the trematode genus glypthelmins stafford, 1905...

116 PROCEEDINGS OF THE HELMINTHOLOGICAL SOCIETY

pleii, in Puerto Rico. Proc. Helm. Soc. Wash.21: 64.

Nasir, P., and M. T. Diaz. 1971. A redescrip-tion of Mesocoelium monas (Rudolphi, 1819)Freitas, 1958, and specific determination inGenus Mesocoelium Odhner 1910 (Trema-toda: Digenea) Rivista Parassit. 32: 149-158.

Pereira, C. 1935. Os Oxyurata parasites de La-certilia do nordeste Brasileiro. Arch. Inst.Biol. 6: 5-27.

Self, J. T., and J. Garcia-Diaz. 1961. Railli-etiella (Heymonsia) hemidactyli Hett, 1934,from Hemidactijlus mabouia in Puerto Rico,with a correction of the identity of R. hebiti-hamata Self and Kuntz, 1961. J. Parasit. 47:912.

Simha, S. S. 1965. Allopharynx leiperi, a newplagiorchiid trematode from an Indian watersnake. J. Parasit. 51: 215-216.

Sharma, P. N., and A. N. Gupta. 1971. Re-view of genus Glossimetra Mehra, 1937 and

proposed synonymy of G. tamiansis Dwivedi,1967 and G. narmadi Dwivedi, 1967 with G.orientalis Mehra, 1937.

Thapar, G. S. 1926. Ozolaimus Dujardin, 1845( — Macracis Gedoelst, 1916), a littl e knownnematode genus from the caecum of Iguanatuberculata. J. Helminth. 4: 69-74.

Webster, J. D. 1943. Helminths from the robin,with the description of a new nematode, Por-rocaecum brevis-piculum. J. Parasit. 29: 161-163.

Whittaker , F. H., G. D. Schmidt, and J. Garcia-Diaz. 1970. Helminth parasites of somebirds in Puerto Rico. Proc. Helm. Soc. Wash.37: 123-124.

Yamaguti, S. 1961. Systema Helminthum Vol.IV . The Nematodes of Vertebrates, parts Iand II , Interscience Publishers, N.Y. pp. 1261.

. 1971. Synopsis of Digenetic Trema-todes of Vertebrates Vol. I. 1074 p., KeigakuPublishers, Tokyo.

The Trematode Genus Glypthelmins Stafford, 1905(Plagiorchioidea: Macroderoididae) with a Redescription ofG. facioi from Costa Rican Frogs1 ~

JAMES J. SULLIVAN S

Department of Zoology, University of Georgia, Athens, Georgia 30601

ABSTRACT: The 28 described species of glypthelminth trematodes are divided into two groups on thebasis of the form of the excretory bladder. Those possessing a Y-shaped bladder are referred to thePlagiorchiidae while those exhibiting an I-shaped bladder are placed in the Macroderoididae. The genericdiagnosis of Glypthelmins is emended and the genus is included in the Macroderoididae. Glypthelminsquieta (Stafford, 1900), G. rugocaudata (Yoshida, 1916), G. staffordi Tubangui, 1928, G. shastai Ingles,1936, and G. facioi Brenes Madrigal, Arroyo Sancho, Jimenez-Quiros, and Delgado Flores, 1959 arerecognized as valid species. A redescription of G. facioi based on specimens collected from Rana pipiensin Costa Rica is presented. The presence of spines or tegumental scales, egg size, size of the testes, andlength of the esophagus are used to distinguish these five species. Reynoldstrema africana (= G. africana)is transferred to the Astiotrematinae (family Plagiorchiidae), and G. diana is designated incertae sedis.

In June, 1969, 34 specimens of Glypthelminsfacioi Brenes Madrigal et al., 1959, were

1 Supported in part by grant AI 10051 (UC ICMR) tothe Department of International Health, School of Medi-cine, University of California, San Francisco, from theNational Institute of Allergy and Infectious Diseases, Na-tional Institutes of Health, U.S. Public Health Service, andin part by Travel Funds provided by the Institute of Ecol-ogy, University of Georgia, Athens.

