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Burhinotaenia colombiana n. sp. (Cestoda, Cyclophyllidea) from the Double-Striped Stone Burhinus bistriatus (Aves, ) in Colombia

Boyko B. Georgiev Bulgarian Academy of Sciences

Éva Murai Hungarian Natural History Museum

Robert L. Rausch University of Washington, [email protected]

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Georgiev, Boyko B.; Murai, Éva; and Rausch, Robert L., "Burhinotaenia colombiana n. sp. (Cestoda, Cyclophyllidea) from the Double-Striped Stone Curlew Burhinus bistriatus (Aves, Charadriiformes) in Colombia" (1996). Faculty Publications from the Harold W. Manter Laboratory of Parasitology. 389. https://digitalcommons.unl.edu/parasitologyfacpubs/389

This Article is brought to you for free and open access by the Parasitology, Harold W. Manter Laboratory of at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Faculty Publications from the Harold W. Manter Laboratory of Parasitology by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. J. Parasitol.,82(1), 1996, p. 140-145 ? AmericanSociety of Parasitologists1996

BURHINOTAENIACOLOMBIANA N. SP. (CESTODA, CYCLOPHYLLIDEA)FROM THE DOUBLE-STRIPED STONE CURLEW BURHINUSBISTRIATUS (AVES, CHARADRIIFORMES)IN COLOMBIA

Boyko B. Georgiev, Eva Murai*, and Robert L. Rauscht Instituteof Ecology,Bulgarian Academy of Sciences, 2 GagarinStreet, 1113 Sofia, Bulgaria

ABSTRACT: Burhinotaeniacolombiana n. sp. (Dilepididae)is describedfrom the small intestine of the double-stripedstone curlewBurhinus bistriatus (Burhinidae) captured at Carimagua,Colombia. The new speciesis distinguishedfrom the most similar Burhinotaeniadelachauxi (Baer, 1925), a parasiteof the Old World Burhinusspp., by the longercirrus-sac (375-590, avg. 514 Aimvs. 322-393, avg. 354 Mm)and longerrostellar hooks (412-451, avg. 440 ,umvs. 358-367, avg. 364 Am).The validity of the BurhinotaeniaSpasskii and Spasskaya,1965 and its genericdiagnosis as proposedby Bona (1994) are confirmed.This is the first recordof a of Burhinotaeniain the New World.

