Proceedings of the Helminthological Society of Washington 35(2) 1968

Volume 35 July 1968 Number '2

PROCEEDINGS

• \'-> .";.<•••"•:.. -r? x_/-,' .-•"•' •-•'.• ':'•,- •• '• ' ?~~ ,••"' ] ;.••'•':••'•.-- •.'.-.•- " ... . • ': ',"- The Helminthologicai Sodety

A semiannual journal of research devoted to Helminfho/ogy and a// branches 6f Parasitology

•^% SuppOrtedun pdrt byithe >'.^.^'f^ Braytqn H. RqnsonT/Aenr^prial Trust Fund

Substriptipn $7.00 a Volume; Foreign, $7^50

A.CHQLONU, ALEXANDER D. Studies on the Freshwater Cercariae of Northern' • •. -/•' " :'" * ^Colorado .T...-7..._-..;...-:...^.:-:,Ji— ____ .^..,:.,. _____ ,.L^.r,r.:,.-j'.'l,^..r. --:... .;•--- J?.:. .^. ; .^259

DASGUPTA, :D. 'R.'j D. J. RASKI, AND Sr A. SHER. A Revision of the (Genus ; •,' fi£[4 Oliveira, 1940 (Nernatoda: Tylenqhidae ) ..^-.' Ii69 ., AND jflAROLD W. MANTER. Some .Digenetie Trernatodes / J of Marine Fishes of. New Caledonia.- '!Part , I. Rucephalidae, ^Monorchiidae, !> V arid Some Smaller 'Families ./l.^-.-l.---^-.^.-...!!!---^..----.-!------___ _•-„ ______: _ 143

;i:r ! THE HELMINTHOLOGICAl SOCIETY OF WASHINGTON / ;-f^ &\j ' '''". -V- "••••. ''•'.'•/"'• W? ' -.C:, A1' " /".' J. Wj " ; •-,_•'"-. £ 'w' ' ; !fHE, SOCIETY /meets once ' a rponth from October ', through' M^y for ; the presentation and discu&iOn of .papers in any and all /branches; of parasitology or related Sciences. All interested ^persons -are invited to attend. j«y '•/ -•• .' .-'"., • : . '; ' -' ' /'';.•' •; ' r '')'/• ",•;- " '.,'. . i ' •.-.': i ' '•.',-':",• ' '•'* ' • .. N • **) - <' '* *T •'=",' 1 '.-•"-.. j •'("•" '- -.•''.,•':- ''Persons interested in -membership -in- the1 Helminthologjcal ^ociety^of Washington ,may obtain .application blanks xfrom .the ;Correspohdirig SeeretaryrTreasurer, '.Miss'1' Edna Jvl. 'Buhrer, Belts- Ville '/iPardsitplogical'. laboratory, .•Agricultural ^Research Center, Beljtsville, Maryland 20705; A; year's subscript|ori to the Proceedings is included in the annual dues ($6.00). ; ,,"..' : ' ; '

^f-'y- ;^;;A';P;'. ; OFFICERS xp THE SOCIETY ifdR i96a

Presfdenfe tJAVID R; LINCICQME /" '''-U'^ ^.•'''i?/: f/0 fe' Vice President: ALAN^C PIPKIN, v, : ' ;" ' '''^ -^'W'^ •'-?•' '!' -V Correspb^Av^g Secretary-Treasurer: ^EDNA M, BUHRER--:/ -.•.; ,'-| x< l^ ^ Assisfcni •Corr^p'ndtn^ Sec^^ •)• Recording Secretary: -E..-.J.. \L. SOUI^By ,-' . '. ;; wV^^ '' 7 : ,^',-:'/:, :^4

/^ECl^U^]&l-|i^ .jr ^^^ ;; ^;;'^^'.'':';:" -rv *' ; Representative to the WasMngton ^cqde^^^ r t i v^, Representative to !the American 'Society pi Parasitologists: CHARLES G. D^JRfelN' •<• 1965-f 7). /Executive, Committee Hembers-at^Large:' ' -GILBERT F. OTTO, ,1968 '}• •> : x' J " ''•';

. .,. , , PROCEEDINGS OF THE^ ÊIMINTHOLOGKAL SOCIETY C*FvWASHINGTO MyCPBDIMGS are published by the ee^minthological j Society o^ Wasi>iHgtorif JHp\vever» ipapers' heed not be /preserited at ,a meeting to ^be published ; in the 'Proceedings. NonTmembers nây publish in the 'Proceedings 'if they vVvill contribute the ;fvill cost |of /publication. Two issues /are .published each '< year; in ^January arid July. -v . / '. -. • ; ', • •.'••,*'• ' '' •'•••'>, :jv ',. ••'•/V1 i .'''':;'h'. •'; •'] '•/'• 7-' •'.;,•',.• ",-•'•.••..'' /-. .;,-'••" .^ /•• '- 'MAiyC/SCRJPTS ,may be subniitted ; to any member of the Editorial iComrnittee. Manu- scripts niust be._ jtypewritten, dpuble -spaced, ;and 'in: .finished; fptmi 'Only the ribbon cppy >will be accepted for publication; it is accepted iWitli the understaitiding that »it will be published ,ohly in the ProceedingsT • / ,• "'•''. ;'' ' ';'• •,-• -t\ ' '•.;'„'• '' . ' • ., • .'^ : .,'".,7 ' ," ^ :• 'REPRINTS may be-wdered ;from the "PRINTER at theAârne^ time 'the 'corrected /proof is returned tOj the EDIITOR. y :l /"!, •''''•.''' ,•• "/'•';'' •/• ' .-.,. • '•'. " ,;/'""' "'•:}/ -'''"''-,;, -e': .'- !-: '•':»•''• BACK VOLUMES, ,6f the Proceedings are available. f!nQViries cpricerning back volumes and current subscriptions^ should be directed toi jMiss Edria M. • Buhrer^ Corresppnding Secretary- Treasurer. !.;, ; "'- . . ^'"'~. >-/••' ' •!.'',-'' . ' '"! '" '! • '-'•• •• • ,!'r-''f' '\'-'J .• .•'!-• EDiTORIAL> CpJVlMlTTEE "'' ''' »;-"•':: ' -VN/' J. TROMB A, Editor, 1970 Beltsville Parasitoljbgical Laboratory " ! AgriculturaMleseareh Center'

'JACOB'H, 'FISCHTHALH972 y" ;//.'. i R. NICKLE^ 1972;'

i HARGIS?JR;,'i971 ^ '^

; :Ll|CKER, ;1Q72 '-; t ' =,; fcA ' , 197Q - :. , 1969

VOLUME 35 JULY 1968 NUMBER 2

Thiabendazole as an Anthelmintic Against Ascaridia columbae in Pigeons1

EVERETT E. WEHR2 AND MERLE L. COLGLAZIER

Ascaridia columbae is a common intestinal The worms passed by each bird were nematode of pigeons. When present in large counted daily. At the termination of treat- numbers, the parasite causes anemia and weak- ment, the pigeons were necropsied and all ness. Often the affected birds are poor eaters worms remaining were counted. and breeders (Levi, 1957). Attempts to control this parasite with piperazine were unsuc- Results and Discussion cessful (Hwang et al., 1958) despite efficacy The data are in Table 1. One hundred per of the drug against A. galli of chickens (Col- cent of the A. columbae were removed by glazier et al., 1960; Horton-Smith and Long, feeding mash medicated with 0.5% thiabenda- 1956; and Shumard and Eveleth, 1955). Inas- zole for a 10-day period. Failure of the drug much as thiabendazole has demonstrated effec- to remove all worms from pigeons treated for tiveness against many intestinal parasites, in- 8 and 9 days reduced its overall efficacy cluding ascarids of livestock and poultry to 96%. This reduced efficiency may be mis- (Brown, 1961; Cuckler, 1961; and Long and leading because 47 of the 62 worms recovered Wakelin, 1964) it was tried against A. colum- in Trial 2 came from one pigeon; this bird bae. The results are reported herein. failed to consume its allotment of medicated mash. Materials and Methods The drug removed immature as well as ma- The 57 White King pigeons used in this ture worms. The greatest number of worms experiment were purchased from a commercial was passed on the 3rd post-treatment day, but grower as 6-week-olcl squabs. They were housed in sterilized wire cages suspended appreciable numbers were still being passed on from the ceiling of the poultry house, and the 6th day. Unfavorable effects were not fed unmedicated turkey-pigeon mash. Fecal observed in any of the birds. examinations were made for a 2-week period to ascertain that the pigeons were free from Summary and Conclusions extraneous parasites. Each bird was then Fifty-seven 15-week-old White King pigeons, inoculated experimentally with approximately experimentally infected with the intestinal 1,000 infective eggs of A. columbae from lab- roundworm, Ascaridia columbae, were used in oratory cultures. four critical trials to assess the anthelmmtic About 50 days later, when the droppings contained eggs of A. columbae, each bird was activity of a 0.5% thiabendazole-medicated put in a separate cage and given free access mash fed ad libitum. Efficacies of 99, 84, 100, to turkey-pigeon mash containing 0.5% thia- and 100% were obtained in birds fed the mash bendazole by weight, for the periods shown in for 8, 9, 10, and 13 days, respectively. The Table 1. average efficacy in the four trials was 96%. Both immature and mature A. columbae were 1 Beltsville Parasitological Laboratory, Animal Disease and Parasite Research Division, Agricultural Research Service, removed. The drug produced no visible un- Beltsville, Maryland 20705. - Retired November 30, 1965. favorable effects. 117

Table 1. Efficacy of 0.5% thiabendazole-medicated mash for the removal of Ascaridia columbae from pigeons.

Parasites Number Treatment of period No. removed No. recovered Efficacy Trial pigeons (days) at necropsy ( per cent ) 1 19 8 669 8* 99 2 15 9 318 62f 84 3 8 10 258 0 100 4 15 13 282 0 100

Total or average 57 1,527 70 96 Worms recovered from one bird only. Worms recovered from five birds (47 from one bird).

Literature Cited Hwang, J. C., D. K. McLoughlin, and E. E. Wehr. 1958. Removal of ascarids from Brown, H. D. 1961. 2-(4'Thiazolyl)-benzimida- zole (thiabendazole)—a new anthelmintic. pigeons. Vet. Med. 53: 263-264. Abstr. of Papers, 140th Mtg. Am. Chem. Soc., Levi, W. M. 1957. The pigeon. Levi Publish- Chicago, Illinois: 28-30. ing Co., Inc., Sumter, South Carolina: pp. Iv Colglazier, M. L., A. O. Foster, F. D. Enzie, and + 667. D. E. Thompson. 1960. The anthelmintic Long, P. L., and D. Wakelin. 1964. The ef- action of phenothiazine and piperazine against fects of thiabendazole upon experimental in- Heterakis gallinae and Ascaridia galli in chick- festations of Ascaridia galli and Capillaria ens. J. Parasit. 46: 267-270. obsignata in the chicken. British Poultry Sci. Cuckler, A. C. 1961. Thiabendazole: a new 5: 187-192. broad spectrum anthelmintic. J. Parasit. 47 Shumard, R. F., and D. F. Eveleth. 1955. A (SuppL): 36-37. Horton-Smith, C., and P. L. Long. 1956. The preliminary report on the anthelmintic action anthelmintic effect of three piperazine deriva- of piperazine citrate on Ascaridia galli and tives on Ascaridia galli (Schrank 1788). Poul- Heterakis gallinae in hens. Vet. Med. 50: try Sci. 35: 606-614. 203-205.

Helminths of Some Wild Mammals in the Southeastern United States1 GROVER C. MILLER AND REINARD HARKEMA Zoology Department, North Carolina State University at Raleigh

During our studies in recent years, some When possible, host animals were obtained local native wild mammals have been exam- alive from trappers during the regular trapping ined for helminthic parasites. Some of these season, but some were shot and others drowned studies have been published including infor- in traps so that immediate examination was mation on helminths of the raccoon (Harkema not always possible. All animals, however, and Miller, 1964) and the mink (Miller and were examined as soon as possible after cap- Harkema, 1964). Information is presented ture. The trematodes, cestodes, and acantho- here on a limited number of four additional cephalans were killed and fixed in AFA (al- species of animals: the otter, Lutra canadensis; cohol, formalin, and acetic acid), stained and bobcat, Lynx rujus; grey fox, Urocyon cine- mounted for further study. The nematodes reoargenteiis; and red fox, Vulpes fulva. were killed in hot 70 per cent alcohol and stored in glycerin alcohol. Large nematodes 1 Supported in part by research grant AI05927 from the National Institutes of Health, U. S. Public Health Service. were cleared in lacto-phenol.

Table 1. Incidence of helminths in 20 otters from the National Museum as USNM Helm. Coll. North Carolina. No. 57415. Species of parasite Number infected Enhydridiplostomum alarioides (Dubois, 1937) was the only other trematode found in Trematoda Btischkirovitrema incrassatum (Diesing, 1850) 14 the otter. It occurred in 7 of 20 hosts and EnhydridvplostomuTn alarioides (Dubois, 1937) 7 varied from 1 to 400 per host. This species Nematoda was reported from the mink, Mustela vison, in Capillaria plica (Rudolphi, 1819) 5 North Carolina by Miller and Harkema (1964). Crenosoma goblei Dougherty, 1945 1 Specimens were collected by Pearson in 1951 Gnathostonia miyazakii Anderson, 1964 4 from the otter in Chapleau, Ontario and catalogued as USNM Helm. Coll. No. 49045. Results and Discussion Permission was obtained from the Museum to stain and mount these for detailed study. Our Twenty-eight speeies of helminths were re- forms agree in all respects. Sawyer (1961) covered from 65 mammals. These are dis- published the first report of this parasite in cussed in relation to the four species of hosts the United States. The type specimen of because most of the parasites encountered EnJujdridiplostomiim fosteri (Mclntosh, 1940) appeared to be fairly host specific. also was compared, and we agree that it is a distinct and separate species. Specimens from Lutra canadensis North Carolina have been deposited as USNM (Table 1) Helm. Coll. No. 57429. Twenty otters were collected from eight Capillaria plica (Rudolphi, 1819) was one of counties in North Carolina in the vicinity of three nematocles collected from the otter. It oc- the Northeast Cape Fear River. About one- curred in 5 of 20 hosts and ranged from 1 to 30 third was from coastal counties, but there per host. The urinary bladders of the raccoon, appeared to be no apparent differences in the bobcat, and foxes also contained this parasite. helminths in hosts from coastal areas versus Crenosoma goblei Dougherty, 1945, was re- those from inland, habitats. Two trematocle covered from the respiratory tract of one host. and three nematode species were collected Only one specimen, a female, was found, and, from the otter. therefore, the specific identification is uncertain. Baschkirovitrema incrassatum (Diesing, 1850) Gnatliostoma miyazakii Anderson, 1964, was was the most common trematocle found in the recovered from the kidneys of four otters. otter. It was present in 14 of the 20 hosts These hosts were from counties adjacent to and varied in number from 1 to 89 per host. the Northeast Cape Fear River. The total The measurements of our specimens compare number of parasites was difficult to ascertain favorably with those of Beverley-Burton (1960). because it was practically impossible to remove This form is found in various species of otters an intact worm from the fibrous tissue of the from several parts of the world including Brazil kidney. Sufficient material was available, (Diesing, 1850), the Sudan (Myers, Wolf- however, to confirm the identity of this species. gang, and Kuntz, 1960), and Southern Rho- The unusual location of these mature gnatho- desia (Beverley-Burton, 1960). More recently stomes invites additional investigation. This is Miller and Harkema (1964) reported the mink apparently the first record of this nematode in in North Carolina as a new host. Specimens the United States. The only other gnatho- from the otter in New York deposited in the stome reported in mustelids in North America National Museum by W. J. Hamilton in 1953 is Gnathostonia sociale (Leidy, 1858). The and. additional specimens from Georgia col- latter species was reclescribed and compared lected by McKeever and Sawyer in 1955 were with G. miyazakii by Anderson (1964). obtained on loan through the courtesy of Dr. Allen Mclntosh and compared with our speci- Lynx rufus mens. All are similar. The only other record (Table 2) of this trematocle in the United States is that Helminths were recovered from 16 bobcats, of Sawyer (1961) from the otter in Georgia. 12 hosts were from counties in eastern North Two of our specimens have been deposited in Carolina and four from two counties in South