2 A portion of a dissertation submitted to the graduatefaculty of the University of Georgia in partial fulfillmen tof the requirements for the degree of Doctor of Philosophyin the Department of Zoology.

;! Present address: Institute for Medical Research, Uni-versity of California, ICMR, Kuala Lumpur, Malaysia.

recovered from the small intestines of 8of 12 Rana pipiens Schreber. Worms wererecovered from frogs collected at Turrialba,approximately 20 miles from the type locality,Coris, Cartago Province, Costa Rica. Study ofwhole mounted and serially sectioned wormsprovided additional data to those of BrenesMadrigal et al. (1959).

Specimens of G. facioi were compared toother glypthelminth trematodes (sensu Byrdand Maples, 1963) collected by the author in

Copyright © 2011, The Helminthological Society of Washington

OF WASHINGTON, VOLUME 43, NUMBER 2, JULY 1976 117

the United States, Costa Rica and Venezuela orborrowed from the U.S. National Parasite Col-lection and collections of other workers. Thesecomparative studies focused attention on thetaxonomic controversy surrounding this groupof widely distributed plagiorchioid trematodes,parasitic in amphibians and reptiles. This con-troversy is evidenced by the inculsion of thegroup's 28 species in either one genus (e.g.,Nasir, 1966), or, at one time or another, in asmany as six genera (e.g., Cheng, 1959; Byrdand Maples, 1963). In view of the previ-ously suggested relationships of these 28 speciesto the genus Glypthelmins Stafford, 1905, indi-cated by vise of the neutral term glypthelminth,the status and familial assignment of the genusGlypthelmins is reexamined to provide a basisfor assessing the relationships of these 28species.

Trematodes were heat-killed in 0.7 per centsaline under slight coverslip pressure, fixed inalcohol-formalin-acetic acid solution (AFA),stained with Harris hematoxylin, and mountedin Canada balsam or Permount. Specimens ofG. facioi sectioned at 8 ^ had been previouslywhole mounted. Figures were drawn with theaid of a Wild drawing tube; unless otherwiseindicated, all measurements are given inmicrons with the mean in parentheses. Indi-vidual measurements of the G. facioi and G.quieta referred to in this study are availableelsewhere (Sullivan, 1972).

Literature citations in Tables 1-3 are listedas they appear in the Index Catalogue of Med-ical and Veterinary Zoology, U.S. GovernmentPrinting Office, Washington, D.C.

Glypthelmins Stafford, 1905, Char. Emend.

SYNONYM: Margeana Cort, 19.19.DIAGNOSIS: Macroderoididae. Body elon-

gate, cylindrical to subcylindrical. Tegumentspined or scaled. Oral sucker subterminal.Aoetabulum medial, in anterior half of body.Pharynx well-developed. Esophagus present.Cecal bifurcation midway between pharynxand acetabulum. Ceca long, reaching to orinto posterior quarter of body. Testes post-acetabular, symmetrical or diagonal in position.Cirrus pouch elongate, usually overlapped byacetabulum. Ovary pretesticular, in close prox-imity to acetabulum. Seminal receptacle andLaurer's canal present. Uterus transverselycoiled, intercecal, reaching to posterior extrem-

ity; uterine coils not developed in pretesticularregion. Matraterm non-muscular, surroundedby numerous gland cells. Genital pore immedi-ately preacetabular. Vitellaria follicular inlateral fields of body, overlapping ceca dorsallyand ventrally, occasionally confluent dorsally;longitudinal extent highly variable, extendingfrom level of oral sucker to posttesticularregion. Excretory vesicle I-shaped. Parasitic inintestine of anurans.

TYPE SPECIES: Glypthelmins quieta (Staf-ford, 1900) Stafford, 1905.

Glypthelmins quieta

(Stafford, 1900) Stafford, 1905

SYNONYMS: Distomum quietum Stafford,1900; Margeana californiensis Cort, 1919;Glypthelmins californiensis (Cort, 1919) Miller,1930; Glypthelmins subtropica Harwood, 1932.