The double-striped stone curlew Burhinus bistriatus (Wagler) DESCRIPTION occurs in Central America and northern South America, from Burhinotaeniacolombiana n. sp. southern Mexico to northern Colombia and northern Brazil (Figs.1-15) (Howard and Moore, 1980). The only record of cestodes from Strobilabandlike, when gravid 66-94 mm (77 ? 9 mm, n = 7) long, this host was reported by Beddard (1913) who described the consistingof 296-356 (314 ? 21, n = 7) proglottides,8-38 (23 + 9, n = = dilepidid tapeworm Eugonodaeum oedicnemi Beddard, 1913 7) of them gravid;maximum width 1,488-2,012 (1,706 ? 180, n Scolex from a at the London Zoo. 7) at gravid proglottides,rarely at pregravidproglottides. (Figs. 1, 2) distinct from neck, roundedlaterally, with maximum width 986- B. bistriatus Three cestode species were found as parasites of 1,135 (1,042 ? 56, n = 7) at level of suckers;apical part forms cuplike in Colombia. The present paper describes 1 of them, which was cavity bearing crown of rostellar hooks; walls of cavity protrusible. found to represent a new species of the genus Burhinotaenia Suckersprominent, oval, highly muscularized,with diameter310-390 = near anterior Spasskii and Spasskaya, 1965. The systematic position of the (340 ? 20, n 28). Rostellumheartlike, wider end, grad- ually taperingposteriorly; in posteriordirection reaching end of scolex other 2 is a of another that will be published species subject study and passing slightly into neck; 734-849 (794 ? 43, n = 7) long and separately. 493-505 (502 ? 6, n = 7) wide. Lackingsaclike rostellarsheath, only stronglydeveloped muscular bundles surround rostellum. Rostellar hooks MATERIALAND METHODS 24-28 (26 ? 1, n = 9) in number,directed anteriorly and arrangedin singlecrown; each hook consists of chitinoidpart (with long and slightly The present descriptionis based on 8 entire specimens, 1 separate curvedblade, shorthandle, and bifid guard),and amorphouscase com- scolex, and a few fragmentscollected by R. L. and V. R. Rausch on 6 prisinghandle and guardof chitinoid part(Figs. 3-7), which takesstain; April 1977 from the small intestine of 1 female B. bistriatus(collection hook measurements:general length (a) 412-451 (440 ? 11, n = 20), no. 42,671) capturedat Carimagua,Colombia. They were fixed in hot length of chitinoid part (b) 336-371 (362 ? 8, n = 20), lengthof blade 10% formalin, stored in 70% ethanol, stained in iron acetocarmine, (c) 246-264 (254 + 4, n = 20), lengthof base of chitinoid part(d) 107- dehydrated,cleared in eugenol, and mounted in Canadabalsam. Two 125 (118 ? 5, n = 19), length of hook base (e) 170-208 (193 + 12, n scoleces were mounted in Berlese'sfluid in order to provide detailed = 20); vary in narrowerlimits in 1 scolex, e.g., (a) 439-451, (b) 356- observationsof rostellarhooks. Mature,pregravid, and gravidproglot- 371, (c) 249-259, (d) 114-123, (e) 192-208 (n = 14). Neck 402-585 tides of 1 specimen were sectioned transverselyby hand with a razor (478 ? 70, n = 7) wide in narrowestpart. Segmentation begins at about blade to facilitatestudy of some anatomicaldetails. 1,444-1,772 (1,590 ? 118, n = 7) from posteriormargin of suckers. Metric and meristic characteristicsare given as the range,the mean Proglottides(Figs. 8-11) craspedote,wider than long; matureproglot- + 1 SD in parentheses,and the number of measurementsor counts tides with ratio 1/4.1-1/2.8 (1/3.3 ? 0.4, n = 10);gravid where otherwise length/width taken (n). The dimensions are in micrometersexcept proglottideswith length/widthratio 1/2.7-1/2.2 (1/2.5 ? 0.2, n = 10). stated. Genital pores irregularlyalternating, mainly in short series, e.g.,... 2, Parasite The type specimens were deposited in the U.S. National 1,2,1, 12,, 4, 4, 1, 1, 1, 111 , 2,, 5,1, 1, 10, 2, 1, 1, 1, 3, 1, 5, Collection, Beltsville (USNPC), the ParasitologicalCollection of the 1, 1, 1, 2, 1, 4, 4, 3, 1, 3; opened in middle of lateralproglottis margin Departmentof Zoology,Hungarian Natural History Museum, Budapest on tall, slightlyexpressed genital papilla. Genital atria (Fig. 12) infun- (HNHM), and the collection of the senior author(BBG). dibular,with well expressedcircular musculature of tubularbase; 38- The type specimens of Burhinotaeniadelachauxi (Baer, 1925) from 48 (43 ? 4, n = 10) deep, with diameter of tubularpart 13-20 (16 ? the collection of the Natural History Museum, Geneva, nos. 6/24-25 2, n = 10). Ventral osmoregulatorycanals 23-59 (38 ? 14, n = 10) (2 slides) and a specimen of Burhinotaeniacoronata from Burhinus wide; with transverseanastomosis along posteriormargin of every pro- oedicnemus(Linnaeus), the Great HungarianPlain, from the Parasi- glottis. Dorsal canals7-13 (9 ? 2, n = 10) wide. Genital Bu- osmoregulatory tological Collection of the HungarianNatural History Museum, ducts between osmoregulatorycanals. dapest, no. 10907 (publishedas B. delachauxiby Muraiet al. [1988]), Testes 17-32 (24 ? 3, n = 38, takenfrom 7 strobila)in number;their were used as comparativematerial. numbervaries in narrowerlimits in 1 strobila,e.g., 17-22 (n = 6), 19- 25 (n = 6), 22-25 (n = 6), 22-27 (n = 6), 24-32 (n = 6); major part of testes (Figs. 8, 9) situated in mature proglottidesin 2 groupslaterally to vitellariumand laterallyand posteriorlyto ovary wings;single testes Received 12 September 1994; revised 21 August 1995; accepted 21 occasionallydisposed posteriorlyand dorsallyto vitellarium;testes do August 1995. not extendlaterally beyond longitudinal osmoregulatory canals; vestigial * Departmentof Zoology,Hungarian Natural History Museum, Baross testes might be presentalong posteriormargin of pregravidand gravid utca 13, H-1088 Budapest,Hungary. proglottides.External vas deferens coiled both ventrally and dorsally t Departmentof ComparativeMedicine SB-42, University of Wash- to cirrus-sac,covered with single layer of large, intensely stained cells ington, Seattle, Washington98195. (Fig. 12);its diameter11-28 (18 ? 6, n = 15). Cirrus-sac(Fig. 12) thick- 140 GEORGIEVET AL.-NEW SPECIES OF BURHINOTAENIA141