Table 2. Incidence of helminths in 16 bobcats was one of four species of cestodes recovered from North Carolina and South Carolina. from the bobcat. It was present in 12 of the Species of parasite Number infected .1.6 hosts from both coastal and inland habitats, though more prevalent in hosts from coastal Trematoda regions. It is interesting that both adults and Alaria canis LaRuc and Fallis, 1936 5 spargana were not found in the same host even Paragonimns rudis (Diesing, 1850) 1 though often present in the raccoon and grey Cestoda fox. It is known that young kittens can be Spirometra mansonoides (Mueller, 1935) 12 infected with spargana by feeding them in- Mesocestoides variabilis Mueller, 1927 1 Taenia macrocystis Diesing, 1850 12 fected copepods. An excellent summary on Taenia rilcyi Loewen,1929 4 the biology of S. mansonoides was recently Acanthocephal a presented by Mueller (1966). Centrorhynchus sp. 1 Mesocestoides variabilis Mueller, 1927, was recovered from only one host taken from Nematoda Fender County, North Carolina. These speci- Toxocaramystax (Zecler, 1800) 12 Ancylostoma caninum (Ercolani, 1859 ) 5 mens were stained and compared with the Ancylostoma braziliense Gomez de Faria, 1910 3 descriptions of Voge (1955). Except for minor Capillaria plica (Rudolphi, 1819) 2 differences they compare favorably. The Anafilaroides rostrata Gerichter, 1949 2 specific identification of members of the genus Molineus barbatus Chandler, 1942 6 Dirofilaria striata (Molin, 1858 ) 2 Mesocestoides is difficult. The genus was reviewed by Voge (1955) who regarded both Mesocestoides manteri Chandler, 1942, and Carolina. Thirteen species of helminths were M. variabilis as synonyms of Mesocestoides recovered including two trematodes, four corti Hoeppli, 1925. cestodes, one acanthocephalan, and seven Taenia macrocystis Diesing, 1850, was one nematodes. of two taenioids found in the bobcat. It was Alaria canis LaRue and Fallis, 1936, was present in 12 of 16 hosts ranging from 2 to 48 recovered from five hosts, all of which were specimens per host. In two hosts from South collected in Brunswick County, North Caro- Carolina and two coastal North Carolina speci- lina. The number of worms varied from 1 to mens there was a mixed infection of T. macro- 159 per host. Pearson (1956) gave a thorough cystis and T. rileiji Loewen, 1929 (see below). description of the life cycle of A. canis and Members of the cestode genus Taenia include also the first report of its presence in bobcats. a number of closely allied species, many of Most of the specimens in the bobcat are which are poorly defined and consequently smaller than the ones recovered from the foxes difficult to identify. Taenia macrocystis was in this area. Otherwise, they are similar. placed in the genus Hijdatigera by Warclle and Paragonimus rudis (Diesing, 1850 (= P. McLeod (1952), Yamaguti (1959), and others. kellicotti Ward, 1908) was recovered from one Taenia is differentiated from Hijdatigera on host collected in Brunswick County, North the basis of a nonsegmented neck region. Carolina. Four mature and two immature There is no segmentation immediately behind worms were found. There are scattered records the suckers on our specimens. Moreover, other of this trematode occurring in other states, but authorities (Cameron, 1926) (Rausch and the only records in North Carolina are Hard- Williamson, 1959) (Esch and Self, 1965) castle (1941), Harkema and Miller (1964), advocate the synonymy of Hijdatigera and and Miller and Harkema (1964) reporting the Multiceps with Taenia. Recent authors have house cat, raccoon, and mink respectively as emphasized the character and size of the hosts. Jordan and Byrd (1958) reported Para- hooks, especially the smaller ones, as being gonimus from the bobcat and cited other the best features for specific identification. references to its presence in swine in Georgia. The study by Riser (1956) on the hooks of Stewart and Jones (1959) also reported it taenioid cestodes from North American felids from the pig in Georgia and used the designa- was the basis for the identification of our tion P. rudis as determined by Allen Mclntosh. specimens. Spirometra mansonoides (Mueller, 1935) Taenia rileiji Loewen, 1929, was recovered

Copyright © 2011, The Helminthological Society of Washington OF WASHINGTON, VOLUME 35, NUMBER 2, JULY 1968 121 from 4 of 16 hosts, varying from 2 to 23 per Table 3. Incidence of helminths in 24 grey foxes host. As stated above they were found in from North Carolina, South Carolina, and Georgia. mixed infections with T. macrocystis. They Species of parasite Number infected are differentiated from the latter on the basis of the larger size, having a more robust body, Trematoda Etirytrema prncyonis Denton, 1942 3 and with a strobilar configuration much like Euryhclmis squamula (Rudolphi, 1819) 1 that of Taenia pisiformis. T. macrocystis is a Sellacotyle mustelae Wallace, 1935 1 rather small, slender member of the genus. Alaria cam's La Rue and Fallis, 1936 1 Procyotrema marsupiformis Harkema and Of significance too is the fact that the large Miller, 1959 1 hooks of T. rileyi are about 0.22-0.24 mm long, Cestoda whereas those of T. macrocystis are 0.32-0.34 Taenia pisiformis (Bloch, 1780) 9 mm in length. Comparison of the large and Mesocestoides variabilis Mueller, 1927 5 small hooks of T. rileiji, T. macrocystis, and Spirometra (?) mansonoides (Mueller, T. pisiformis (the latter collected locally from 1935) spargana 2 Nematoda dogs) indicates that the size and shape of the Physaloptera maxillaris Molin, 1860 9 small hooks provide the best diagnostic fea- Molineus patens (Dujardin, 1845) 2 tures for separating these forms. This is ap- Ancylostoma caninum (Ercolani, 1859) 17 parently the first report of T, macrocystis and Capillaria aerophila (Creplin, 1839) 1 Capillaria plica (Rudolphi, 1819) 5 T. rileyi from the southeastern United States. Trichuris vulpis (Froelich, 1789) 1 One immature specimen of Centrorhynchits Dirofilaria immitis (Lcidy, 1856) 2 sp. was the only acanthocephalan recovered. Members of this genus are commonly found in birds, and it is presumed that this form is recovered from the lungs of two hosts. Klewer an accidental parasite of the bobcat. (1958) reported its presence in the lungs of Toxocara mystax (Zeder, 1800) was the 23 of 24 bobcats in Virginia. He also reported most common nematode in the bobcat and that its life cycle involves the slug Limax was recovered from 12 of 16 hosts and varying maximus as an intermediate host. from 1 to 40 per host. This species is cos- Molineus barbatus Chandler, 1942, was re- mopolitan, having been reported many times covered from 6 of 16 hosts with from 1 to 23 from felids. Much of the biology on this and per host. This form is commonly found in related species has been studied by Sprent raccoons (Harkema and Miller, 1964) and has (1956, 1958). been reported from the skunk (Babero, 1960). Ancylostoma caninum (Ercolani, 1859) was Gupta (1961, 1963) studied the life history present in 5 of 16 hosts with 4 of 5 infected and biology of this species. Schmidt (1965) hosts being from South Carolina. There were recently described a new species from the from 1 to 23 per host with an average of weasel and included a key to the genus. This about 7 indicating that the dog hookworm is is apparently the first report of the bobcat as not a common parasite of the bobcat. a host for this species. Ancylostoma braziliense Gomez de Faria, Dirofilaria striata (Molin, 1858) was re- 1910, was recovered from three hosts collected covered from two South Carolina hosts. One in the vicinity of Dorchester, South Carolina. host contained two worms, the other one nine. The largest number from a single host was Both sexes were recovered from the sub- eight. The cat hookworm appears to be an cutaneous region of the thigh. Orihel and even less of a problem than the dog hookworm. Ash (1964) reported this species in a bobcat Capillaria plica (Rudolphi, 1819) was found from Louisiana. The average length of three in the urinary bladders of only two hosts. Read females was 27.8 cm, of six males, 8.8 cm (1949) stated that capillarids appear to have which compares favorably with the measure- little host specificity though fairly specific for ments given by Orihel and Ash (1964). their location within the host. This seems to be the case with C. plica as we have found it Urocyon cinereoargenteus in the bladders of the raccoon, bobcat, otter, (Table 3) red and grey foxes. The 24 grey foxes were obtained as follows: Anafilaroides rostrata Gerichter, 1949, was 15 from North Carolina, 7 from Georgia, and

2 from South Carolina. The helminths re- and Miller, 1964). Wallace (1934) reported covered from this host appear to be similar an accidental infection in fox. A second in all three states, but there were some isolated species, Sellacotyle vitellosa Sogandares-Ber- instances as noted below. Fifteen species of nal, 1961, was described from mink collected helminths, including five trematodes, three in Louisiana. cestodes, and seven nematodes, were collected Maria canis LaRue and Fallis, 1936, was from this host. collected from one fox from Fender County, Eunjtrema procyonis Denton, 1942, a trem- North Carolina with only five specimens pres- atode from the pancreatic duct, was recovered ent. These specimens from the grey fox are from three hosts from North Carolina. The much larger and more robust than those from number per host was 107 and 110 for two of the bobcat. Size, however, appears to be the the three hosts. only difference. Pearson (1956) presented an E. vulpis Stunkard, 1947, was described excellent review of this species. The present from a red fox as a provisional new species, report extends the geographical range distribu- but subsequently Stunkard and Goss (1950) tion of this trematode. placed it in synonymy with E. procyonis. This Procyotrema marsupifonnis Harkema and species has been found in the raccoon and in Miller, 1959, was recovered from the gall grey and red foxes from a number of localities. bladder of one fox from Hertford County, Denton (1944) reported on its life history. North Carolina, which is an endemic area for E. procyonis was transferred to the genus this parasite. Only two specimens were found. Concinnum by Travassos (1944) and Yamaguti This form was described from specimens from (1958). However, as noted by Stunkard the pancreatic duct of the raccoon. Miller and (1947), the validity of Concinnum is doubtful Flarkema (1964) also reported this species in and features listed by Travassos to separate the gall bladder of the mink and commented the two genera are inadequate. on the locations in the hosts as being unique Seventy-four specimens of Euryhelmis squa- for strigeoid trematodes. More recently, Locke mula (Rudolphi, 1819) were recovered from and Brown (1965) reported it from a raccoon one fox collected in the vicinity of Millhaven, collected in Maryland and discussed its path- Georgia. This species was reported from ogenicity. The life history of this species has Virginia (Mclntosh, 1936), North Carolina been completed in our laboratory and will be (Parker, 1950; Harkema and Miller, 1964), published separately. Georgia (Babero and Shepperson, 1958), Taenia pisiformis (Bloch, 1780) was re- Oregon (Senger and Neiland, 1955), and covered from 9 of 24 hosts with 2 to 150 per more recently Anderson and Pratt (1965) host. This species is cosmopolitan in distribu- contributed information on its life history. tion and a common parasite of canine animals. Adults were obtained in our laboratory by Mesocestoides variabilis Mueller, 1927, was feeding cricket frogs, Acris gryllis, to white recovered from five hosts and varied in number mice, both of which are new experimental from 1 to 37 per host. Babero and Shepperson hosts. Although Baer (1931) reported this (1958) identified specimens from Georgia helminth from the European fox, Vulpes vul- raccoons as Mesocestoides lineatus. Witenberg pes, this is apparently the first report of a fox (1934) chose to include both the European as host in the United States. and American representatives as forms of M. Sellacotyle mustelae Wallace, 1935, was re- lineatus. Flowever, the best summation of this covered from the same fox from Georgia that group is by Voge (1955). Pending further contained E. squamula. Seventeen specimens definitive work the authors will retain the were found. This is the most common trema- specific name variabilis for previously stated tode occurring in mink in North Carolina reasons (Harkema and Miller, 1964). (Miller and Harkema, 1964), and this host Spirometra mansonoides (Mueller, 1935) appears to be the usual definitive host (Wal- spargana were recovered from the subcuta- lace, 1935; Erickson, 1946). S. mustelae has neous tissues of two North Carolina hosts. also been reported from the raccoon from Since adults were not obtained our specific Georgia (Sawyer, 1958) and from Georgia, identity is based on our previous findings of Virginia, North and South Carolina (Harkema both spargana and adults in the raccoon.