DESCRIPTION: See Miller (1930).HOSTS AND LOCALITIES: Table 1.SPECIMENS: USNM Helm. Coll. No. 72268-

72271. Other specimens in the author's col-lection.

Glypthelmins rugocaudata(Yoshida, 1916) Yahata, 1934

SYNONYM: Enodiotrema rugocaudatiimYoshida, 1916.

DESCRIPTION: See Yoshida (1916) andYahata (1934).

HOSTS AND LOCALITIES: Rana nigromaculataHollowell, Osaka, Japan (Yoshida, 1916) andP'yongyang (Heijo), North Korea (Ogata,1937); R. rugosa Schlegel, Hiroshima (Yahata,1934) and Kyoto, Japan (Yamaguti, 1936).

SPECIMENS: No specimens of this specieswere available for study.

Glypthelmins staffordi Tubangui, 1928

DESCRIPTION: See Fischthal and Kuntz(1967).

HOSTS AND LOCALITIES: Table 2.SPECIMENS: Four specimens (USNM Helm.

Coll. No. 61702) were studied. (Approximately200 amphibia, including Bufo melanostictus,Rana cancrivora, R. erythraea, R. limnocharisand R. macrodon, from various localities inMalaysia as well as certain of Yuen's (1962)Singapore localities were examined for G. staf-fordi during the last six months of 1974. To



Table 1. Glypthelmins quieta: Hosts and localities.|F. A. Christian; **new locality records.)

(Specimens from the collections of *E. E. Byrd and

Host

BufonidaeBufo americanus Holbrook

HylidaeAcris crepitans BairdHijla crucifcr Weid

( — II. pickcringii Kennicott )

Pseudacris nigrita (LeConte)P. triseriata Weid

Locality

Presque Isle, Maine

IowaCanadaWestern MassachusettsAthens, Clarke Co., Ga.Athens, Clarke Co., Ga.Iowa

Author

Bouchard (1951)

Ulmer (1970)Stafford (1905)Hankin (1945)Byrd & Maples (1963)Byrd & Maples (1963)Ulmer (1970)

RanidaeRana aurora (Baird & Girarcl)

R. boylii (Baird)

R. catesbeiana Shaw

R. clamitans Latreille

R. montezumae BairdR. palustris LeConteR. pipiens Schreber

(= R. cirescen-s Garman)

R. septentrionalis BairdR. sphenocephala Cope

San Francisco, Ca.San Diego Co., Ca.Butte Co., Ca.Marin & Sonoma Co., Ca.CanadaUrbana, Illinoi sHouston & Huntsville, TexasCleveland Co., OklahomaBeaufort Co., N.C.FloridaLouisianaAthens, Clarke Co., Ga.Gaspe Peninsula, CanadaAmherst, MassachusettsNorth CarolinaSeattle, WashingtonHavana, CubaIowa

*Burke, Chatham &Taliaferro Co., Ga.

*Oconee & Screven Co., Ga.*Terrebonne & East Baton

Parishes, La.*Oktibbeha Co., Miss.Western MassachusettsPresque Isle, Maine

*DeKalk & Oglethorpe Co., Ga.* Warren Co., New JerseyMexico, D.F.Presque Isle, MaineCanadaMexico, D.F.

* Franklin Co., Ohio*Alamance Co., N.C.* Franklin Co., Tenii.Presque Isle, MaineCleveland Co., OklahomaHouston & Huntsville, Texas

Cort (1919)Ingles (1936)Ingles (1936)Lehmann (1960)Stafford (1905)Mille r (1930)Harwood (1932)Trowbridge & Hefley (1934)Brandt (1936)Manter (1938)Bennett (1938)Parker (1941)Rankin (1944)Rankin (1944)Rankin (1944)Rankin (1944)Odening (1968)Ulmer (1970)

:::Present study

Present studyfPresent stvidy

'•'Present studyRankin (1945)Bouchard (1951)Present studyPresent studyCaballero y C. & Sokoloff (1934)Bouchard (1951)Stafford (1905)Caballero y C. & Sokoloff (1934)

•{•Present study'^Present studyPresent studyBouchard (1951)Trowbridge & Hefley (1934)Harwoocl (1932)

Table 2. Glypthelmins staffordi: Hosts and localities.