.,.

3 4 5 6 7

FIGURES1-7. Burhinotaenia colombiana n. sp. 1, 2. Scoleces, scale bar 250 um. 3-7. Rostellar hooks, scale bar = 100 um. 142 THEJOURNAL OF PARASITOLOGY, VOL. 82, NO. 1, FEBRUARY1996

8

9

10

I I

FIGURES8-11. Burhinotaenia colombiana n. sp. 8, 9. Mature proglottides. 10. Postmature proglottis. 11. Gravid proglottis. All views dorsal. Scale bar = 250 um. GEORGIEVET AL.-NEW SPECIES OF BURHINOTAENIA143

12

FIGURES12-15. Burhinotaeniacolombiana n. sp. 12. Terminalgenital ducts, dorsal view, scale bar = 100 um. 13. Detail of female genital ducts, ventral view, scale bar 50 = ,m; abbreviations:MG, Mehlis' gland;OV, ovary;SR, seminal receptacle;UT, uterus. 14, 15. Eggs,scale bar = 20 jMm. walled, highly elongate, claviform (with wider aporal end), often sig- + 4, n = 10) long and 12-17 (14 ? 2, n = 10) wide at x base; thoroughly moid; its measurements375-590 54-85 (514 ? 44 x 66 + 7, n = evaginated cirrus occurs rarely, cylindrical, 77-107 (93, n = 3) in narrowerlimits in 1 x = long, 49), vary strobila,e.g., 375-491 54-67 (n with diameter 12-15 (13, n = 3); armamentobserved neither on evag- 6), 456-532 x 63-72 (n = 9), 483-554 x 67-81 (n = 8), 528-590 x inated cirrusnor in ductus cirri. = 58-67 (n 8); situated obliquely, cross poral longitudinalosmoregu- Vitellariumcompact, transversely elongate, usually reniform, situated latory canals and about a third or a quarterof it lies in median field. near middle of posteriorproglottis margin. Ovary (Fig. 8) occupies Internalvas deferens 1 or a few only forming coils in aporal part of cirrus- centralthird of median field;consists of 2 separatewings connectedby sac. Cirrususually slightly evaginated,then almost conical, 16-25 (19 narrowisthmus; wings slightly lobed; aporal wing more anteriorthan 144 THEJOURNAL OF PARASITOLOGY, VOL. 82, NO. 1, FEBRUARY1996 poral.Mehlis' gland distinct, globular (Fig. 13). Seminalreceptacle from gary determinedas B. delachauxiand possessingrostellar hooks with fusiformto lemon-shaped,situated in poralhalf of medianfield, anterior length of the blade 115-160 and length of the handle 120-150 (Murai to poral wing of ovary and posteriorto cirrus-sac;201-254 (233 ? 17, et al., 1988) belongs to B. coronata. n = 10) long, 54-81 (66 ? 8, n = 10) wide. Vaginal orifice (Fig. 12) Dr. F. V. Bona (pers. comm.) examined the rostellarhooks of the posteriorto male pore; vagina passingposteriorly to cirrus-sac;vaginal type of B. coronatafrom the Collectionof the NaturalHistory Museum, lumen with diameter 3-5 (4.5 ? 0.7, n = 10), with distinct muscular Geneva (no. 28/42) and confirmedthat they correspondto the drawing walls surroundedby thick cellularsleeve; diameterof vagina (together of Mahon (1954) for the same material;he found that the ratio length with cellularsleeve) 25-32 (29 + 2, n = 10). of blade: length of base of these hooks is 0.85-1.1:1. Uterusinitially appears in matureproglottides (Fig. 9) as transversely Becausethe synonymyproposed by Spasskayaand Spasskii(1978) is elongatesac along anteriorproglottis margin, ventral to ovary;its ends based on a study of the literature,it can be acceptedonly conditionally do not cross longitudinalosmoregulatory canals. This sac expandsgrad- and needs furtherconfirmation grounded on a reexaminationof the ually in posteriordirection and its posterior margin forms numerous relevant specimens. Moreover, the data on the strobilaranatomy of sacculationsin postmatureproglottides (Fig. 10). Gravid uterus (Fig. most of the nominal taxa consideredas synonymsof B. coronataor B. 11) thick-walled,occupies median field, its ends sometimes overlap delachauxiare scarce,and only the morphologyof rostellarhooks can longitudinal ventral osmoregulatorycanals; small portions of uterus be used for reliablecomparisons. The synonymyof Spasskayaand Spas- rarelyin lateralfields; walls persistentup to end of uterinedevelopment. skii (1978), however, reflectsthe presenceof 2 types of rostellarhook Fully developed eggs (Figs. 14, 15) almost globularor with irregular shapein the knownnominal species of Burhinotaenia,and this character shape, with granularouter envelop (not clearlyseen in whole mounts); is the main one applied furtherin the differentialcomparison of the diameter 50-63 (56 ? 