Physaloptera maxillaris Molin, 1860, was Table 4. Incidence of helminths in five red foxes recovered from nine hosts with 1 to 12 speci- from North Carolina and Georgia. mens per host. P. rara from the raccoon and Species of parasite Number infected P. turgida from the opossum appear to be the more common members of the genus in this Cestocla region, but only P. maxillaris was recovered Taenia pisiformis (Bloch, 1780) Nematoda from the fox. Physaloptera maxillaris Molin, 1860 Molineus patens (Dujardin, 1845) was re- Molineus patens (Dujardin, 1845) covered from two hosts with four and nine Uncinaria stcnocephala (Railliet, 1884) specimens respectively. This was the second Toxocara canis (Werner, 1782) most abundant nematode in North Carolina Capillaria plica (Rudolphi, 1819) mink (Miller and Harkema, 1964) although Morgan and Hawkins (1949) considered it worm, this parasite has been reported from a to be rare in the mink. Since Skinker's (1932) wide range of mammalian species including first report of this species in North America the cat, wolf, coyote, coati mundi, muskrat, it has been reported from a number of mustelid and others. The fox does not appear to be as animals. To our knowledge this is the first susceptible as the dog. Erickson (1944) ex- report of its presence in the grey fox. Babero amined 120 red foxes, 26 grey foxes, 61 coy- (1960) reported M. patens to be highly path- otes, and 18 timber wolves in Minnesota and ogenic to skunks in Louisiana. Schmidt (1965) found no heartworms. In the same paper he recently described a new species, M. mustclae, did report D. immitis from a single red fox in from the long-tailed weasel in Montana and New York state. Walton, Glover, and Upham published a key to the species of Molineus. (1963) did not find heartworm microfilariae Ancylostoma caninnm (Ercolani, 1859) was in the blood of 48 grey foxes and two red foxes the most commonly encountered nematode in whereas 41 of 61 dogs from the enzootic area this host being present in 17 of 23 hosts from were positive in a study at Fort Stewart, all three states. It varied in number from 1 Georgia. They were led to postulate a possible to 200 per host. physiologic resistance to the helminth by foxes. Capillaria aerophila (Creplin, 1839) was Schlotthauer (1964) found D. immitis in the found in one North Carolina fox which con- heart of 4 of 83 red foxes and none in nine tained three specimens. Commonly called the grey foxes. He remarked that wild mammal fox lungworm it has been reported a number populations probably are not highly significant of times from Europe and North America. reservoirs of heartworm infection. Christenson (1938) discussed its life history and epidemiology and Read (1949) presented Vulpes fulva a key to the species of Capillaria in North (Table 4) American mammals. Only five reel foxes were examined during Capillaria plica (Rudolphi, 1819) was col- this study. Four of these were from North lected from the urinary bladders of five foxes. Carolina and one from Georgia. One species As discussed above, we have found this species of cestode and five species of nematodes were to be fairly common in this area. recovered. With the exception of the hook- Trichuris vulpis (Froelich, 1789), the dog worm, Uncinaria stenocephala (Railliet, 1884), whip worm, was recovered from one host and the helminths were similar to those found in only two specimens were present. It is surpris- the grey fox. The species recovered and ing that it was not encountered more often number of hosts infected were as follows: since the species is apparently cosmopolitan Taenia pisiformis (3), Physaloptera maxillaris in distribution. Miller (1947) discussed the (1), Molineus patens (2), Uncinaria steno- life history of this species. cephala (3), Toxocara canis (1), and Capil- Dirofilaria immitis (Leidy, 1856) was col- laria plica (1). lected from two hosts, one in North Carolina with 13 specimens and one in South Carolina Summary with one specimen. Although the dog is gener- Studies were conducted on the incidence ally the recognized definitive host of the heart- and distribution, of helminths in four species

Copyright © 2011, The Helminthological Society of Washington 124 PROCEEDINGS OF THE HELMINTHOLOGICAL SOCIETY of wild mammals. Sixteen bobcats, twelve Diesing, K. M. 1850. Systema helminthum. Vol. from North Carolina and four from South I: 679 p. Vindobonae. Carolina, contained the following parasites: Erickson, A. B. 1944. Helminths of Minnesota Trematoda: 2 species; Cestoda: 4 species; Canidae in relation to food habits, and a host Acanthocephala: 1 species; Nematoda: 7 spe- list and key to the species reported from North cies. Twenty otters from North Carolina con- America. Am. Midi. Nat. 32: 358-372. . 1946. Incidence of worm parasites in tained the following: Trematoda: 2 species; Minnesota Mustelidae and host lists and keys Nematoda: 3 species. Twenty-four grey foxes to North American species. Am. Midi. Nat. from North Carolina, South Carolina, and 36: 494-509. Georgia contained: Trematoda: 5 species; Esch, G. W., and J. T. Self. 1965. A critical Cestoda: 3 species; Nematoda: 7 species. Five study of the taxonomy of Taenia pisiformis red foxes, four from North Carolina and one Bloch, 1780; Midticeps multiceps (Leske, from Georgia, contained: Cestoda: 1 species; 1780); and Hydatigera taeniaeformis Batsch, Nematoda: 5 species. 1786. J. Parasit. 51: 932-937. The 65 mammals harbored 28 different Gupta, S. P. 1961. The life history of Molineus species of helminths. The bobcats collected barbatus Chandler, 1942. Can. J. Zool. 39: from coastal counties contained more hel- 579-587. minths, both in kinds and numbers, than did . 1963. Mode of infection and biolgy of infective larvae of Molineus barbatus Chand- the otters or foxes. The otters contained the ler, 1942. Exp. Parasit. 13: 252-255. fewest parasites, and the foxes, although col- Hardcastle, A. B. 1941. Paragonimus in a cat lected from widely distributed areas, never from North Carolina. J. Parasit. 27: 541. showed heavy infections. Harkema, R., and G. C. Miller. 1964. Hel- minth parasites of the raccoon, Procyon lotor, Literature Cited in the southeastern United States. J. Parasit. Anderson, G. A., and I. Pratt. 1965. Cercaria 50: 60-66. and first intermediate host of Euryhelmis Jordan, H. E., and E. E. Byrd. 1958. Para- squamula. J. Parasit. 51: 13-15. gonimus in wild and domesticated animals in Anderson, R. C. 1964. Gnathostoma miyazakii Georgia. J. Parasit. 44: 470. n. sp. from the otter (Lutra c. canadensis) Klewer, H. L. 1958. The incidence of helminth with comments on G. sociale (Leidy, 1858) lung parasites of Lynx rufus rufus (Schreber) of mink (Mustela. vison). Can. J. Zool. 42: and the life cycle of Anafilaroides restrains 249-254. Gerichter 1949. J. Parasit. 44 (Suppl.): 29. Babero, B. B. 1960. A survey of parasitism in Locke, L. N., and E. E. Brown. 1965. Occur- skunks (Mephitis mephitis) in Louisiana, with rence of the fluke, Procyotrema marsupiformis observations on pathological damages due to Harkema and Miller, 1959 in a Maryland rac- helminthiasis. J. Parasit. 46: 26-27. coon. J. Parasit. 51: 355. , and J. R. Shepperson. 1958. Some Mclntosh, A. 1936. The occurrence of Eunj- helminths of raccoons in Georgia. ]. Parasit. helmis squamula (Rudolphi 1819) in the 44: 519. United States. J. Parasit. 22: 536. Baer, J. G. 1931. Quelques helminths rares ou . 1940. Some helminth parasites of the peu connus du putois. Rev. Suisse Zol. 38: Panama otter. J. Parasit. 26: 219. 313-334. Beverly-Burton, M. 1960. Some trematodes from Miller, G. C., and R. Harkema. 1964. Studies otters in Southern Rhodesia including a new on helminths of North Carolina vertebrates. strigeid, Prudhoella rhodesiensis, n. gen., n. sp. V. Parasites of the mink, Mustela vison, Schre- Proc. Helm. Soc. Wash. 27: 129-134. ber. J. Parasit. 50: 717-720. Cameron, T. W. M. 1926. Observations on the Miller, M. J. 1947. Studies on the life history genus Echinococcus, Rudolphi. J. Helm. 4: of Trichocephalus vulpis, the whipworm of 13-22. dogs. Can. J. Research. 25: 1-11. Christenson, R. O. 1938. Life history and epi- Morgan, B. B., and P. A. Hawkins. 1949. demiological studies on the fox lungworm, Veterinary helminthology, 400 pp. Burgess Capillaria aerophila (Creplin, 1839) Livro Publishing Company, Minneapolis. Jub. Travassos pp. 119-136. Mueller, J. F. 1966. The laboratory propaga- Denton, J. F. 1944. Studies on the life history tion of Spirometm mansonoides (Mueller, of Eurytrema procyonis Denton, 1942. J. 1935) as an experimental tool. VII. Im- Parasit. 30: 277-286. proved techniques and additional notes on the

biology of the cestode. J. Parasit. 52: 437- Skinker, M. S. 1932. Molineus patens (Du- 443. Jardin, 1845) Skrjabin and Schulz, 1926 col- Myers, B. J., W. Wolfgang, and R. E. Kuntz. lected in the United States. J. Parasit. 19: 91. 1960. Helminth parasites from vertebrates Sprent, J. F. A. 1956. The life history aand de- taken in the Sudan (East Africa). Can. J. velopment of Toxocara cati (Schrank, 1788) Zool. 38: 833-836. in the domestic cat. Parasitology 46: 54-78. Orihel, T. C., and L. R. Ash. 1964. Occur- . 1958. Observations on the development rence of Dirofilaria striata in the bobcat of Toxocara canis (Werner, 1782) in the dog. (Lynx rufus) in Louisiana with observations Parasitology 48: 184-209. on its larval development. J. Parasit. 50: 590- Stewart, T. B., and D. J. Jones. 1959. Oc- 591. currence of the lung fluke, Paragonimus rudis Parker, M. V. 1950. Euryhelmis squamula (Diesing, 1850) in native pigs in Georgia. J. (Rudolphi, 1819) reported from a raccoon. Parasit. 45: 548. J. Parasit. 36: 89. Stunkard, H. W. 1947. A dicrocoelid trema- Pearson, J. C. 1956. Studies on the life cycles tode, Eurytrema vulpis n. sp. provis., from and morphology of the larval stages of Alaria the pancreatic ducts of the red fox. J. Parasit. arisaemoides Augustine and Uribe, 1927 and 33: 459. Alaria canis La Rue and Fallis, 1936 (Trem- , and L. J. Goss. 1950. Eurytrema atocla: Diplostomidae). Can. J. Zool. 34: 295- brumpti Railliet, Henry and Joyeux, 1912 387. (Trematoda: Dicrocoeliidae) from the pan- Rausch, R. L., and F. S. L. Williamson. 1959. creas and liver of African anthropoid apes. Studies on the helminth fauna of Alaska. J. Parasit. 36: 574. XXXIV. The parasites of wolves Canis lupus. Travassos, L. 1944. Revisao da familia Dicro- J. Parasit. 45: 395-403. coeliidae Odher, 1910. Monog. Inst. Osw. Read, C. P. 1949. Studies on North American Cruz. 2: 357 p. helminths of the genus Capillaria Zeder, 1800 Voge, M. 1955. North American cestodes of the (Nematoda): I. Capillarids from mammals. genus Mesocestoides. Univ. Calif. Publ. Zool. J. Parasit. 35: 223-230. 59: 125-156. Riser, N. W. 1956. The hooks of taenioid ces- Wallace, F. G. 1934. The life history of Sella- todes from North American felids. Am. Midi. cotyle mustelae a trcmatode parasite of our Nat. 56: 133-137. native mink. Ann. Meet. Minn. Acad. Sci. Sawyer, T. K. 1958. Metagonimoides oregon- (Rochester, Apr. 21) pp. 6-7. ensis Price, 1931 from a Georgia raccoon with . 1935. A morphological and biological a note on Sellacotyle mustelae, Wallace 1935. study of the trematode Sellacotyle mustelae, J. Parasit. 44: 63.' n. g., n. sp. J. Parasit. 21: 143-164. . 1961. The American otter, Lutra cana- Walton, B. D., J. E. Glover, and R. W. Upham. densis vaga as a host for two species of trem- 1963. Lack of Dirofilaria immitis infection atodes previously unreported from North in wild foxes from an enzootic area. J. Parasit. America. Proc. Helm. Soc. Wash. 28: 175- 49: 526. 176. Wardle, R. A., and J. A. McLeod. 1952. The Schlotthauer, John C. 1964. Dirofilaria im- zoology of tapeworms. University of Minne- mitis in the red fox (Vulpes fulva) in Minne- sota Press, Minneapolis, Minn., 780 p. sota. J. Parasit. 50: 801. Witenburg, G. 1934. Studies on the genus Schmidt, G. D. 1965. Molineus mustelae sp. n. Mesocestoides. Arch. Zool. Ital. Torino. 20: (Nematoda: Trichostrongylidae) from the 467-508. long-tailed weasel in Montana and M. cha- Yainaguti, S. 1958. Systema helminthum. Vol. bmidi nom. n. with a key to the species of I. Digenetic trematocles. Part I. Interscience Molineus. ]. Parasit. 51: 164-168. Publishers, N. Y., London, 979 p. Senger, C. M., and K. A. Nciland. 1955. . 1959. Systema helminthum. Vol. II. Helminth parasites of some fur-bearers of The cestodes of vertebrates. Interscience Pub- Oregon. J. Parasit. 41: 637-638. lishers, N. Y., London, 860 p.

Digenetic Trematodes of Some Freshwater and Marine Fishes from Ghana1

JACOB H. FISCHTHAL AND J. D. TuoMAs2

The trematodes of this report were collected (1967) reported that metacercariae found en- in part by the junior author while a faculty cysted in Epiplatys senegalensis and E. sexfas- member in the Department of Zoology, Uni- ciatits (Steindachner) in Sierra Leone de- versity of Ghana, Legon; additional collec- veloped into Posthodiplostomum nanum when tions were made by the senior author during fed to day-old chicks and cattle egrets. The 1965-66 while a Fulbright lecturer in Zoology presence of an unencysted metacercaria in the at University College of Cape Coast, Cape small intestine of Heterobranchus longifilis Coast. Specimens have been deposited in the probably represents an accidental infection U. S. National Museum Helminthological Col- resulting from the ingestion of a fish harboring lection as noted. All measurements are in the encysted stage. Five of 20 E. senegalensis microns, unless otherwise indicated. The were infected with one (in three), three, and nomenclature of the hosts from Ghana and eight metacercariae, respectively, one of two their identification are based primarily on the T. zillii with one, and one of eight PI. fasciatus keys and descriptions given by Norman, by with one. Norman and Irvine, and by Trewavas and Irvine, in the book by Irvine (1947). Family Maseniidae Eumasenia ghanensis n. sp. Family Dlplostomatidae (Figs. 1, 2) Metacercaria of Posthodiplostomum HOST: Heterobranchus longifilis Cuvier and nanuin Dubois, 1937 Valenciennes (Clariidae). SYNONYM: Posthodiplostomum antillanum HABITATS: Duodenum and adjacent intestine. Perez Vigueras, 1944. LOCALITIES: Tributary to Pra river near HOSTS: Epiplatys senegalensis (Steinclach- Beposo; tributary to Wynne river near Kwese- ner) (Cyprinodontidae); Tilapia zillii (Gervais), mintin; stream near Ekotsi; stream one-half Plemichromis fasciatus Peters (Cichlidae); mile east of Ekumfi; Ghana. Heterobranchus longifilis Cuvier and Valen- DATES: 2, 9, 22, 25 November 1965. ciennes (Clariidae). SPECIMENS: USNM Helm. Coll. No. 63174 HABITATS: Encysted in mesenteries and (holotype); No. 63175 (paratypes). liver; free in small intestine of H. longifilis. DIAGNOSIS (based on 81 adult specimens; LOCALITIES: Kakum river estuary, Iture (E. 15 measured): Body 543-1,044 by 177-350 senegalensis); stream near Ekotsi (T. zillii}; at gonadal level, extremities rounded, lateral tributary to Wynne river near Kwesemintin bulgings of body anteriorly caused by size and (H. fasciatus, H. longifilis); Ghana. shape of oral sucker. Tegument spined as far DATES: 2, 7, 9 November 1965. posteriorly as midlength of posttesticular space, SPECIMENS: USNM Helm. Coll. No. 63170 spines sparser posteriorly. Forebody 182-315 (E. senegalensis); No. 63171 (T. zillii); No. long, hindbody 288-605 long. Oral sucker 63172 (H. fasciatus); No. 63173 (H. longifilis). subterminal ventral, 94-140 by 77-123, same DISCUSSION: This species was originally de- length and width to longitudinally or trans- scribed in the adult form from a heron from versely elongate, funnel-shaped; 56 circumoral Brazil, and later was reported from Cuba. spines in double row of 28 each, alternating, Williams and Chaytor (1966) and Williams interrupted dorsally, wedge-shaped, oral row spines 10-16 long, aboral row spines 8-12 1 Contribution from the Department of Biology, State University of New York at Binghamton, Binghamton, New long. Acetabulum 73-134 by 72-136, round York 13901 (J. H. Fischthal). to very slightly transversely elongate, center 2 Address of J. D. Thomas: School of Biological Sciences, The University of Sussex, Falmer, Brighton, Sussex, En- at level of anterior two-fifths of body length. gland. This study was supported in part by a State University Sucker length ratio 1:0.69-0.96. Prepharynx of New York Faculty Research Fellowship (FRF67-40-004 ) awarded to the senior author. 12-31 long; pharynx 33-49 by 34-54, round

