneity and well-defined host-parasite relation- ships. The absence of P. perspicua in N. erythrogaster flavigaster may be explained on the basis of feeding and habitat preferences of the host species. A key distinguishes the 19 species of Proteoceph- alus parasitizing herptiles in North America. A number of proteocephalid cestodes have lrThis study was conducted in part by a grant entitled hppn dpspribprl as nflrasitpQ nf rprvtilpc. anrl "Studies on helminths of the northern Gulf of Mexico" °, af scnoea as parasites or reptiles- ana from the state of Mississippi to Robin M. Overstreet. amphibians in North America. All have med- " Present address: Department of Biology, University 11 j j • i i- i L r of Mississippi, University, Mississippi 38677 ' ullary gonads and simple scohces, but for most Copyright © 2011, The Helminthological Society of Washington PROCEEDINGS OF THE HELMINTHOLOGICAL SOCIETY of these species little descriptive information Proteocephalus amphiumicola sp. n. exists other than brief original descriptions. (Figs. 1-6) Because of this, considerable differences of opinion have resulted concerning their sys- DESCRIPTION (based on 20 specimens): tematic positions. The present work describes Strobila up to 300 mm long. Scolex aspinose, three new species and presents supplemental 309-390 (352) wide, with indistinct apical data for 16 more; a diagnosis accompanies organ and four shallow, padlike suckers; suck- species for which new material was collected; ers 222-297 (253) long by 117-155 (133) salient features useful in deducing phyloge- wide. Immature proglottids wider than long. netic relationships are provided for other spe- Mature proglottids 662-1,576 (1,040) long by cies. Based on that data, the study also ex- 394-820 (678) wide. Ratio of proglottid width amines the generic status of all proteocephalids to length 1:0.9-3.8 (1:1.9). Testes 63-136 with medullary gonads and simple scolices; it (.1.05), 7-18 (13) preporally, 25-53 (37) further considers the interspecific relationships postporally, 31-70 (55) antiporally; 31-68 in of those species parasitizing reptiles and am- diameter. Cirrus sac 110-205 (157) long by phibians in North America. 63-95 (80) wide; ratio of cirrus sac length to proglottid width 1:2.5-4.2 (1:3.4); basal Materials and Methods portion of cirrus thick-walled, up to 25 thick. Genital atrium without papillae. Genital pores Cestodes were relaxed in cold tap water or alternating irregularly in anterior 12-28% saline, fixed with AFA, and stored in 70% (22%) of proglottids. Ovary bilobed with ir- ethanol. After staining with Harris' hema- regularly shaped lobes expanded anteriorly, toxylin, Mayer's hematoxylin, or Ehrlich's acid 371-476 (425) wide; lobes 148-340 (250) hematoxylin, worms were mounted in Histo- long. Vagina anterior to and never crossing clad. Unless otherwise stated, measurements cirrus sac; muscles surrounding vagina near of holdfast structures were based on examina- terminal end acting as sphincter; seminal re- tion of 25 scolices, and those of proglottids ceptacle present. Uterus preformed in mature and their organs on 100 proglottids, since mean proglottids. Vitelline follicles extending along values for certain characteristics were found entire length of proglottid; follicles not in to be taxonomically useful. The cirrus of pro- single file; follicles and ovarian lobes proxi- teocephalids typically comprises a basal and mate posteriorly. Gravid proglottids 820-3,814 distal portion. The thickness of the basal por- (1,850) long by 394-820 (678) wide; ratio tion appears to be related to the length of the of proglottid width to length 1:2.5-5.0 (1: cirrus, but since it is difficult to obtain speci- 3.8). Uterus with 39-98 (59) lateral branches mens with everted cirri, cirrus length has not occupying 41-84% (65%) of proglottid width; proved to be useful taxonomically; I have, preformed ventral uterine pores lacking. Eggs however, noted which examined species have 49-62 in diameter, oncospheres 25-37. Ex- thick- and thin-walled basal portions. Nota- cretory system composed of paired dorsal and tions for deposited specimens are: USNM ventral spiraling lateral longitudinal medul- Helm. Coll. for United States National Mu- lary ducts and cortical network of reticulate seum Helminthological Collection, Beltsville, tubules; tubules anastomosing near posterior Maryland and HWML for Harold W. Man- end of proglottid. ter Laboratory, Division of Parasitology, Uni- HOST: Amphiuma means Garden, two-toed versity of Nebraska State Museum, Lincoln, conger eel. Nebraska. Measurements are in micrometers SITE OF INFECTION: Upper small intestine. unless otherwise noted, with averages in pa- LOCALITIES AND PREVALENCE: St. Andrew's, rentheses and previously known data in brack- Jackson County, Mississippi (2/4 (type); Fowl ets; figures were drawn with the aid of a River, Theodore, Mobile County, Alabama drawing tube. (1/D. Figures 1-6. Proteocephalus amphiumicola. 1. Scolex. 2. Ootype region. 3. Terminal genitalia. 4. Immature proglottid. 5. Mature proglottid. 6. Gravid proglottid. Copyright © 2011, The Helminthological Society of Washington OF WASHINGTON, VOLUME 45, NUMBER 1, JANUARY 1978 Oi 0 100pm Copyright © 2011, The Helminthological Society of Washington PROCEEDINGS OF THE HELMINTHOLOGICAL SOCIETY 8 Copyright © 2011, The Helminthological Society of Washington OF WASHINGTON, VOLUME 45, NUMBER 1, JANUARY 1978 HOLOTYPE: USNM Helm. Coll. No. 74330. expanded anteriorly, 532-798 wide; lobes 228- PARATYPES: USNM Helm. Coll. No. 74331; 342 long. Vagina anterior or posterior to cir- HMWL No. 20286 and 20324. rus sac, crossing it or not; vaginal sphincter ETYMOLOGY: The specific name means absent, seminal receptacle present. Uterus "Amp/m«7ia-inhabiting," and is derived from preformed in mature proglottids. Vitellaria the Greek "amphi" meaning on both sides, extending
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