3, n = 10). Embryophorevery thick, highly new species. refractive,oval or goblet-like,with diameter 39-46 (44 ? 2, n = 10). The shape of the rostellarhooks of B. colombianan. sp. is close to Oncosphereround to oval or with irregularshape near to oval, thin- those of B. delachauxi.The hitherto reportedlengths of the hooks of walled,with diameter27-34 (31 + 2, n = 10). Embryonichooks 3 pairs, this species are 370 (Baer, 1925), 360-388 (Mahon, 1954), and 366- central pair 18-19 (18.8 ? 0.4, n = 8) long, lateralpairs 15-16 (15.9 375 (Baer, 1959). The present observationson type specimens of B. + 0.4, n = 8) long. delachauxirevealed that their hooks are 358-367 (364, n = 8) long;Dr. F. V. Bona (pers. comm.) measureda minimum length of 337 for the Taxonomic summary rostellarhooks of types of B. delachauxi.The new specieswith rostellar hooks 412-451 is from B. B. bistriatus (440) long clearly distinguished delachauxi Type host: Double-stripedstone curlew, (Wagler). this character.A full betweenthe of Colombia. by comparison anatomy proglottides Typelocality: Carimagua, of B. colombianaand B. delachauxiis difficultdue to the rathercon- Site: Small intestine. tractedcondition of the types of the latterand the scarceliterature data. Specimens: Holotype USNPC 85055 (1 slide). ParatypesUSNPC a considerabledifference in the dimensions of the cirrus 12 HNHM 67008/1-3 Nevertheless, 85056-85057 (2 specimens, slides); (3 specimens, sac was 375-590 x 54-85 x ofB. entire 1 and 2 established,i.e., (514 66) colombiana 3 slides); BBG 1995.6.6.1-4 (2 specimens, scolex, frag- and 322-393 x 40-45 x n = of the of B. 4 (354 42, 6) types delachauxi. ments; slides). In the scolex from of B. available refers to the addition, only the types delachauxi Etymology: The specific name, colombiana, country for has metric charactersunder the of dimensions of where the were collected. measuring range specimens the new species, i.e., diameterof scolex 975, diameterof suckers250- 268 (258, n = 4), rostellum 562 long and 459 wide. Remarks The genus BurhinotaeniaSpasskii and Spasskaya,1965 was erected DISCUSSION for 2 species describedfrom of the family Burhinidae(Charad- riiformes):B. delachauxi(Baer, 1925) (type species)and Burhinotaenia After the separation of Burhinotaenia from Paricterotaenia were separatedfrom the genus megistacantha(Fuhrmann, 1909). They by Spasskii and Spasskaya (1965), its validity was supported ParicterotaeniaFuhrmann, 1932 on the basisof theirenormous rostellar hooks (about3 times largerthan the hooks of the type-speciesP. porosa only in a subsequent publication of the same authors (Spasskaya (Rudolphi, 1810))and some peculiaritiesof genitalorgans, in particular and Spasskii, 1978). Schmidt (1986) recognized B. delachauxi the structureof the graviduterus considered to form egg capsules(Spas- and B. coronata as members of the genera Polycercus Villot, skii and Spasskaya,1965). Accordingto Spasskayaand Spasskii(1978), 1883 (syn. Paricterotaenia) and Choanotaenia Railliet, 1896, the Burhinotaeniaincludes 2 valid species: (1) Burhinotaenia genus he Burhinotaenia as a of delachauxi (Baer, 1925) Spasskii and Spasskaya,1965, synonyms Ic- respectively, i.e., regarded synonym