Host Locality Author

BufonidaeBufo melanostictus Schneider

RanidaeOoeidozijgd lima (Gravenhorst)Rana cancrivora GravenhorstR. crythraea SchlegelR. Hmnocharis vittigera Weigmann

(= R. vittigera Weigmann)R. macrodon Dumeril & BibronR. tigrimi rugulosa Weigmann

( = R. rugulosa Weigmann)

Taihoku, Formosa

Hanoi Prov., North VietnamRepublic of SingaporeRepublic of SingaporeLaguna Prov., Luzon, Philippine Is.Manila, Lu/.on, Philippine Is.Republic of SingaporeAmoy and Canton, ChinaHanoi Prov., North Vietnam

Yamaguti & Mitunajia (1943)

Odening (1968)Yuen (1962)Yuen (1962)Tubangui (1928)Fischthal & Kuntx (1967)Yuen (1962)Li (1937a)Odening (1968)



Table 3. Comparison of body and esophageal lengths (in microns) for Glypthelmins staffordi and G.rugocaudata.

Body

Glypthclminx staffordi Glypthelmins rugocaudata

Esophagus Author Body Esophagus Author

2090-4150

1940-4960

1454-1960

40

38-210

10-85

Tuhangui (1928)

Yuen (1962)

Fischthal & Kuntz (1967)

Up to 3200 160-300 Yoshida (1916)

date, the author has not found this species inPeninsular Malaysia or Singapore.)

Glypthelmins shastai Ingles, 1936

DESCRIPTION: See Ingles (1936). Certainmeasurements of the type specimen of G.shastai (USNM Helm. Coll. No. 8925) werenot in accord with the size ranges presentedby Ingles (1936). Since other aspects of thetype's morphology agreed with the originaldescription, only the measurements of the type,figured by Ingles (his pi. 16, fig. 1), are in-cluded here.

Body 3,200 long by 810 wide. Oral sucker210 by 270. Acetabulum 160 by 160. Pharynx110 by 140. Esophagus 300 long. Ovary 210by 210. Anterior testis 320 by 270. Posteriortestis 340 by 320. Cirrus pouch 570 by 120.

HOST AND LOCALITY : Bufo boreas Baird andGirard, Shasta County, California (Ingles,1936).

SPECIMENS: Only the type specimen wasavailable for study.

Glypthelmins facioi

Brenes Madrigal, ArroyoSancho, Jimenez-Quiros, and

Delgado Flores, 1959(Figs. 1-4)

DESCRIPTION (measurements based on 29specimens): Body elongate, 1,820-3,440(2,800) long by 390-900 (640) wide. Tegu-ment entirely scaled, with scales diminishingin number posteriorly. Oral sucker subterminal,170-240 (200) by 170-250 (220). Acetab-ulum medial, in middle third of body, 90—140(120) by 80-150 (120). Ratio of oral suckerto acetabulum 1:0.58 ± 0.03. Prepharynxshort. Pharynx muscular, 70-110 (90) by 110-160 (130). Esophagus 110-280 long. Cecaterminating near posterior extremity. Testes

smooth, diagonal, in mid-region of body; ante-rior testis dextral, 130-240 (200) by 100-230(170), in immediate postovarian zone; poste-rior testis 130-290 (240) by 110-250 (190),with its anterior border extending into the zoneof the anterior testis. Cirrus pouch 150-340(270) by 50-100 (70), extending from pre-acetabular region to ovarian level, containingsaccular seminal vesicle; cirrus eversible, un-armed. Ovary sinistral, pretesticular, spherical,overlapped by acetabulum or not, 100—240(170) by 90-210 (160). Laurer's canal andseminal receptacle present. Uterus intercecal,with numerous transverse coils reaching toposterior extremity of body; pretesticular coil-ing absent. Metraterm as long as or slightlylonger than cirrus pouch. Metraterm and cirruspouch opening separately into a small genitalatrium; genital pore median, immediately pre-acetabular. Vitellaria follicular, commencingat level of cecal bifurcation and terminating ata level behind posterior testis; follicles distrib-uted laterally, overlapping ceca dorsally andventrally, and often confluent dorsally at vary-ing levels throughout their extent. Eggs opercu-late, 27-32 (29) by 12-20 (15). Excretorybladder I-shaped, reaching to level slightly be-hind posterior testis. Excretory system meso-stomate; common collecting tubules receivinganterior and posterior main collecting tubulesin immediate postovarian region.