Copyright © 2011, The Helminthological Society of Washington OF WASHINGTON, VOLUME 35, NUMBER 2, JULY 1968 127 to transversely elongate, four-lobed anteriorly posterior limits of extension from level of with dorsal lobe longer and ventral shorter anterior margin of acetabulum to that of than lateral ones; esophagus 5-37 long; cecal posterior testis, varying slightly within these bifurcation 21-72 preacetabular; ceca cell- limits; vitelline reservoir median to ovary. lined, extending to level of posterior margin Uterus filling most of available space in hind- of posterior testis (rarely) or short distance body, coils may enter forebody, ascending posttesticular (frequently). ventral to gonads and part of cirrus sac, met- Testes two, smooth, usually diagonal, in raterm unclifferentiated, opening into genital contact with one another, both or either one atrium. Eggs numerous, yellow-brown, oper- alone may be in contact with ovary, anterior culate, 25 measuring 21-26 by 13-18. testis sinistral and posterior testis tending to Excretory bladder I- to Y-shaped, few cells be median; rare amphitypy occurring; testes lining walls; stem wide, extending to posterior tandem in one specimen with anterior testis testis; arms short stubs or somewhat longer, on in median position wedged between ovary and either side of posterior testis, sometimes one posterior testis; in several specimens left testis arm a short stub and other longer; narrow duct displaced posteriorly, its anterior margin lying connecting stem to terminal excretory pore. posterior to anterior margin of other testis. DISCUSSION: Seven of 14 fish examined Anterior testis 52-111 by 64-122, 5-54 harbored this species, being infected with one postacetabular; posterior testis 57—131 by (in two), three, six (in two), 12 and 52 65-140, 28-125 postacetabular; posttestic- worms, respectively. Three species have been ular space 197-355 long. Cirrus sac 262-480 described: E. moradabadensis N. N. Srivas- (longitudinal extent) by 46-87, thick-walled, tava, 1951 (India); E. bangiveulensis Bever- muscular, widest at level of anterior cham- ley-Burton, 1962 (Zambia); E. vital Agrawal, ber of seminal vesicle, commencing post- 1964 (India). The first two are from clariid acetabular and median to ovary, proximal fishes and the third from a bagrid fish. Our part an elongated oval extending preace- species appears closest to E. moradabadensis, tabular, containing seminal vesicle, pars the latter differing in having 52 circumoral prostatica and prostate cells, distal part very spines, a diverticulum from the pars prostatica, long and narrow, with a posteriorly directed and the cirrus sac extending only to the pos- loop, containing ejaculatory duct and some terior margin of the acetabulum. None of the prostate cells, opening into shallow genital previously described species were noted as atrium middorsally in region of interrupted having the pharynx lobed anteriorly, but this circumoral spines. Seminal vesicle thick- may have been overlooked. walled, muscular, bipartite, both chambers oval, posterior chamber 60-114 by 23-59, Superfamily Plagiorchioiclea anterior chamber 53-92 by 31-55. Pars pros- Orientocreadium indicuni Paiide, 1934 tatica 52-81 (longitudinal extent) by 15-38, HOST: Heterobranchus longifilis Cuvier and an elongate, cell-lined, thick-wallecl vesicle; Valenciennes (Clariidae). diverticulum lacking. Ejaculatory duct very HABITATS: Small and large intestine. long, with a posteriorly directed loop. Prostate LOCALITIES: Tributary to Wynne river near cells surrounding anterior chamber of seminal Kwesemintin; stream near Ekotsi; Ghana. vesicle, pars prostatica and proximal end of DATES: 7, 9, 22 November 1965. ejaculatory duct. Genital pore median, dorsal SPECIMENS: USNM Helm. Coll. No. 63176. to oral sucker in region of interrupted cir- DISCUSSION: Three of 14 fish examined cumoral spines. Ovary 74-104 by 59-102, smooth, round to harbored two adult, five adult, and 14 adult usually slightly longitudinally elongate, over- and 28 immature specimens, respectively. In lapping acetabulum. Mehlis' gland median to life the forebody is extremely active, trembling, ovary. Seminal receptacle dorsal to portions elongating greatly, and probing. Our speci- of gonads. Laurer's canal not seen. Vitelline mens readily fit the description of O. indicum, follicles relatively few, in lateral extracecal except that in the former the pharynx is four- fields, invade intercecal space to variable lobed anteriorly, the dorsal lobe being longer extent without becoming confluent, anterior- and ventral shorter than lateral ones; this

Copyright © 2011, The Helminthological Society of Washington OF WASHINGTON, VOLUME 35, NUMBER 2, JULY 1968 129 condition could have been overlooked in the SPECIMENS: USNM Helm. Coll. No. 63177 original description of this species from Rita (holotype); No. 63178 (paratype). buchanani Bleeker (Bagridae) from India. Re- DIAGNOSIS (based on one adult worm, two examination of specimens of O. batrachoides young adults just starting egg production, and Tubangui, 1931, reported from Egypt by 75 immature specimens from one of three fish Fischthal and Kuntz (1963b) show that the examined; adult measured): Body 899 by 430, pharynx is also four-lobed anteriorly. Agrawal elongate oval. Tegument completely covered (1964) declared O. indiciim (cirrus and met- with fine spines. Forebody 276 long, hindbody raterm spined) a synonym of O. batrachoides 552. Gland cells in preacetabular parenchyma, (without such spination) probably because primarily extra- and prececal. Eyespot pig- Fischthal and Kuntz (1963b) had placed O. ment granules lying lateral and posterolateral to raipurensis Saksena, 1958, and O. daijalai Sak- pharynx, rarely anterolateral to latter. Oral sena, 1958 (cirrus only spined) in synonymy sucker 109 by 138, usually subterminal ventral, with O. batrachoides; she indicated that the may be terminal when retracted into body; presence of a spined cirrus and metraterm are with single row of 70-75 circumoral spines, variable characters. Inexplicably, Fischthal latter club-shaped, elongate, sharply pointed, and Kuntz overlooked the presence of a spiny length of seven 11-14. Acetabulum 71 by 74, cirrus in Saksena's species; the latter (1965) embedded in body parenchyma, center at level noted this fact. Because of the spiny cirrus of anterior one-third of body length. Sucker and metraterm, the present authors recognize length ratio 1 : 0.65. Prepharynx 32 long; O. indicwn as a valid species. Additionally, pharynx 74 by 80; esophagus 15 long (con- because the cirrus only is spined, we agree tracted); prepharynx and esophagus conspicu- with Saksena (1965) that O. raipurensis is a ous even in immature specimens; cecal bifur- valid species, but we regard O. daijalai and cation 44 preacetabular; ceca conspicuously lined with large elongated cells, terminating O. umadasi Saksena, 1960, as synonymous 72 from posterior extremity, divergent in ace- until such time as their life cycles are eluci- tabular region, convergent at testicular level dated. (even in immature specimens). Testes two, symmetrical, longitudinal axis of Family Cryptogonimiclae each testis lying in anteromedian-posterolateral Paraeryptogonimus ghanensis n. sp. plane, mainly extracecal, anteromedian part (Fig. 3) dorsal to ceca, posttesticular space 230 long; HOST: Lutjanus guineensis Bleeker, lagoon right testis 205 by 119, left testis 174 by 121. snapper (Lutjanidae). Cirrus sac and cirrus lacking. Seminal vesicle HABITAT: Small intestine. tripartite (conspicuous in older immature spec- LOCALITY: Kakum river estuary, Iture, imens), overlapping ovary and acetabulum Ghana. clorsally. Pars prostatica forming elongate, cell- DATE: 26 November 1965. lined vesicle 48 by 35. Ejaculatory duct very

Figures 1-16. 1. Eumuscnia ghanensis, holotype, ventral view. 2. Same. Terminal genitalia, holotype, ventral view. 3. Paracryptogoninms ghanensis, holotype, dorsal view. 4. Monodhelmis torpedinis, ventral view. 5. Same. Terminal genitalia, ventral view. 6. Same. Pharynx, ventral view. 7. Pycnadenoides ghanensis, holotype, ventral view. 8. Same. Terminal genitalia, paratype, ventral view. 9. Pclorohelmins ghanensis, holotype, ventral view. 10. Same. Terminal genitalia, holotype, ventral view. 11. Pelorohelmins pala- u'anensis, holotype, ventral view of anterior portion of body. 12. Pleorchis ghanensis, holotype, dorsal view. 13. Same. Terminal genitalia, paratype, dorsal view. 14. Same. Pharynx, paratype, dorsal view. 15. Mesolecitha ghanensis, holotype, ventral view. 16. Same. Terminal genitalia, holotype, ventral view. Abbreviations: AO, accessory organ; C, cirrus; CL, cirrus lobe; CM, distal muscular portion of genital cone; CS, cirrus sac; ED, ejaculatory duct; GA, genital atrium; GC, gland cells; GP, genital pore; HC, proximal hermaphroditic chamber of genital cone; HV, middle hermaphroditic vesicle of genital cone; M, metraterm; PAO, projection of accessory organ; PC, prostate cells; PGA, plaited wall of genital atrium and accessory organ; PP, pars prostatica; SV, seminal vesicle; U, uterus.

Copyright © 2011, The Helminthological Society of Washington 130 PROCEEDINGS OF THE HELM1NTHOLOG1CAL SOCIETY short, opening into shallow genital atrium. Family Fellodistomaticlae Prostate cells surrounding anterior chamber of Markevitschiella sp. seminal vesicle, pars prostatica, and ejaculatory HOST: Pagrus ehrenbergi Cuvier and Valen- duct. Genital pore just preacetabular, median. ciennes, golden-headed sea bream (Sparidae). Ovary deeply multilobed, 123 by 189, HABITAT: Small intestine. median, mainly intercecal, may overlap ceca LOCALITY: Tema, Ghana. ventrally, in contact with both testes, lying 57 SPECIMENS: USNM Helm. Coll. No. 63179. postacetabular. Seminal receptacle not seen. DISCUSSION: Three immature specimens were Vitelline follicles in adults and immature speci- found in one host. They resemble M. naka- mens distributed from pharyngeal or esopha- zaivai (Kobayashi, 1921) Skrjabin and Koval, geal level to level of anterior margin of testes, 1957 (syn. Steringotrema n. Kobayashi, 1921), lateral fields confluent or nearly so clorsally from sparid fishes from Japan, but specific between acetabulum and testes; right and left identification can not be made. Kobayashi vitelline ducts uniting dorsal to ovary. Uterus (1921) noted that Steringotrema Odhner, descending and ascending between testes to 1911, might be identical to Pycnadena Linton, posterior extremity, slightly overlapping testes 1911 (syn. Didymorchis Linton, 1910, pre- ventrally, filling posttesticular space inter- and occupied). Yamaguti (1958) listed them as extracecally, ascending further between right separate genera. Skrjabin and Koval (1957) testis and ovary to right of latter, then proceed- created the genus Markevitschiella for Kobay- ing across body at anterior margin of ovary to ashi's species. Our specimens do not possess left side of latter, continuing ascent antero- a postoral muscle ring, a significant feature of medianly to genital atrium, windings on each Pycnadena. The latter genus is transferred by side of ovary to lateral body walls, slightly us in this paper to the family Opistholebetidae. overlapping ovary dorsally. Eggs numerous, yellowish, 10 measuring 14—19 by 8-13. Monascus typicus (Odhner, 1911) Excretory bladder Y-shaped, cell-lined; stem Looss, 1912 extending to ovary and arms to sides of pharynx; short, narrow duct connecting blad- SYNONYMS: Monascus monenteron Looss, der to terminal excretory pore. 1907 (nom. nud.); Haplocladus typicus Odh- DISCUSSION: The young adults measure 745 ner, 1911. by 287 (17 eggs in uterus) and 715 by 255 HOSTS: Decapterus rhonchus (Geoffroy St. (36 eggs), respectively. P. ghanensis is closest Hilaire), mackerel scad, Selar crumenophthal- to P. leilae (Nagaty, 1957) Manter, 1963 mus (Bloch), goggle-eye scad, Chloroscombrus (Red Sea) and P. sp. Manter, 1963 (Fiji). chrysiirus (L.), bumper (Carangidae). Manter (1963) noted that P. sp. is very close HABITAT: Small intestine. to, if not identical with, P. leilae (syn. Meta- LOCALITIES: Tema (Decapterus, Selar), dena I. Nagaty, 1957); his Fijian specimens Cape Coast (Chloroscombrus); Ghana. DATES: 1954 (Tema), 16 December 1965 differ from the latter in having the vitelline (Cape Coast). follicles confluent, or nearly so, near the ace- SPECIMENS: USNM Helm. Coll. No. 63180 tabulum, and a slightly different sucker ratio. (D. rhonchus); No. 63181 (S. crumenoph- No description or illustration was given for thalmus); No. 63182 (C. chnjsurus). P. sp. On the basis of the distribution of the DISCUSSION: Our collection consists of two vitelline follicles we do not believe P. sp. iden- specimens from two D. rhonchus, one from S. tical with P. leilae. The number of circumoral crumenophthalmus, and two from one of 28 spines and seminal vesicle chambers are not C. chnjsurus. These represent new host and recorded for the latter. Our species differs geographic distribution records. There are from the latter in having the vitelline follicles extensive variations in our present material as confluent, or nearly so, between the acetab- well as in specimens reported by Skrjabin and ulum and testes rather than being in separate Koval (1957) and Fischthal and Kuntz lateral fields, and in the excretory arms extend- (1963a). The redescription of Monascus fili- ing to the pharyngeal level rather than to the formis (Ruclolphi, 1819) Looss, 1907, by Doll- oral sucker. fus (1947) indicates characteristics which