terotaeniadelachauxi Baer, 1925; Choanotaeniadelachauxi (Baer) Lo- Polycercus although the generic name was not included in the pez-Neyra, 1935; Paricterotaeniacoronata of Mahon (1954) and Baer list of synonyms. Recently, Bona (1994) validated Burhino- (1959); and (2) Burhinotaeniacoronata (Creplin, 1829) Spasskayaand taenia and amended its generic diagnosis. Taeniacoronata 1829;Choanotaenia Spasskii,1978, synonyms Creplin, The present study confirms the peculiar structure of the ros- coronata(Creplin) Fuhrmann, 1909; Paricterotaenia coronata (Creplin) Fuhrmann, 1932; Choanotaeniamegistacantha Fuhrmann, 1909; Ic- tellar apparatus of Burhinotaenia spp., characterized by the ab- terotaeniamegistacantha (Fuhrm.) Baer, 1925; Paricterotaeniamegis- sence of a saclike rostellar sheath. It is completely different from tacantha (Fuhrm.) Fuhrmann, 1932; Sacciuterina megistacantha the rostella of the majority of dilepidid genera, including these (Fuhrm.)Mathevossian, 1963; Burhinotaeniamegistacantha (Fuhrm.) of Paricterotaenia, Choanotaenia and Polycercus. According to and 1965; Choanotaeniadelachauxi var. mesacan- Spasskii Spasskaya, Bona this of termed as onder- tha Lopez-Neyra,1935; Choanotaeniamagnihamata Burt, 1940. (1994), type rostellar apparatus, Fuhrmannand Baer (1943) re-examinedthe originalCreplin's spec- stepoortioid, occurs only in the genera Burhinotaenia, Onder- imens and recognizedP. delachauxias a junior synonymof P. coronata. stepoortia Ortlepp, 1938, and Spreotaenia Spasskii, 1969. How- This synonymy was followed by Mahon (1954) and Baer (1959). In ever, Burhinotaenia differs obviously from the latter 2 genera contrast to the conclusion of Mahon her of (1954), precise drawings by its strobilar morphology. Therefore, the present study on the rostellarhooks from the types of P. coronataand from her specimens from a charadriiformhost from Zaire clearlydemonstrated 2 different new species and its comparison with the types of the type species types of hook shape. On the basis of this publication,Spasskaya and of Burhinotaenia confirm the opinion of Bona (1994) about the Spasskii (1978) considered the shape of rostellarhooks as the main validity of this genus and do not support the generic allocations distinguishingcharacter between B. delachauxi and B. coronata:the of its species as proposed by Schmidt (1986). formeris characterizedwith ratio of blade: of base about length length The results are also in with the 1.5:1;the latter has rostellarhooks with almost equal lengths of blade present agreement generic and base. Consideringthis differentiatingfeature, the conclusionis that diagnosis of Burhinotaenia proposed by Bona (1994). However, the specimen of Burhinotaeniafrom Burhinusoedicnemus from Hun- there is an exception with respect to the structure of the gravid GEORGIEVET AL.-NEW SPECIES OF BURHINOTAENIA145 uterus. According to the key of Bona (1994), the uterine walls (Geneva) for lending the type specimens of B. delachauxi. This of Burhinotaenia are not persistent and the eggs are loose in investigation was supported by the National Scientific Research parenchyma in the final stage of uterine development. In well Foundation of the Republic of Bulgaria, grants B-44/1991 and stained specimens of the new species, the uterine walls are seen B-404/1994 (to B.B.G.). up to the end of the development of the organ. The peculiar scolex structure of Burhinotaenia and the other LITERATURECITED 2 genera with an onderstepoortioid rostellar apparatus makes BAER, J. G. 1925. Cestodes nouveaux du Sud-Ouest de l'Afrique. questionable their position within the family Dilepididae be- Revue suisse de Zoologie 31: 529-548. cause most of the genera (including the type genus Dilepis Wein- ? 