HOST: Rana pipiens Schreber.SITE OF INFECTION: Small intestine.LOCALITIES: Coris and Turrialba, Cartago

Province, Costa Rica.SPECIMENS: USNM Helm. Coll. No. 72275.

Other specimens in the author's collection.

DiscussionOf the 28 described species of glypthel-

minths, the form of the adult excretory vesicleis unrecorded for Glypthelmins facioi, G.



Figures 1—4. Glypthelmins facioi from Turrialba , Costa Rica (scales in millimeters). 1. Ventral view;note uterine configuration. 2. Ventral view showing form of the excretory bladder. 3. Section throughthe genital atrium; note unarmed cirru s and three adjacent tegumental scales. 4. Section showing dor-sally confluent vitellaria .

palmipedis, G. parva, G. proximus, G.pseudium, G. shastai, G. simulans, G. vitellino-philum, and Choledocystus intermedius. Lif ehistories are known only for G. quieta (Rankin,1944; Leigh, 1946), G. hyloreus (Martin,1969) and C. pennsylvaniensis (Sullivan andByrd, 1970), all three of which are charac-terized by I-shaped bladders in both cercarialand adult stages. Y-shaped bladders arerecorded for G. repandum and G. linguatula(Travassos, 1924), G. subtropica (Harwood,1932), G. africana (Beverly-Burton, 1963), G.

incuroatum and G. ramitesticularis (Nasir,1966), and Rauschiella tineri (Babero, 1951).The excretory vesicle of G. staff or di is variouslydescribed as Y-shaped (Tubangui, 1928; Yuen,1962), as a central canal with two large col-lecting trunks (Li , 1937), or as I-shaped(Odening, 1968). The bladder of G. rugo-caudata is described as extending as far forwardas the testes and dividing into two short lateralbranches (Yoshida, 1916), as Y-shaped (Yahata,1934), or as club-shaped (Yamaguti, 1936).Similarly, Travassos (1926) described the blad-



der of G. elegans, later designated the seniorsynonym of both Choledocystus eucharis andC. vesicalis by Ruiz (1949), as Y-shaped. Thebladders of C. eucharis and vesicalis were de-scribed, respectively, as tubular (Pereira andCuocolo, 1941) and as Y-shaped with shortarms and a long trunk (Ruiz and Leao, 1942).However, Ruiz (1949) considered the bladderin elegans as transitional between the I- andY-shaped forms. Finally, Cordero's (1944)descriptions of G. sera and G. festina, while notstating the bladder forms, suggested that theyare Y-shaped.

Comparison of the shape of the excretorybladder in the glypthelminth trematodes (sensulato) indicates that they are divisible into twogroups: 1) those possessing Y-shaped, and 2)those possessing I-shaped bladders. SouthAmerican forms, represented by 15 "species"(two of which were also reported from CentralAmerica and one only from Mexico), can beincluded in the first group. North Americanforms, as well as one Central American form,and those reported from Southeast Asia, Japanand Korea, can be included in the secondgroup. Although exhibiting a Y-shaped blad-der, the sole representative of the group de-scribed from Africa, Reynoldstrema africanaCheng, 1959 (= Glypthelmins a. Dollfus,1950), demonstrated certain characters whichsuggest its removal from consideration with thisgroup (see below).

While placement of the glypthelminths in thesuperfamily Plagiorchioidea Dollfus, 1930, isaccepted (Schell, 1962; Byrd and Maples,1963; Odening, 1964; Sullivan and Byrd,1970), various members of the group, or thegroup as a whole, have been included in theMacroderoididae McMullen, 1937 (Schell,1962; Odening, 1964; Sullivan and Byrd, 1970)or in the Plagiorchiidae Luhe, 1901 (Fischthaland Kuntz, 1967; Martin, 1969; Ulmer, 1970).However, in establishing the Macroderoididae,McMullen (1937) separated it from the Plagior-chiidae primarily on the basis of the I-shapedexcretory vesicle of the former as opposed tothe Y-shaped vesicle of the latter. Since theglypthelminths are divisible by the I- or Y-shapeof the bladder and exhibit other morphologicalcharacters of the Plagiorchiidae and theMacroderoididae, as emended by Odening(1964) and Sullivan and Byrd (1970), respec-