Copyright © 2011, The Helminthological Society of Washington OF WASHINGTON, VOLUME 35, NUMBER 2, JULY 1968 131 overlap those of M. typicus. The latter may lobed anteriorly; entrance to pharyngeal lumen be a synonym of the former, but final decision surrounded by cup-like modification composed must await life cycle studies. Dawes (1946) of longitudinal muscles, some of lateral fibers indicated that some synonymy of species prob- of cup continuing posteriorly through center ably exists. of each pharyngeal lobe, remainder of lobe with circular muscles. Esophagus thin-walled Family Hirudinellidae posteriorly and muscular anteriorly, latter with Hirudinella ventricosa (Pallas, 1774) inner longitudinal and outer circular muscles, Baird, 1853 muscular part 61 (contracted)—176 (extended) HOST: Acanthocybium solandri (Cuvier and long. Cecal bifurcation 680-800 preacetab- Valenciennes), wahoo (Scombridae). ular; ceca cell-lined, extending to posterior HABITAT: Stomach. extremity (at level of excretory pore) or not LOCALITY: Tema, Ghana. so far, postcecal space 51-210 long. DATE: 31 December 1964. Testes two, about same size, smooth, longi- SPECIMENS: USNM Helm. Coll. No. 63183. tudinally elongate, intercecal, in contact with DISCUSSION: We are identifying our three ceca or not, symmetrical or anterior margin of specimens from one host as //. ventricosa as a right testis slightly ahead of left, in contact or result of the analysis of the genus by Nigrelli not, partly overlapping anterior part of ace- and Stunkard (1947). One flattened worm tabulum; right testis 242-270 by 138-191, left measured about 47 mm long. The testes are testis 242-270 by 140-187. Terminal male extracecal and symmetrical, one on each side genitalia dextral, between accessory genital of the acetabulum; the ovary is postacetabular; organ and right cecum. Cirrus sac and cirrus and the genital pore is submedian to the left lacking. Seminal vesicle 167-246 by 52-100, at the level of the cecal bifurcation. saccular, undivided, thick-wallecl, muscular, commencing ventral to anterior part of right Family Monodhelminthidae testis or in contact with latter, ventral to ovary. Monodhelmis torpedinis Dollfus, 1937 Pars prostatica an elongate vesicle, 146—218 by (Figs. 4-6) 37-68, thick-walled, muscular, cell-lined, sur- HOST: Anus heudeloti Cuvier and Valen- rounded by compact layer of prostate cells. ciennes, sea catfish (Ariidae). Ejaculatory duct observed in two specimens, HABITAT: Small intestine. 22 by 11 and 29 by 15, very short, opening LOCALITIES: Tema, Cape Coast, Iture, into right side of genital atrium. Latter 116- Elmina; Ghana. 148 by 121-160, very large, cavity large, dorso- DATES: 13 January (Elmina), 21 February ventrally oriented; in ventral view walls com- (Iture), 16 March (Cape Coast); 1966. posed of outer moderately thick and inner thin SPECIMENS: USNM Helm. Coll. No. 63184. dorsoventrally oriented muscle layers, very DESCRIPTION (based on 20 adult specimens; thick anteroposteriorly oriented ring of circular 10 measured): Body 2,353-2,615 by 550-640, muscles between them, inner walls much widest at gonadal level, ends blunt. Tegument plaited; muscles layers, plaited walls and entirely covered with fine spines. Forebody lumen continuous with those of accessory 1,215-1,360 long; hindbody 926-1,045 long, genital organ. Latter 205-242 by 148-182, shorter than forebody. Oral sucker subterminal large, flask-shaped, very thick walled, antero- ventral, somewhat funnel-shaped, 230-265 by posteriorly oriented, may overlap left cecum 215-242, longer than wide, with wide opening veiitrally, lumen 58—77 wide posteriorly; into prepharynx; preoral lobe 6-24 long. Ace- muscle layers of genital atrium change their tabulum 203-218 by 196-220, round to longi- orientation with regard to long axis of body tudinally elongate, center at level of anterior as they bend sharply posteriorly and continue one-half to three-fifths of body length. Sucker into accessory organ: outer and inner layers length ratio 1:0.81-0.93. Prepharynx very now longitudinal muscles, middle layer in short, thick-walled, muscular. Pharynx 121— dorsoventrally oriented ring; outer longitudinal 138 by 96-116, longitudinally elongate, con- muscles on one side continuous around pos- spicuously constricted or with only very slight terior end of accessory organ and are longi- indication of constriction near midlength, un- tudinal muscles of other side; at posterior end

Copyright © 2011, The Helminthological Society of Washington 132 PROCEEDINGS OF THE HELMINTHOLOGICAL SOCIETY a broad band of some of the outer longitudinal was inadequately described from a single worm muscle fibers from each side turn anteriorly from an electric ray, Narcacion torpedo Klein through the midcenter of solid concentric (probably a synonym of Torpedo narke Risso) circular muscles which completely surround (Torpedinidae), from Mauritania, by Dollfus this band and continue into a large projection (1937). We agree with Yamaguti (1952) that lying free in lumen, projection covered by thin, the ray probably is an accidental host, having compact circular muscle layer, projection 100- ingested a teleost harboring the adult worm. 145 by 51-60 (at base); a very thin band of The two other species in the genus, M. aril muscle fibers extending from posterior limits Yamaguti, 1952 (Borneo) and M. philippinen- of inner longitudinal muscles of one side of sis Velasquez, 1961 (Philippines), as well as lumen to those of other side, passing through our specimens of M. torpedinis, are all from midcentral band of fibers entering projection; sea catfishes, Arius spp. Dollfus (1937) stated thick mass of gland cells externally capping that the postcecal space in his specimen is posterior end of accessory organ, some cells on 0.055 mm long, but his illustration shows that other parts of latter. Genital pore median, it is at least three times longer. We noted in 335-420 preacetabular, 815-970 from anterior our Tema specimens that the postcecal space extremity, at level of anterior one-third to two- is 100-210 long, resembling Dollfus' worm, fifths of body length, half way between cecal whereas in the Cape Coast area specimens it bifurcation and acetabulum or slightly more is only 51-75 long; in our opinion these dif- anteriorly. ferences represent intraspecific population Ovary 172-201 by 160-196, trilobed, dex- variations. While Dollfus does not state that tral, in contact with right testis or overlapping the ovary is trilobed, his illustration appears to it and accessory organ dorsally, lying 23-70 show this condition; we are unable to reconcile preacetabular. Seminal receptacle and Laurer's the difference in appearance of the oral sucker. canal not seen. Vitelline follicles in extracecal Dollfus shows the pharynx partially projecting fields, some few overlapping ceca ventrally, into the oral cavity. While this phenomenon extending from level of anterior part of testes was not noted in our specimens, the opening or more posteriorly at acetabular level to 165- into the oral sucker is certainly wide enough 345 from posterior extremity, anteriormost as to allow the pharynx to enter it. We believe well as posteriormost levels of each field sub- that the constriction noted by us on the equal; right and left vitelline ducts coursing pharynx is the level to which it enters the anteriorly ventral to cecum on its side, arching oral cavity; it probably is then held in this medianly anterior to testis on its side, uniting position by the contraction of the oral sucker in notch between testes to form vitelline musculature around the pharynx. We also reservoir. Uterus mainly intercecal in hind- believe that the cup-like modification of the body, posteriorly overlapping ceca and barely entrance to the pharynx assists in feeding extending extracecally in some, descending and when the pharynx is projecting into the oral ascending between testes, dorsal to accessory cavity. organ, opening into genital atrium sinistrally. We have examined the holotype and a para- Eggs numerous, yellowish, moderately thick- type specimen of M. philippinensis (USNM shelled, operculate, 27 measuring 32-39 by Helm. Coll. No. 39478), and find them much 21-28. flattened and difficult to interpret. The illus- Excretory bladder V-shaped, cell-lined, arms tration of this species by Velasquez (1961) dorsal and intercecal in hindbody, ventral and shows the holotype specimen. In the explana- extracecal in forebody, crossing ceca ventrally tion of the plate it is stated that the figure is at ovarian level, extending anteriorly to esoph- a ventral view, whereas it actually is a dorsal ageal level; excretory pore subterminal dor- view. The structure labeled SR is not the sal, 46—80 from posterior extremity. seminal receptacle, but the ovary which ap- DISCUSSION: Our collection consists of six pears to be bi- or trilobed. A seminal recep- specimens from one host from Tema, and tacle was not seen. The structure labeled OV three, five and six, respectively, from three of is not the ovary, but the seminal vesicle. The five fish examined from the Cape Coast area structure labeled VS is not the seminal vesicle, (includes Iture and Elmina). M. torpedinis but some dark staining granules along the wall

of the lumen of the accessory genital organ; 194-290, somewhat pyriform, longitudinally or the seminal vesicle lies clextrally between the transversely elongate, embedded in body fold accessory organ, ovary, and right cecum. The containing radial muscles, facing anteroven- accessory organ, as far as can be ascertained, trally on anterior robust part of body, opening is basically similar to that described by us for transversely elongate, center at level of anterior M. torpedinis. The excretory arms do not three- to four-tenths of body length. Sucker extend to the oral sucker, but only to the length ratio 1 : 1.39-1.70. Prepharynx 32-48 esophageal level. The eggs are operculate, long; pharynx 93-145 by 93-145, round to 12 measuring 53-63 by 24-30. M. philip- longitudinally or transversely elongate; esoph- pinensis differs from our specimens in having agus 36-53 long, thick-walled, muscular; much longer eggs, a round oral sucker, and cecal bifurcation overlapping anterior part of an entirely muscular esophagus, and in the acetabulum; ceca wide, conspicuously cell- vitelline follicles not extending as far pos- lined, extending to posterior extremity, post- teriorly. M. arii differs from our specimens in cecal space 60-82 long. having an unlobed ovary, a round oral sucker, Testes two, usually diagonal but may be the pharynx trilobecl anteriorly, and an entirely tandem, in contact, smooth, round to longi- muscular esophagus, and in the vitelline fields tudinally or transversely elongate; anterior being relatively short. M. philippinensis differs (left) testis 116-180 by 107-222, 73-230 from M. arii in being larger, in the greater postacetabular; posterior (right) testis 115— length of the vitelline fields, the more posterior 177 by 121-232, 160-350 postacetabular; post- extension of the ceca, and in having the ovary testicular space 92-235 long. Cirrus sac 304- lobed, and the pharynx bilobecl anteriorly. 542 by 70-136, thick-walled, muscular; commencing 39-152 postacetabular, sinistral or Family Opistholehetidae dorsal to ovary, extending anterosinistrally to Pycnadenoides ghanensis n. sp. open into shallow genital atrium lying sinistral (Figs. 7, 8) to pharynx; containing convoluted seminal HOSTS: Umbrina cirrosa (L.) (type), U. vesicle, short pars prostatica, prostate cells, ronchus Valenciennes, drum (Sciaenidae); and long, muscular, protrusible cirrus. Genital Pomadasys jubelini (Cuvier and Valenciennes), pore sinistral to pharynx, 36—90 posterior to burro (Pomadasyidae); Drepane punctata oral sucker, 61-97 preacetabular. (L.), spadefish (Drepanidae). Ovary 83-145 by 73-125, longitudinally HABITAT: Small intestine. elongate, smooth, dextral, separated from LOCALITIES: Tema (U. ronchus, P. jubelini), testes, overlapping acetabulum 10 to lying up Cape Coast (U. cirrosa, P. jubelini, D. punc- to 125 postacetabular. Oviduct emerging from tata); Ghana. posteromeclian margin of ovary. Mehlis' gland DATES: 16 December 1965 (U. cirrosa); posterior and posteromedian to ovary. Seminal 3, 25 March 1966 (P. jubelini, Cape Coast); receptacle 75-90 by 145—222, transversely 21 March 1966 (D. punctata). elongate, large, dorsal, overlapping right SPECIMENS: USNM Helm. Coll. No. 63185 cecum, ovary and anterior testis. Laurer's (holotype, U. cirrosa); No. 63186 (paratypes, canal not seen. Vitelline follicles extending U. ronchus); No. 63187 (paratype, P. jube- extracecally and cecally from level of posterior lini); No. 63188 (paratype, D. punctata). margin of acetabulum to posterior extremity, DIAGNOSIS (based on nine adult specimens; filling posttesticular space, confluent dorsally seven measured): Body 961-1,264 by 500- from ovarian or anterior testis level posteriorly, 830, ovoid, flat to short distance preacetabular, may overlap lateral margins of testes and remainder round and robust, ends blunt; tegu- posterior margin of posterior testis ventrally; ment thick, unspined. Forebody 250-340 long, vitelline reservoir posteromedian to ovary, hindbody 460-730 long. Gland cells distrib- overlapping anterior testis. Uterus ventral, uted throughout parenchyma. Oral sucker intercecal but may slightly overlap ceca, be- 145-191 by 152-230, usually transversely tween anterior margin of posterior testis and elongate, subterminal ventral; preceeded by acetabulum but may be entirely pretesticular; very short preoral lobe. Postoral circular metraterm thick-walled, muscular, commencing muscle ring present. Acetabulum 206-298 by dorsal to acetabulum, opening into genital

Copyright © 2011, The Helminthological Society of Washington 134 PROCEEDINGS OF THE HELMINTHOLCGICAL SOCIETY atrium. Eggs relatively few, yellow-brown, 15 1911, Pijcnadenoides, and Propijcnadenoides measuring "67-80 by 32-41. Fischthal and Kuntz, 1964, from the family Excretory bladder Y-shaped, cell-lined, wide, Fellodistomatidae to the family Opistholebet- dorsal, stem extending to about middle of idae. As a matter of convenience, we prefer anterior testis, arms extending to anterior at this time to recognize the above discussed margin of anterior testis; short, narrow duct genera as valid. Sogandares (1959) described connecting bladder to subterminal dorsal ex- Pachycreadium lerneri from a single specimen, cretory pore. stating that the ovary is smooth and globular; DISCUSSION: Our collection consists of three the holotype specimen shows the ovary as specimens from one of five. U. cirrosa ex- being deeply six-lobed. amined, two from one 17. rhonchus, one each from three of eight P. jubelini, and Family Pelorohelminthiclae Peloroheltnins ghanensis n. sp. one from one of 15 D. punctata. Two species have been described: P. pagrosomi (Figs. 9, 10) Yamaguti, 1938 (Japan); P. calami Manter, HOST: Arius latiscutatiis Giinther, sea cat- 1947 (Florida). Both are from sparid fishes. fish (Ariidae). Our form appears closest to P. pagrosomi, the HABITAT: Swim bladder. latter differing in the extension of the cirrus LOCALITY: Tema, Ghana. sac just posterior to the anterior margin of DATE: 17 December 1964. the acetabulum, the vitelline follicles being SPECIMENS: USNM Helm. Coll. No. 63189 absent dorsal to the gonads, and the acetab- (holotype); No. 63190 (paratype). ulum being near midbody and transversely DIAGNOSIS (based on one adult worm, one oval in shape. young adult just starting egg production, and Cable (1956) transferred the thick-bodied two immature specimens; adult measured): species of the genus Pachycreadium Manter, Body 11,015 by 4,370; tegument thick, un- 1954, from the family Opecoelidae to the armed; hindbody wide and bluntly rounded at family Opistholebeticlae because of the pres- extremity, forebody tapering slightly. Fore- ence of a postoral ring, black pigment spots, body 1,895 long, hindbody 8,285, forebody to and other features. Siddiqi and Cable (1960) hindbody ratio 1 : 4.37. Preoral lobe present. noted that the two species of Pijcnadenoides Oral sucker subterminal ventral, 580 by 667; Yamaguti, 1938, show many similarities to acetabulum 975 by 798, center at about level species of Pachycreadium. They stated: "If, of anterior one-fifth of body length; sucker as seems very likely, a postoral ring is present length ratio 1 : 1.68 (1 : 1.30 in young adult in the two species, Pachycreadium should be measuring 6,020 in total body length). Pre- reduced to synonymy with Pijcnadenoides if, pharynx lacking; pharynx 287 by 353, anterior indeed, both genera are not synonymous with three-fifths overlapping oral sucker clorsally; Pycnadena, a genus that differs from Pachy- esophagus 166 long (contracted anteriorly); creadium. only in the symmetrical arrangement cecal bifurcation 1,012 preacetabular, dorsal of the testes." Pijcnadenoides ghanensis n. sp. to level of anterior one-third of genital atrium; has a postoral ring. Examination by us of ceca conspicuously cell-lined, winding to specimens of Pachycreadium crassigulum (Lin- within 897 of posterior extremity (closer to ton, 1910) Manter, 1954 (USNM Helm. Coll. latter in other specimens). Nos. 8461, 39350), P. gastrocotylum (Manter, Testes two, symmetrical, narrow, much 1940) Manter, 1954 (No. 9346), P. lerneri longitudinally elongate, serpentine, anterior- Sogandares, 1959 (No. 38872), Pijcnadenoides most margin posterolateral to acetabulum, calami Manter, 1947 (No. 37039), Pycnadena mainly extracecal; right testis 3,465 in longi- lata (Linton, 1910) Linton, 1911 (No. 8463), tudinal extent, varying in width from 130-498; P. piriformis Price, 1934 (No. 38713), and left testis 2,990 in longitudinal extent, varying Propycnadenoidcs philippinensis Fischthal and in width from 115-445. Vasa efferentia long, Kuntz, 1964 (No. 37888) show that a postoral emerging from anterior margin of each testis. ring is present in all of them. In light of the Seminal vesicle large, bipartite; proximal part above we are transferring Pycnadena Linton, long, relatively thin-walled, winding antero-