1959. Helminthes Parasites. In Explorations des Parcs Na- land, 1858) possess rostella with saccular sheaths. The charac- tionaux du Congo Belge. Mission J. G. Baer-W. Gerber (1958). teristics that the absence of a saccular rostellar sheath, in com- Fascicule 1. Institut des Parcs Nationaux du Congo Belge, Bruxelles, Belgium, p. 1-163. bination with the saclike uterus without a paruterine organ and BEDDARD,F. E. 1913. Contribution to the anatomy and systematic the avian host, resemble cestodes of the family Metadilepididae. arrangement of the Cestoidea. XI. On a new tapeworm from Oed- The metadilepidids are a small family (8 genera) of avian par- icnemus. Proceedings of the Zoological Society of London 58: 861- asites morphologically similar to both Dilepididae and Paru- 877. F. V. 1994. Railliet & 1909. In L. can be from the former the BONA, Family Dilepididae Henry, terinidae; they distinguished by F. Khalil, A. Jones, and R. A. Bray (eds.). Keys to cestode parasites absence of a saccular rostellar sheath and the position of the of vertebrates. CAB International, Wallingford, U.K., p. 443-554. developing uterus dorsal to the ovary, and from the latter by FUHRMANN, O., ANDJ. G. BAER. 1943. Mission biologique Sagar-Omo the absence of a paruterine organ (Kornyushin and Georgiev, (Ethiopie meridionale), 1939, dirigee par le professeur Eduardo Bulletin de la Soci6te Neuch&teloise des Sci- 1994). In spite of the similarity of Burhinotaenia with the me- Zavattari. Cestodes. ences Naturelles 68: 113-140. tadilepidids, it cannot be placed among them because of the HowARD, R., ANDA. MooRE. 1980. A complete checklist of the birds ventral position of its uterus to the female glands. It seems that of the world. Oxford University Press, Oxford, U.K., 701 p. none of the cyclophyllidean families erected up to the moment KoRNYUSHIN,V. V., AND B. B. GEORGIEV1994. Family Metadilepi- may harbor the genus Burhinotaenia, but the erection of a new didae Spasskii, 1959. In L. F. Khalil, A. Jones, and R. A. Bray (Eds.). Keys to cestode parasites of vertebrates. CAB International, family could be well grounded only as a part of a thorough Wallingford, U.K., p. 585-593. revision of the order. Therefore, Burhinotaenia is tentatively MAHON,J. 1954. Tapeworms from the Belgian Congo. Annales du retained in the Dilepididae pending further revisions ofthe order Mus6e Royal du Congo Belge, C, Zoologie, Series V 1: 135-263. Cyclophyllidea. In its present composition, the family Dilepi- MURAI,E., T. SULGOSTOWSKA,AND F. TENORA 1988. Tapeworm par- asites of Burhinus oedicnemus(L., 1758) in Hungary (Cestoda: didae is a heterogeneous group. This is also the opinion of Bona Progynotaeniidae and Dilepididae). Acta Zoologica Hungarica 34: (1994) who believed that the 100 genera referred to this family 379-392. do not form a monophyletic assemblage and suggested further SCHMIDT,G. D. 1986. Handbook of tapeworm identification. CRC studies in order to elaborate a new suprageneric arrangement Press, Boca Raton, Florida, 675 p. of the group. SPASSKAYA,L. P., ANDA. A. SPASSKII.1978. Cestodes of birds in the USSR. of aquatic birds. Izdatel'stvo 'Nauka,' Moscow, Until now, the species of the genus Burhinotaenia were known Dilepidids 315 p. [In Russian.] only from Europe, Africa, and south Asia (Spasskaya and Spas- SPASSKII,A. A., AND L. P. SPASSKAYA.1965. Revision of the genus skii, 1978; Schmidt, 1986). This is the first record of cestodes Paricterotaenia (Cestoda: Dilepididae). Parazity Zhivothykh i Ras- of this genus in the New World. teniy 1: 84-103. [In Russian.]

ACKNOWLEDGMENTS We are grateful to F. V. Bona (Torino) for providing unpub- lished information on Burhinotaenia spp. and to C. Vaucher