tively, it is proposed that those glypthelminthswith a Y-shaped bladder be restricted to thePlagiorchiidae, whereas those with an I-shapedbladder be referred to the Macroderoididae.The taxonomy of those glypthelminths pos-sessing a Y-shaped bladder as well as that ofCholedocijstus pennsylvaniensis Byrd andMaples, 1963 (= Glypthelmins p. Cheng,1961) and Glypthelmins hyloreus Martin, 1969,wil l be considered elsewhere.

Citing Manter's (1969) statement, "To dis-tinguish between a 'tubular excretory vesicle'and a Y-shaped vesicle is not realistic. Al lexcretory vesicles are tubular, whether I-shaped, Y-shaped, or V-shaped," Nasir andDiaz (1970) emended the diagnosis of theMacroderoididae to include forms possessing atubular bladder rather than only an I-shapedbladder as originally proposed by McMullen(1937). Considering the morphological simi-larities of the adult worms assigned to the twofamilies, such an all inclusive treatment ofexcretory vesicles could render the familyMacroderoididae a synonym of the Plagior-chiidae, if the intramolluscan stages were notknown (Yamaguti, 1971). Although Manter'sstatement regarding separation of tubular andY-shaped bladders is well founded, the basicI-, Y-, or V-shape of the bladder is, as Mantersuggests, independent of its tubular nature.

The author accepts the conclusions of Miller(1930) in considering Marge ana Cort, 1919, asynonym of Glypthelmins, and also agrees withByrd and Maples (1963) in suppressing thereinstatement of Margeana by Cheng (1959).

Although Miller (1930) differentiated G.californiensis from G. quieta by the anteriorextent of the vitellaria in G. californiensis,Caballero y C. and Sokoloff (1934) noted thatthe vitellaria in their G. californiensis did notconform with Cort's (1919) account and attrib-uted this difference to the effects of flattening.The variability in the position of the vitellaria inspecies of Glypthelmins, which was emphasizedby Miller (1930), as well as the findings ofCaballero y C. and Sokoloff (1934) with regardto G. californiensis, leads to the conclusion thatvitelline position is insufficient for the sepa-ration of G. californiensis from G. quieta. Nasirand Diaz (1970) considered these two speciesconspecific, a conclusion which is supported bythe author.



In separating G. subtropica from G. quieta,Harwood (1932) maintained that G. subtropica

. . . most closely resembles G. quieta, but itmay be ... distinguished from this form bythe transverse band of vitellaria, and thelocation of the testes behind the transversevitelline duct, and the tendency of the uterusto pass ventral to the testes rather thanbetween them.

Olsen (1937), in bis key to the species ofGlypthelmins, further distinguished G. sub-tropica from G. quieta by indicating that theformer possessed a pharynx larger than theventral sucker.

Concurring with Miller's (1930) findingsregarding morphological variability in G. quieta,Rankin (1944) also figured nine specimens(his figs. 11-19) of G. quieta from asingle natural infection and called attentionto the vitelline distribution; three speci-mens (his figs. 14, 17, 19) showed markedtransverse bands of vitellaria, obviating one ofHarwood's (1932) distinctions between G.subtropica and G. quieta. Further, Harwood'suse of the relative position of the testes andtransverse vitelline ducts is considered highlyquestionable in view of 1) Stafford's (1900)inability to accurately demonstrate the ductsthemselves or their position, and 2) Rankin's(1944) findings which indicate that the relativepositions of the ducts and testes depend to somedegree on the oblique or symmetrical positionof the testes. Ventral overlap of the testes bythe ascending uterus is constant although thedegree of overlap is highly variable. No taxo-nomic significance, therefore, is attached to a"tendency" of the uterus to pass ventral to thetestes rather than between them. Preliminarystudy of size allometry, based on measurementsof 74 G. quieta, indicates that although pharyn-geal size is greater than that of the acetabulumin smaller specimens, this size relationship isreversed in larger specimens (Sullivan, 1972).Therefore, the ratio of pharyngeal size toacetabular size cannot be considered a validtaxonomic character as proposed by Olsen(1937) for the separation of G. subtropica andG. quieta. Since the characters used by Har-wood (1932) and Olsen (1937) are insuffi-cient for the separation of these two species andno characters could be found to distinguish

them from each other, it is suggested that G.subtropica be considered a synonym of G.quieta as proposed by Manter (1938).