Copyright © 2011, The Helminthological Society of Washington OF WASHINGTON, VOLUME 35, NUMBER 2, JULY 1968 135 clextral to acetabulum, inflated with sperm; small, numerous, 13 measuring 23—27 by distal part much shorter, thick-walled, mus- 13-17. cular, tubular, with single dextral loop. Pars Excretory bladder long, sinuous, thick- prostatica thick-walled, muscular, cell-lined, walled, cellular except for short, muscular part sinuous, much shorter than seminal vesicle, closest to excretory pore; short, narrow duct distal part enlarged into a chamber, entirely connecting bladder to subterminal dorsal ex- surrounded by thick layer of prostate cells, cretory pore; arms uniting dorsally between sinistral to metraterm. Ejaculatory duct short, anterior part of esophagus and pharynx. very muscular, with much enlarged muscular DISCUSSION: The type species of the genus, sphincters at each end, posterior sphincter P. palawanensis, was described by Fischthal projecting into chamber of pars prostatica, and Kuntz (1964) from a single specimen anterior sphincter projecting into proximal from the small intestine of Gazza minuta chamber of genital cone. Genital cone 298 by (Bloch) (Leiognathidae) from Palawan Island, 74, considerably modified into three parts: Philippines. Without placing it in a family, proximal part round, muscular hermaphroditic but within the superfamily Hemiuroidea, they chamber 74 in diameter into which ejaculatory created the subfamily Pelorohelminthinae for duct and metraterm open independently; the genus. Freitas and Kohn (1967) described middle part 106 by 66, an elongate, thick- Dollfustravassosius moniliovatiis n. gen., n. sp., walled, muscular hermaphroditic vesicle; distal from the swim bladder of Arius ( = Tachy- part a large, very muscular sphincter, 158 by surus) grandicassis Valenciennes (Ariidae) 121, projecting considerably into genital from Marambaia Island, Brazil, and placed it atrium. Latter large, elongate, 628 by 175, in a new subfamily Dollfustravassosiinae within muscular, with inner ladderlike arrangement the family Isoparorchiidae Poche, 1926. We of strong circular muscle bundles which are declare Dollfustravassosius and Dollfustravas- relatively widely spaced from one another, and sosiinae synonyms of Pelorohelmins and Pe- outer thick layer of longitudinal muscles; entire lorohelminthinae, respectively. For reasons atrium surrounded by thick layer of longitudi- discussed by Fischthal and Kuntz (1964), nal muscle fibers lying free in parenchyma, excluding the reference to symmetrical testes, fibers proceed posteriorly beyond genital cone. the genus can not be placed in the family Genital pore a narrow, transverse slit, median Isoparorchiidae. In light of the description of to slightly submeclian, prebifurcal, ventral to Pelorohelmins moniliovatiis (Freitas and Kohn, anterior part of esophagus, 107 from oral 1967) n. comb., and P. ghanensis, we have sucker. reexamined the type species (USNM Helm. Ovary 324 by 505, smooth, median, inter- Coll. No. 60181), and find that the testes in cecal, 3,298 posttesticular, postovarian space, the latter are symmetrical, narrow, much longi- 1,588 long. Oviduct from ventral surface of tudinally elongate, and serpentine (Fig. 11) ovary, enlarging into vesicle before narrowing rather than tandem and dextral as originally and becoming surrounded by Mehlis' gland. described. The so-called "tandem testes" Latter well developed, juxtaposed postero- actually represents the right testis alone; a vas sinistral to ovary, sinistral to ootype. Laurer's efferens was observed emerging from the canal not seen. Vitellaria dendritic, with multi- anterior end of this testis, and was traced lobed branches, inter- and extracecal, 1,755 anteriorly to the acetabulum. Also, the seminal posttesticular, three vitelline ducts entering vesicle in the type species is preacetabular ootype complex. Uterus coiled in hindbody rather than being dextral to the acetabulum from level of cecal ends to acetabulum, dorsal as originally described. The generic and sub- to ceca and vitellaria, ascending dorsal to family diagnoses given by Fischthal and Kuntz acetabulum; metraterm dextral to pars pros- (1964) need to be emended to state: Testes tatica, composed of two muscular parts, distal two, postacetabular, symmetrical, longitudi- 187 by 96, more muscular than proximal, a nally elongate, serpentine. Immediately follow- large, very muscular sphincter separates proxi- ing, in the generic diagnosis only, it should mal and distal parts and projects into latter; state: Seminal vesicle extending anteriorly few gland cells surrounding metraterm. Eggs from acetabulum, bipartite.

P. ghanensis differs from P. palawanensis in Family Pleorchiidae having larger eggs, and the Mehlis' gland next Pleorchis ghanensis n. sp. to or in contact with the ovary rather than (Figs. 12-14) relatively far removed; further, in the former HOSTS: Type, Cynoscion macrognathuv the ootype lies between the ovary and Mehlis' (Bleeker), large-mouth weakfish (Sciaenidae); gland, the secretions of the latter flowing Pomadasys jubelini (Cuvier and Valencien- dextrally into the ootype, whereas in the latter the Mehlis' gland lies between the ovary and nes), burro (Pomadasyidae). ootype, its secretions flowing sinistrally into HABITAT: Small intestine. the ootype. P. ghanensis and P. palawanensis LOCALITY: Tema, Ghana. differ from P. moniliovatits in having the eggs SPECIMENS: USNM Helm. Coll. No. 63191 haphazardly arranged in the uterus rather than (holotype, C. macrognathus); No. 63192 moniliform, the genital atrium (as defined by (paratypes, C. macrognathus); No. 63193 vis) elongate rather than globular and appear- (paratypes, P. jubelini). ing as a genital bulb, and the excretory pore DIAGNOSIS (based on 18 specimens; eight subterminal dorsal rather than terminal. P. adults measured): Body 4,660—5,247 by palawanensis differs further from P. monili- 1,310-1,550, flat, elongate, sides nearly par- ovatus in having the Mehlis' gland relatively allel, tapering slightly preacetabularly to far removed from the ovary rather than next to rounded anterior extremity, posterior extremity or in contact with the latter. The structural de- more or less truncate and slightly emarginate tails of the terminal genitalia in P. moniliovatus at excretory pore. Tegument spined except for either differ significantly from the other two preoral lobe, spines scalelike ventrally and species or, as is probable, are not presented. dorsally, long and pointed laterally and sub- With the presence of three species in the laterally. Gland cells in preacetabular paren- genus, we believe that the erection of a new chyma. Eyespots, or rarely scattered pigment family for them is warranted. Therefore, we granules, just posterior to oral sucker or on are raising the subfamily to family status. each side of pharynx. Forebody 878-1,235 long, hindbody 3,145-3,995 long; preoral lobe Pelorohelmiiithidae n. fain. 13-32 long. Oral sucker 258-314 by 276-354, DIAGNOSIS: Hemiuroidea. Body large; tegu- transversely elongate, subterminal ventral. ment thick. Oral sucker subterminal ventral, Postoral circular muscle ring present. Acetab- followed directly by pharynx and esophagus. ulum 224-298 by 218-310, round to slightly Ceca winding to near posterior extremity. longitudinally or transversely elongate, center Acetabulum in anterior body third. Testes two, at level of anterior one-fifth to one-fourth of symmetrical, serpentine, postacetabular. Semi- body length. Sucker length ratio 1 : 0.83-1.02. nal vesicle, pars prostatica and ejaculatory Prepharynx wide, 155—346 long, muscular; duct present. Genital cone in three parts: pharynx large, usually slightly wider than long, proximal hermaphroditic chamber into which 165-250 by 195-236, with well-developed and ejaculatory duct and metraterm project and conspicuous anterior circular muscle ring, re- open independently; middle hermaphroditic mainder weakly muscular or muscles degen- vesicle; distal muscular part projecting into erated and replaced by vesicular cells, sur- muscular genital atrium. Genital pore median rounded by gland cells; esophagus 110-165 to slightly submedian, between oral sucker and long, slightly muscular; cecal bifurcation 103- cecal bifurcation. Ovary round, posttesticular. 272 preacetabular; ceca extending to posterior Vitellaria dendritic, multilobed, between testes extremity, a single pair of large, elongate, and cecal ends, inter- and extracecal. Uterus saccular diverticula extend to prepharynx postacetabular, between acetabulum and cecal level, many smaller diverticula present on outer ends, inter- and extracecal. Metraterm of two sides of ceca from postacetabular level to near muscular parts. Eggs small, numerous. Ex- cecal ends, ceca and diverticula conspicuously cretory bladder long, sinuous, part near excre- cell-lined. tory pore muscular. Includes only one genus, Testes 44 in number in 17 specimens and Pelorohelmins Fischthal and Kuntz, 1964. 46 in one, lying in intercecal space on each

side of excretory bladder in two ventral and with small operculum, 23 measuring 52—66 by two superimposed dorsal longitudinal rows, 33-47 (averaging 61 by 39). extending from just postovarian to 820-960 Excretory bladder I-shaped, elongate, tubu- from posterior extremity, 22 testis on right lar, extending between rows of testes to ovary; and 22 on left in 12 individuals, 20 and 24 a pair of collecting tubules extend anteriorly in four, 24 and 20 in one, and 22 and 24 in to approximate level of posterior margin of oral one; testes measuring 132-220 by 140-254, sucker; excretory pore terminal. usually transversely elongate but may be DISCUSSION: Our collection consists of 11 round to longitudinally elongate, smooth to adult and five immature worms from C. mac- slightly lobed. Cirrus sac 563-766 (longitudi- rognathus, and one adult and one immature nal extent) by 133-177, relatively thick-walled, worm from P. jubelini. The body spines ap- proximal extremity 242-416 postacetabular, pear to be easily lost. The gland cells in the curving dorsal to dextrolateral part of acetab- preacetabular parenchyma were observed in a ulum, lying dorsal to uterus, separated from few specimens only. Five species of Pleorchis ovary by uterine coils, containing seminal Railliet, 1896, are recognized: Type, P. polij- vesicle, pars prostatica, cirrus, and prostate orchis (Stossich, 1888) Railliet, 1896 (Adriatic cells. Seminal vesicle bipartite, relatively thin- Sea); P. americanus Liihe, 1906 (U.S. Atlan- walled, posterior chamber 188-386 by 125- tic); P. sciaenae Yamaguti, 1938 (East China 162, smaller anterior chamber 122-224 by Sea); P. californiensis Manter and Van Cleave, 60-115. Pars prostatica relatively thin-walled, 1951 (U. S. Pacific); P. magnipoms Aral, 1962 cell-lined, 106-168 by 24-67. Cirrus muscular, (Mexican Pacific). All are from sciaenid longer than par prostatica. Prostate cells sur- teleost fishes; an additional host for the last rounding anterior chamber of seminal vesicle, named species is a dasyatid sting-ray. The pars prostatica and part of cirrus. Genital latter probably is an accidental host, having atrium wide, shallow. Genital pore median, ingested the teleost harboring the adult stage. immediately preacetabular. Examination of specimens of P. americanus Ovary 246-416 by 391-525, deeply multi- (USNM Helm. Coll. No. 8400) and P. califor- lobed, wider than long, median, lying 331-452 niensis (No. 38872) reveal the presence of a postacetabular, intercecal, may partly overlap postoral muscle ring, an inconspicuous circular anteriormost testes. Oviduct thick-walled, muscle ring on the anterior end of the pharynx, muscular, arising from ovary dorsum. Laurer's and an operculum on the eggs; no mention canal thick-walled, muscular, sinuous, opening was made of these features for any of the on dorsal surface over ovary. Seminal recep- species in the genus. Our species appears tacle lacking; uterus containing sperm. Mehlis' closest to P. sciaenae and P. magnipoms. P. gland poorly developed, lying anterodorsal to sciaenae differs in having only the anterior ovary. Uterus may fill intercecal space be- part of the body spined, lacking a circular tween ovary and acetabulum, occasionally muscle ring on the anterior part of the pharynx overlapping both dorsally as well as lying (or if one is present, it probably is incon- sinistral to acetabulum. Metraterm short, com- spicuous), the esophagus being practically mencing dorsal to acetabulum, surrounded by absent, the ovary being shallowly lobed, the gland cells. Vitelline follicles small, numerous, vitelline follicles not being partly confluent, extending from posterior extremity to level of or nearly so, ventral and dorsal to the testes, cecal bifurcation or slightly more posteriorly, and the eggs being longer (69-72). P. mag- extending to body margins, filling posttesticular space, may be confluent ventral and niporus differs from our species in lacking a dorsal to entire testicular field or more pos- circular muscle ring on the anterior part of terior part of this region, frequently present the pharynx (or if one is present, it probably is a ladderlike arrangement of follicles ventrally is inconspicuous), the ovary being rosette- and dorsally in testicular field as bridges of shaped, the vitelline follicles extending ante- follicles extend from between testes across riorly only to the level of the posterior margin excretory bladder; right and left vitelline ducts of the acetabulum, and not being confluent, unite posterodorsal to ovary to form small or nearly so, in the testicular region, and in vitelline reservoir. Eggs thin-shelled, yellowish, the eggs averaging longer (75).

Family Proctoecidae vesicle, 78-112 by 30-36, longer than seminal Mesolecitha ghanensis n. sp. vesicle. Cirrus short, muscular, lying lobelike (Figs. 15, 16) in genital atrium or protruding lobelike HOST: Acanthurus monroviae (Steindach- through genital pore, with long, slender, blunt- ner), surgeon-fish (Acanthuridae). tipped spines. Prostate cells surrounding ante- HABITAT: Small intestine. riormost part of seminal vesicle and length of LOCALITY: Cape Coast, Ghana. pars prostatica. Genital atrium large, trans- DATE: 3 March 1966. versely elongate, thick-walled muscular. Gen- SPECIMENS: USNM Helm. Coll. No. 63194 ital pore anterodextral to acetabulum. (holotype); No. 63195 (paratypes). Ovary 75-86 by 72-80, round to slightly DIAGNOSIS (based on four adult and three longitudinally elongate, smooth; 17-76 post- immature specimens from one of two fish acetabular, anterosinistral to and contiguous examined; adults measured): Body 1,262- with anterior testis, may slightly overlap latter 1,394 by 240-270, elongate, narrow, ends or may be entirely ventral to level of contiguity round. Tegument finely spined. Forebody of anterior and posterior testes. Seminal recep- 370-412 long, hindbody' 735-825 long. Oral tacle lacking. Laurer's canal opening dorsal sucker 157-169 by 140-155, longitudinally to acetabulum. Mehlis' gland well developed, elongate, subterminal ventral; preoral lobe anterior to ovary, overlapping latter and ace- present or not. Acetabulum 155-157 by .1.40— tabulum. Vitelline follicles relatively few, in- 155, almost round to slightly longitudinally conspicuous, extending in lateral extracecal elongate; compact circular muscle band within and cecal fields from level of posterior margin lateral and anterior margins, absent poste- of acetabulum to 22-109 posttesticular, pos- riorly; opening a narrow, transverse slit, lumen terior limit of fields in each specimen unequal, very narrow and cryptlike; center at about fields confluent dorsal to gonads; vitelline level of anterior one-third of body length. reservoir large, anterior to ovary, lying in space Sucker length ratio 1 : 0.92—1.0. Prepharynx between latter and acetabulum to being en- very short; pharynx 65-69 by 78-82; esopha- tirely dorsal to acetabulum. Uterus filling most gus 85-99 long, longer than pharynx, thick- available space in hindbody, sperm in many walled, muscular; cecal bifurcation preacetab- uterine coils, ascending dextral to cirrus sac; ular, in some bifurcation delayed for short metraterm with fine spines for short distance, distance which adds cell-lined, nonmuscular opening into genital atrium. Eggs relatively portion to length of esophagus; ceca wide, few, yellow-brown, operculate, 15 measuring conspicuously cell-lined, unequal in length, 39-58 by 20-27, all but three of latter between 47-54 long. terminating 133—293 from posterior extremity. Excretory bladder Y-shaped, cell-lined; stem Testes two, smooth, tandern to very slightly long, wide, extending to posterior part of oblique, contiguous, may overlap slightly, may acetabulum; arms extending to oral sucker; overlap cecum ventrally; anterior testis 99—121 narrow duct connecting bladder to terminal by 95-114, 13-98 postacetabular; posterior excretory pore. testis 116-133 by 93-116, 135-185 postace- DISCUSSION: The genus contains a single tabular; posttesticular space 415-535 long. Vas species, M. linearis Linton, 1910, from Acan- efferens emerging from anterior margin of each thurus coerulens Bloch and Schneider from testis, contiguous for short distance before Florida, Puerto Rico and Jamaica. It was entering cirrus sac. Latter 165-215 by 65-78, placed in a new family Proctoecidae by Cabal- widest at midlength, thin-walled; commencing lero (1959). Nahhas and Cable (1964) re- postacetabularly or just anterior to its posterior examined the type of Proctoeces neomagnonis margin, proximal end median or slightly sinis- Sidcliqi and Cable, 1960, declaring it a syn- tromedian, extending diagonally and terminat- onym of M. linearis. Siddiqi and Cable (1960) ing at genital atrium lying just anterodextral stated that in P. neomagnonis the vitelline to acetabulum; containing seminal vesicle, pars follicles are distributed in two lateral fields prostatica, cirrus and prostate cells. Seminal from the ovary to the anterior testis; from an vesicle 66-88 by 41-52, oval, thick-walled. examination of their specimen (USNM Helm. Pars prostatica a straight, cell-lined, elongate Coll. No. 39337) and their illustration it was

Copyright © 2011, The Helminthological Society of Washington OF WASHINGTON, VOLUME 35, NUMBER 2, JULY 1968 139 noted that the left field extends beyond the thick-shelled, yellowish abnormal eggs were midlength of the posterior testis, while the noted in our specimens, 16 measuring 16—20 right field extends posterior to the posterior by 8-11; Manter (1947) found such eggs testis. Examination of the type specimen of in his new species, Diphtherostomum ameri- M. Unearis (USNM Helm. Coll. No. 8471) camim. shows that the vitelline follicles completely encircle the body between the acetabulum and Zoogonus minis Looss, 1901 anterior testis; in Siddiqi and Cable's specimen HOST: Brachydeuterus auritiis (Cuvier and the vitelline follicles are confluent ventrally Valenciennes), burrito (Pomadasyidae). between the ovary and anterior testis, and HABITAT: Small intestine. dorsally at the level of the anterior testis only. LOCALITY: Cape Coast, Ghana. Our species differs in being much smaller, in DATE: 19 January 1966. having the vitelline follicles in separate fields SPECIMENS: USNM Helm. Coll. No. 63197. ventrally, in the ovary being contiguous with DISCUSSION: Two adult specimens were re- the anterior testis or both testes, and in having covered from one of 29 fish examined. The an acetabulum with compact circular muscles pharynx is six-lobed anteriorly; there are two within its lateral and anterior margins. lateral lobes on each side as well as one dorsally and one ventrally. There has been much Family Zoogoiiidae discussion in the literature regarding the syn- Diphtherostomum anisotremi onymy of Z. minis with Z. rubellus (Olsson, Nahhas and Cable, 1964 1868) Odhner, 1902, but it is generally agreed HOST: Pomadasys jubelini (Cuvier and that a completely satisfactory conclusion can Valenciennes), burro (Pomadasyidae). not be reached until their life cycles are HABITATS: Small intestine and pyloric ceca. elucidated. LOCALITY: Cape Coast, Ghana. DATE: 3 March 1966. Literature Cited SPECIMENS: USNM Helm. Coll. No. 63196. Agrawal, V. 1964. On some new trematodes DISCUSSION: Our collection consists of one, from fresh water fishes of Lucknow. Indian three, and 21 adult worms, respectively, from J. Helm. 16: 82-99. Caballero y C., E. 1959. Consideraciones en three of seven fish examined. This species torno a la familia Xenoperidae Poche, 1925 was described by Nahhas and Cable (1964) (Trematoda, Digenea Cams, 1863). Rev. Biol. from Anisotremus virginicus (L.) (Pomadasy- Trop. 7: 57-61. idae) from Jamaica, West Indies. They state Cable, R. M. 1956. Opistholebes diodontis that the hindbody is unspined, but examination 11. sp., its development in the final host, the of the holotype specimen (USNM Helm. Coll. affinities of some amphistomatous trematodes No. 60279) shows spines, although sparse, to from marine fishes and the allocreadioicl prob- be present; our specimens are also entirely lem. Parasitology 46: 1-13. spined. In both the metraterm is enlarged Dawes, B. 1946. The Trematoda. 644 p. Univ. distally. Our specimens differ from the original Press, Cambridge. Dollfus, R. P. 1937. Les trematodes Digenea description in having a shorter hindbody, the des selaciens (plagiostomes). Catalogue par cirrus sac always overlapping the acetabulum hotes. Distribution geographique. Ann. Para- (as far as miclacetabular level), the genital sit. 15: 164-176. pore usually prebifurcal but sometimes bifur- . 1947. Sur Monascus filiformis (Ruclol- cal, and the sucker length ratio 1:1.46-2.0. We phi 1819) A. Looss 1907, trematode de 1'in- can not determine whether the differences testin de Cepola rubescem (L.) en Mediter- noted are due to development in different ranee. Ann. Parasit. 22: 319-323. hosts species or to two different species being Fischthal, J. H., and R. E. Kuntz. 1963a. involved; life cycle studies could supply the Trematode parasites of fishes from Egypt. Part IV. A redescription of Monascus tijpicus answer. The pharynx in our worms is four- (Odhner, 1911) (Fellodistomidae). Proc. lobed anteriorly, with the dorsal lobe longer Helm. Soc. Wash. 30: 177-182. and ventral shorter than the two lateral ones; , and . 1963b. Trematode para- we could not verify this feature in the holo- sites of fishes from Egypt. Part VII. Ori- type. In addition to the normal eggs, smaller, entocreadium batrachoides Tubangui, 1931

(Plagiorchioidea) from Glorias lazera, with Saksena, J. N. 1965 An analysis of the trem- a review of the genus and related forms. J. atodes genus Orientocreadium Tubangui, Parasit. 49: 451-464. 1931. Proc. Natl. Acad. Sci. India, sec. B —, and . 1964. Digenetic trema- 35: 367-370. todes of fishes from Palawan Island, Philip- Siddiqi, A. H., and R. M. Cable. 1960. Di- pines. IV. Some immature Didymozoidae, a genetic trematodes of marine fishes of Puerto bucephalid; a new hemiuroicl genus and Rico. Sci. Surv. Porto Rico and Virgin Is. 17: subfamily. J. Parasit. 50: 253-260. 255-369. Freitas, J. F. T. de, and A. Kohn. 1967. Skrjabin, K. I., and V. P. Koval. 1957. Fam- Dollfustravassosius moniliovatus gen. n., sp. n. ily Fellodistomatidae Nicoll, 1913. In Skr- (Trematoda, Isoparochiidae), un trematode jabin, K. I. [Trematodes of animals and man]. parasite de poisson marin. Ann. Parasit. 42: Moskva. 13: 163-452. (Russian text.) 313-319. Sogandares-Bernal, F. 1959. Digenetic trema- Irvine, F. R. 1947. The fishes and fisheries of todes of marine fishes from the Gulf of the Gold Coast. 352 p. Crown Agents for Panama and Bimini, British West Indies. Colonies, London. Tulane Stud. Zool. 7: 69-117. Kobayashi, H. 1921. On some cligenetic trem- Velasquez, C. C. 1961. Some digenetic trematodes in Japan. Parasitology 12: 380-410. atodes of Philippine food fishes. J. Parasit. 1920. 47: 521-526. Manter, H. W. 1947. The cligenetic trematodes Williams, M. O. 1967. The neascus (postho- of marine fishes of Tortugas, Florida. Am. diplostomulum) stage of Posthodiplostomum Midi. Nat. 38: 257-416. nanum Dubois and an experimental determi- . 1963. Studies on digenetic trematodes nation of part of the life cycle. J. Helm. 41: of fishes of Fiji. III. Families Acanthocolp- 269-276. idae, Fellodistomatidae, and Cryptogonimidae. , and D. E. B. Chaytor. 1966. Some J. Parasit. 49: 443-450. helminth parasites of fresh water fishes of the Nahhas, F. M., and R. M. Cable. 1964. Di- Freetown peninsula, Sierra Leone. Bull. Inst. genetic and aspidogastrid trematodes from Fond. Afrique Noire, ser. A 28: 563-575. marine fishes of Curagao and Jamaica. Tulane Yamaguti, S. 1952. Parasitic worms mainly from Stud. Zool. 11: 169-228. Celebes. Part I. New digenetic trematodes Nigrelli, R. F., and H. W. Stunkard. 1947. of fishes. Acta Med. Okayama 8: 146-198. Studies on the genus Hinidinella, giant trem- . 1958. Systema helminthum. Vol. I. Di- atodes of scombriform fishes. Zoologica genetic trematodes of vertebrates. Parts I and (N. Y.) 31: 185-196. II. 1,575 p. Interscience Publ., N. Y.

Levinseniella hunteri sp. nov., a New Species of Microphallid Trematode from the Wilson's Plover, Charadrius wilsonia Ord1 RICHARD W. HEARD, III University of Georgia Marine Institute, Sapelo Island, Georgia

In June of 1966 during a parasitologic survey details of the excretory and reproductive of shorebirds on Grand Terre Island, Louisi- systems. They were then killed in hot saline ana, the ceca of one of three Wilson's Plovers under slight cover slip pressure and imme- examined harbored five specimens of an un- diately fixed in AFA. Each specimen was described species of Levinseniella Stiles and stained with Harris' hematoxylin and mounted Hassall, 1901. Living worms were examined in Canada balsam via standard procedures. with light and phase microscopy to determine Measurements are given in microns; the size range first followed by the average in paren- 1 This study was supported in part by an appropriation from the Congress of the United States to the Southeastern theses. Cooperative Wildlife Disease Study, School of Veterinary Medicine, University of Georgia, Athens, with funds admin- The author wishes to express sincere appre- istered and research coordinated by the Bureau of Sport ciation to Dr. T. B. Ford and Mr. J. G. Broom Fisheries and Wildlife, Department of the Interior, through Contract No. 14-16-0008-676. of the Louisiana Wildlife and Fisheries Com-

Figure 1. Levinseniella hunteri sp. nov. drawn with microprojector (scale in mm). Ventral aspect of holotype; note position of large male papilla within genital atrium.

gland cells. A thin, nonmuscular membrane appears to surround seminal vesicle and pars prostatica complex. Genital pore a slack slit, lying immediately sinistral to the acetabulum. Genital atrium sinistral, and adjacent to acetabulum. Male genital papilla a relatively large blunt, muscular cone, with its broad base anchored in right wall of genital atrium directly adjacent to acetabulum and dorsal to genital pore. Ductus ejactulatoris penetrates male papilla anteriorly from side, approximately one- third the distance from papilla's base. Upon penetrating male papilla, ductus ejactulatoris Figure 2. Levinseniella hunteri sp. nov. drawn turns distally, terminating in small opening at with microprojector (scale in mm). Terminal tip of papilla. When within genital atrium, genitalia of paratype showing male papilla pro- male papilla is directed laterally and lies at truding through genital pore. right angle to long axis of body. In this position its rounded distal end is proximal to lateral glandular wall of genital atrium which bears 9 mission for making available the facilities of well defined atrial (male) pockets. No scle- the Marine Biological Laboratory on Grand rotized "hooks" observed in atrial pockets. Terre Island during the study. Special thanks When male papilla protrudes through genital are given Dr. S. Deblock for examining the pore, portion of genital atrium containing specimens used in this study and confirming pockets becomes constricted. Metraterm ex- their taxonomic status. I would also like to tending from intertesticular region to genital thank Dr. E. E. Byrd for helpful suggestions atrium; entering atrium from dorsal aspect, on the preparation of the manuscript. midway between base of male papilla and lateral wall of atrium. Metraterm ending in Levinseniella hunteri sp. nov. dilated opening with glandular lining. Ovary (Figs. 1-2) 85-90 (88) by 36-44 (41), lateral to acetab- DESCRIPTION (based on five specimens): ulum and anterior to right testis. Ootype and Small linguiform distomes 620-730 (680) long associated Mehlis' gland, Laurer's canal and by 280-370 (320) at widest point (level of fertilization chamber in intertesticular region. testes). Cuticular spines largest in region of Vitellaria post-testicular, acenose, composed of oral sucker, extending from anterior end to a group of 6—7 large follicles (30-40 in diam- level of seminal vesicle. Oral sucker subter- eter) on each side of body. Uterus post- minal, 75-92 (85) wide by 70-80 (75) long, acetabular, with lateral loops occasionally ex- with postoral sphincter present. Acetabulum tending over anterior border of testes. Eggs 62-75 (70) in diameter, located slightly less brown, operculate, 15-17 long by 8-10 wide. than two-thirds of body length from cephalic- Excretory pore subterminal. Bladder V or U end. Prepharynx well developed, 30-50 (41) shaped. Flame cell formula 2 [(2 + 2) + (2 long. Pharynx well developed, 45—55 (48) + 2)] = 16. long by 27-30 (29) wide. Esophagus 80-125 HOST: Charadrius wilsonia Ord. (111) long. Ceca 170-210 (190) long, "A" LOCALITY: Grand Terre Island, Jefferson shaped, extending posterolaterally to near body Parish, Louisiana. margins, ending at acetabular level. Seminal SITE OF INFECTION: Ceca. vesicle, 62-100 (88) long by 31-45 wide (38), HOLOTYPE (No. 70965) and Paratype (No. kidney shaped, lying just anterior to acetab- 70966) in USNM Helm. Coll., Beltsville, Mary- ulum and slightly dextral to middle of body. land. Vasa efferentia joining just posterior to seminal This species is named in honor of Dr. Wanda vesicle to form greatly reduced vas deferens. S. Hunter. Pars prostatica well developed, 70-110 (88) Levinseniella hunteri sp. nov. differs from long, surrounded by numerous relatively small all the other described species of Levimeniella

Copyright © 2011, The Helminthological Society of Washington OF WASHINGTON, VOLUME 35, NUMBER 2, JULY 1968 143 by possessing a massive conical male papilla hunteri has "numerous male pockets" and no penetrated from the side by the ductus ejac- female pouch, and therefore, is placed in ulatoris. It more closely resembles L. indica Group IV of this scheme with L. polydactyla. Lai, 1936, L. pohjdactijla Deblock and Rose, 1982, and L. carteretensis Coil and Heard, Summary 1966, than any of the other members of the Levinseniella hunteri sp. nov. is described genus. Though the description for L. indica from the Wilson's Plover, Charadrius wilsoni is vague, L. hunteri differs from it in having Ord., collected in Louisiana. It differs from no female pouch and nine (rather than four all other described species of Levinseniella by or five) male pockets. Levinseniella poly- possessing a massive conical male papilla dactyla differs from L. hunteri by possessing penetrated from the side by the ductus ejac- a larger number of atrial (male) pockets (12) ulatoris. Levinseniella hunteri has nine male and by the ductus ejaculatoris penetrating its atrial pockets and no female pouch, and there- comparatively small male papilla through the fore is placed in Group IV of the subgeneric base. Levinseniella carteretensis, also described scheme proposed by Heard (1968). from the Wilson's Plover, is differentiated from Literature Cited L. hunteri by possessing a rudimentary male Heard, R. W., III. 1968. Parasites of the clap- papilla and a well-developed female pouch. per rail, Rallus longirostris Boddaert. I. The Heard (1968) divided the genus Levinsen- current status of the genus Levinseniella with iella into four morphological groups based on the description of Levinseniella byrdi n. sp. the number of male pockets and the presence (Trematoda: Microphallidae). Proc. Helm. or absence of a female pouch. Levinseniella Soc. Wash. 35(1): 63-68.