Tubangui (1928) distinguished G. staffordifrom G. quieta by vitelline extent, arrangementof testes and ovary, and egg size. Yahata(1934), in transferring Enodiotrema rugo-caudatum to Glypthelmins, separated thisspecies from G. quieta by the relative positionsof the ovary and ventral sucker, as well as theposition of the genital pore and that of thetestes. The author's inability to obtain sufficientspecimens of G. staffordi or G. rugocaudataprecludes any assessment of morphologicalcharacters which could be used to differentiatethese two forms from each other or from G.quieta, G. shastai or G. facioi. It should benoted that the literature concerning the glypt-helminths contains no statements which purportto distinguish G. staffordi from G. rugocaudata.Examination of pertinent figures and descrip-tions of G. rugocaudata (Yosbida, 1916; Yahata,1934) and G. staffordi (Tubangui, 1928; Yuen,1962; Fischthal and Kuntz, 1967; Odening,1968) and examination of four specimens ofG. staffordi (USNM Helm. Coll. No. 61702)indicated that the only discernible differencebetween the two species was the relative lengthof the esophagus (Table 3). Although not giv-ing measurements, Yahata (1934) figuredspecimens of G. rugocaudata which show arelatively longer esophagus than that of G.staffordi. In view of the lack of supportiveevidence to the contrary and the wide geo-graphic separation of these two species fromthe North and Central American species, bothG. staffordi and G. rugocaudata are tentativelyaccepted as valid species. Of Parentheticinterest in this regard is the report of a Glypt-helmins sp. from Rana temporaria in easternSiberia (Milogradova and Spasskii, 1957)which extends the northern range of the genusin Asia. Further study of this northern formcould provide data for evaluating both thesystematics of the Asian species of Glypthelminsand their relation to the North and CentralAmerican forms.

Nasir and Diaz (1970) considered G. shastaia synonym of G. linguatula, a species trans-ferred to Choledocijstus by Byrd and Maples(1963) and the position with which the author



is in accord. It is difficul t to determine on whatbasis the synonymy was advanced since Nasirand Diaz' key was constructed around thoseforms with a known cercaria and those forwhich the cercaria was unknown. Study of theholotype of G. shastai indicates that the specieswas described from immature forms, judging bythe development of the uterus. However, thepresence of G. shastai in a bufonid host as wellas the large testes suggests that this speciesmay be more closely allied to Choledocystusthan is presently suspected. Since the form ofthe excretory bladder is unknown, G. shastai isretained in Glypthelmins pending furtherstudy. As such, it can be distinguished fromG. facioi by the presence of tegumental spinesrather than scales and by the relatively greatersize of the testes, a character together with eggsize which distinguishes it from G. quieta.

Glypthelmins facioi can be distinguishedfrom G. quieta by the presence of scales (Fig.3) as well as by egg size. It should be noted,however, that although Brenes Madrigal et al.(1959) reported that G. facioi possessed eggsmeasuring 33-47 by 20-21, eggs of specimenscollected at Turrialba measured 27-32 by12-20.

Cheng's (1959) establishment of Reynold-strema for Glypthelmins africana has much tosupport it; however, its relationship to the otherglypthelminths is questionable. He maintainedthat "The posteriorly located uterus and theposteriorly situated testes are sufficient criteriato justify the erection of ... Reynoldstrema."Cheng regarded this genus more closely alliedto Brachycoelium (Dujardin, 1845), Glypt-helmins and Marge ana than to AstiotremaLooss, 1900, since Reynoldstrema lacked aprepharynx, long Laurer's canal, and an excre-tory bladder reaching the seminal receptacle,characteristics of Astiotrema as outlined byOlsen (1937). However, the prime genericcharacters of Reynoldstrema strongly suggesta closer relationship to Astiotrema than thatadvanced by Cheng (1959).

While Beverly-Burton (1963) noted thatDollfus (1950) included Glypthelmins in theBrachycoeliidae Johnston, 1912, on the basis ofthe I-shaped excretory bladder, she agreed withYamaguti (1958) and Skrjabin and Antipin(1958) in assigning Glypthelmins to thePlagiorchiidae, after she demonstrated a Y-

shaped bladder in G. africana, recovered fromRana adspersa (Tschudi) and Mabuya striata(Peters) in Southern Rhodesia. She furthermaintained that Plagiorchis himalayi (Jordan,1930), P. momplei Dollfus, 1932, and P.ramlianus (Looss, 1896) were more similar toG. africana "than . . . the accepted species ofGlypthelmins . . . ," in that the uterus filled thearea posterior to the ceca and vitellaria. Con-sequently, she suggested that a separate genusmight be erected for some of these forms.

In reviewing Reynoldstrema, Fischthal andThomas (1968), after considering the sug-gestions of Beverly-Burton (1963), indicatedthat Cheng (1959) had previously establisheda new genus for G. africana. These authorsdisagreed with the placement of G. africana inPlagiorchis Luhe, 1899, by Vercammen-Grandjean (1960) or Plaplometra Looss, 1899,by Manter and Pritchard (1964). Fischthaland Thomas maintained the validity ofReynoldstrema to which they transferredPlagiorchis laurenti Vercammen-Grandjean,I960, and P. berghei Vercammen-Grandjean,1960.

In view of the generic revision of Plagiorchisby Odening (1959) and the findings ofBeverly-Burton (1963) and Fischthal andThomas (1968), the author does not acceptthe synonymy of Reynoldstrema with Plagior-chis proposed by Yamaguti (1971). The Y-shaped excretory vesicle exhibited by species ofReynoldstrema indicates placement in thePlagiorchiidae rather than the Macroderoididaeas proposed by Odening (1964). Additionally,the morphological characters of Reynoldstremaare sufficient to allocate this genus to the sub-family Astiotrematinae Baer, 1924.

Skrjabin and Antipin (1958) recorded a V-shaped excretory bladder for Glypthelminsdiana Belouss, 1959. The presence of a V--shaped bladder in this species precludes itsplacement in either the Plagiorchiidae or theMacroderoididae. Accordingly, G. diana isremoved from Glypthelmins, which is charac-terized by an I-shaped bladder. However, theV-shaped bladder of this species appears toindicate a lecithodendriid relationship, but sincethe species does not conform to any of thepresently known lecithodendriid genera, itseems advisable to designate the speciesincertae sedis pending further investigation.



Acknowledgments

I am grateful to the late Dr. Elon E. Byrd,Department of Zoology, University of Georgia,for his comments and suggestions during thestudy, and to Drs. F. A. Christian, BiologyDepartment, Southern University, M. J.Ulmer, Department of Zoology, Iowa StateUniversity, G. W. Martin, Department ofZoology, University of Montana, for loaning ordonating specimens and J. R. Lichtenfels forloaning specimens from the USNM Helmin-thological Collection. I am indebted to Mr.Arnold Ericson of the IICA, Turrialba, CostaRica for his cooperation. I would also like tothank Drs. C. P. Goodyear, Tenn. Coop. Fish-eries Unit, Tenn. Tech. University, Ow-YangChee Kong, Institute for Medical Research,Kuala Lumpur, Lim Chuan Fong, Departmentof Zoology, University of Singapore, andMessrs. Mohamed Yusof bin Mohd. Idris,Chong Kon Chu, and Ching Kee Chai of theUC ICMR staff for help in collecting am-phibians, and Dr. David Stiller, UC ICMR, forreading the manuscript and Mrs. Margaret Limfor typing. I would like to thank the Directorand staff of the Institute for Medical Research,Kuala Lumpur, for their help and cooperation.

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the trematode genus glypthelmins stafford, 1905...

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