Some Digenetic Trematodes of Marine Fishes of New Caledonia. Part I. Bucephalidae, Monorchiidae, and Some Smaller Families1

WALTER O. DuRio2 AND HAROLD W.

The trematodes were collected by H. W. Dr. M. Legancl, Institut Francais d'Oceanie, Manter in June and July 1963. All the fishes Noumea. were from near Noumea, New Caledonia, and The trematodes were killed in A. F. A. (50% most of them obtained from the city market alcohol, 100 parts; formalin, 6l/-> parts; glacial where many fishes are sold alive each morning. acetic acid, 2M> parts) under a cover glass. Facilities and space for examination of the Specimens were stained with Delafield's or fishes were provided at the Noumea Aquarium. Ehrlich's hematoxylin. Special thanks are due Mrs. Rene Catala, Act- Some of the species are recorded also from ing Director of the Aquarium, who aided in Australia. The Australian collections were the procurement of fishes and in many other made by II. W. Manter in 1963, unless other- ways. About 40 species of fishes were exam- wise indicated. ined and 46 species of Digenea collected. Holotypes of new species are deposited in One handicap in the work was the lack of the United States National Museum Helmin- a good authority for the identification of the thological Collection. Most paratypes and fish hosts. In some cases, only the common, other specimens will be in the Manter Collec- local name was ascertained. A number of fishes were preserved and later identified by tion at the University of Nebraska. All measurements are in millimeters, unless 1 Studies from the Department of Zoology, University of otherwise stated. Nebraska, No. 386. Supported in part by National Science Foundation Grant No. GB 468. Names of new genera and new species are - University of Southwestern Louisiana, Lafayette, La. :] University of Nebraska, Lincoln. in boldface type, not italicized.

L 4

BUCEPHALIDAE Poche, 1907 type. Intestinal cecum and uterus extending Neidhartia coronata n. sp. into zone of vitellaria; uterus may extend (Figs. 1-3) slightly anterior to vitellaria. Eggs 33-38 by HOST: Serranidae; probably Epinephelus sp. 17—22 p.. Excretory pore terminal; anterior LOCATION: Intestine. extent of vesicle not determined. NUMBER: Eight (two immature) from one The name coronata is for the ring of rhynchal host. spines. HOLOTYPE: USNM Helm. Coll. No. 63302. DISCUSSION: The small spines on the rhyn- DESCRIPTION (based on six somewhat mac- chus are easily lost but in most specimens erated, extended specimens): Body elongate, scars indicate the ridge or row where the spines truncate at anterior end; most body spines lost; had been located. length 1.392-1.949; width 0.193-0.287. Rhyn- The genus Neidhartia Nagaty, 1937, has chus conical, flattened at anterior end, with been distinguished from Prosorhynchus Odh- dorsal and ventral lobe; bearing single row of ner, 1905, chiefly on the basis of the ovary spines (lost on holotype) and spine-bearing being at the intertesticular level rather than ridge (Figs. 2-3). Anterior half of rhynchus being pretesticular or opposite the anterior with sublongitudinal muscles curving to be- testis. come almost semicircular in side view near The genus Dollfustrema Eckmann, 1934, has edge; tapered part of rhynchus with large, three to six rows of rhynchal spines, and usually wide, thin-walled, irregularly spaced segments an intertesticular ovary. Thus, it seems to be (muscles). a near relative of Neidhartia. The type species Mouth in posterior one-fourth to one-fifth of Dollfustrema, D. vaneiji (Shen, 1930), has of body length; pharynx subspherical, 0.058- a cone-shaped rhynchus, and the ovary may 0.065 in diameter; intestinal cecum long, ex- be mostly anterior to the testes. It does have tending anteriorly almost to midbody. a distinctive three rows of spines on the Testes tandem to slightly diagonal on right rhynchus, and the rhynchus seems to have side of body, slightly separated, posterior testis peculiar transverse bands of muscles. All other species in the genus have a more lenticular dorsal to pharynx. Cirrus sac 0.226-0.229 by rhynchus. 0.074-0.099; overlapping posterior testis; containing ovoid seminal vesicle, straight pars Myorhynchus pritchardae n. gen., n. sp. prostatica, and numerous prostatic cells; wall (Figs. 4-6) rather thick, with diagonal muscles. Genital HOST: Serranidae; commonly called "leche." pore ventral, near posterior end of body. LOCATION : Intestine. Ovary subspherical, between testes but on NUMBER: One specimen. left side of body. Uterus not extensive, extend- HOLOTYPE: USNM Helm. Coll. No. 63303. ing anteriorly to about midbody, with loop DESCRIPTION: Body elongate, broad and between testes, then along left side of cirrus truncate at anterior end, more tapered poste- sac. Vitellaria immediately pretesticular ex- riorly; spined, except at flat anterior surface tending anteriorly to near midbody, forming of rhynchus. Length 1.427; greatest width, a broad band across body; 26 follicles in holo- just anterior to midbody, 0.470. Rhynchus

All figures were drawn with the aid of a camera lucida. The scale values are in mm. Abbreviations: c, cirrus; dl, dorsal lobe; esv, external seminal vesicle; ex, excretory vesicle; ga, genital atrium; in, mouth; mt, metraterm; pph, prepharynx; prv, prostatic vesicle; sr, seminal receptacle; sv, seminal vesicle; to, terminal organ; ut, uterus; vl, ventral lobe. Figures 1-8. 1. Neidhartia coronata. Holotype. Ventral view. 2- Same. Anterior end. Dorsal view showing dorsal lobe and spines. 3. Same. Anterior end of a paratype. Lateral view. 4. Myorhynchus pritchardae. Holotype. Ventral view. 5. Same. Anterior end, showing musculature of the rhynchus. Ventral view. 6. Same. Eggs. 7. Prosorhynchus loiigisaccatus. Holotype. Ventral view. 8. Prosorhynchus serrani. Holotype. Ventral view.

(Fig. 5) with flattened anterior surface bear- near midbody; cirrus sac large. Ovary far ing ventral flap broadly incurved at posterior anterior, near rhynchus. Vitellaria lateral, edge; 0.303 long by 0.303 wide; tapering entirely postovarian. Egg shell covered with posteriorly; strongly muscular and markedly minute spines. Type species: M. pritchardae bilateral; with six pairs of muscles on each n. sp. side of median line as follows: (1) at anterior DISCUSSION: This genus is related to Proso- end a group of short diagonal fibers; (2) a rhynchus, but has three distinguishing charac- small group of dorsoventral muscles; (3) ters: (1) the complex structure of the rhyn- curved, more or less longitudinal muscles; (4) chus; (2) the location of the ovary anterior to a long row of dorsoventral muscles extending all vitellaria; and (3) the spiny eggs. to posterior end of rhynchus where they inter- The name Myorhynchus is from myo — grade with (5) transverse muscles along outer muscle and rhynchos — snout. The species is side of rhynchus; (6) curved, more or less named for Mary Hanson Pritchard, University longitudinal muscles along lateral sides of of Nebraska. rhynchus. Longitudinal fibers in middle of ventral flap of rhynchus. Prosorhynchus longisaccatus n. sp. Mouth median, slightly anterior to midbody; (Fig. 7) pharynx 0.066 in diameter; esophagus present; HOST: Serranidae; commonly called "leche." cecum anterior to pharynx, extending more or LOCATION: Intestine. less laterally to left, broadly rounded. Testes NUMBER: Two from one host. spherical, smooth, slightly diagonal; anterior HOLOTYPE: USNM Helm. Coll. No. 63304. testis to left, partly dorsal to pharynx; posterior DESCRIPTION: Body elongate, anterior end testis near midbody. Cirrus sac 0.554 long by truncate, posterior end broadly rounded; 0.140 wide; to left of midline; anterior end length 1.096-1.201; almost uniformly wide; near midbody, just posterior to posterior testis. width 0.331-0.348. Rhynchus well developed, Seminal vesicle straight, elongate, extending cone-shaped, with flattened top, 0.267-0.283 about half-length of cirrus sac; pars prostatica long by 0.267-0.271 wide at anterior end; extending anteriorly to near base of sac, then anterior edge broadly indented ventrally. looping back and widening to form a terminal Muscles of rhynchus cone bilaterally arranged, portion lined with microvilli. One ventral and with lateral transverse bands and two sub- two smaller a trial lobes. Genital pore ventral, median, more or less diagonal or dorsoventral 0.118 from posterior end of body. Ovary bands. Muscles of anterior edge of rhynchus spherical, median, far anterior, immediately more or less perpendicular to surface ventrally, posterior to rhynchus, separated from anterior but dorsally arranged in paired groups of testis by intestinal cecum; Mehlis' gland im- diagonal muscles. mediately postovarian. Vitelline follicles lat- Mouth near midbody; pharynx 0.065-0.074 eral, in two widely separated rows; 10 on right in diameter; cecum anterior to pharynx, short, side, 13 on left; entirely posterior to ovary; wide, extending almost halfway between mostly at testicular level. Uterus with one pharynx and rhynchus. slender loop extending anteriorly to level of Testes ovoid, diagonal, close together; pos- base of rhynchus; mostly to right of cirrus sac; terior testis to right of, or immediately posterior not extending appreciably posterior to genital to, pharynx. Cirrus sac to left of midline, pore. Eggs relatively wide, 25-30 by 20-22 ^; 0.477-0.552 long by 0.141-0.152 wide; thick- shell covered with small spines or pointed walled; extending to midlevel of posterior testis projections (Fig. 6). Excretory pore terminal; and to, or almost to, pharynx. Seminal vesicle anterior extent of vesicle not determined. tubular, extending about one-third length of GENKRIC DIAGNOSIS OF Myorhynchus: Buce- cirrus sac, then bending directly forward as a phalidae; Prosorhynchinae. Rhynchus conical, sperm-free tube, joining pars prostatica near flattened anteriorly, with ventral fold or lobe; basal (proximal) end of cirrus sac; pars pros- without tentacles or suckers; strongly bilateral, tatica long, relatively narrow, with dorsoventral muscles in paired bilateral sets, mostly diagonal loop near middle of cirrus sac. Genital pore or dorsoventral. Mouth near midbody; cecum ventral, slightly to left of midline, near pos- directed anteriorly. Testes slightly diagonal, terior end of bodv.

Ovary ovoid, to right of, or partly posterior between the testes. At the present time, this to, anterior testis. Vitellaria in anterior third difference seems to be the best character to of body, mostly lateral, confluent at base of separate the two genera. rhynchus; 13 follicles on each side; posterior Prosorhynchus longisaccatus occurred in the follicles overlapping anterior part of cecum, same host with Myorhynchus pritchardae. not reaching testis. Mehlis' gland overlapping right posterior edge of ovary, or (in paratype) Prosorhynchus serrani n. sp. dorsal to left side of posterior testis. Uterus (Fig. 8) extending anteriorly along left side of cecum SYNONYM: P, crucibulus (Rud.) of Nagaty, but not anterior to it, filling most of body to 1937, nee Rudolphi, 1819, in Serranus ( = right of cirrus sac; not extending postatrially. Variola) louti (Forskal), Red Sea. New syn- Eggs 30-33 by 17-23 p,. Excretory pore ter- onymy. minal; vesicle along left side of atrium; anterior HOST: Serranus louti (Forskal); Serranidae. extent not determined. NUMBER: Thirteen from one host. The name longisaccatus refers to the long HOLOTYPE: USNM Helm. Coll. No. 63305. cirrus sac. DESCRIPTION: Body elongate, about uni- DISCUSSION: This is one of those species of formly wide; anterior end truncate; posterior Prosorhynchus with cone-shaped rhynchus, end broadly rounded; length 1.027-2.245; vitellaria forming an arc and ovary not anterior width 0.226-0.487. Rhynchus conical, flat- to testes. It seems most closely related to P. tened anterior fold with broad ventral indenta- epinepheli Yamaguti, 1939. A species we con- tion, 0.185-0.267 long by 0.156-0.217 wide; sider to be P. epinepheli was collected (34 cone relatively narrow. Mouth median at mid- specimens) from Epinephelus merra Bloch at body; pharynx 0.065-0.080 in diameter; esoph- Heron Island, Australia. It agrees with P. agus short; cecum extending anteriorly, length longisaccatus in that the gonads are clustered variable, 0.096-0.301. close together in the region of the pharynx and Testes rounded to ovoid, tandem to diagonal, cecum. The ovary may be anterior to both to right of midline, separated by two coils of testes, but usually it overlaps the anterior testis. uterus; anterior testis immediately posterior to Yamaguti's (1939) figure shows one testis midbody. Cirrus sac to left of midline, 0.361— rather widely separated from the other, but 0.574 long by 0.092-0.174 wide; thick-walled; this character was not mentioned in his de- always reaching or overlapping level of pos- scription. In the Australian material, the testes terior testis; separated from posterior testis by are close together or separated only by a single uterus and excretory vesicle; containing curved, coil of the uterus. Two characters separate P. tubular seminal vesicle about one-third length longisaccatus from P. epinepheli: (1) the of cirrus sac; short but sometimes looped, uterus does not extend even to midatrial level, sperm-free duct and pars prostatica often but whereas in all specimens of P. epinepheli it not always constricted near middle to appeal- extends postatrially; (2) the rhynchus is wider, bipartite. Genital atrium spacious, with large and the arrangement of muscles at its anterior atrial lobe. Genital pore ventral, about 0.1,5 edge gives a distinctive appearance. The from posterior end of body. cirrus sac is thick-walled in both New Cale- Ovary immediately pretesticular, near mid- donian and Australian material. body, to right of midline, lateral to pharynx. Prosorhynchus longisaccatus is also similar Mehlis' gland and yolk reservoir near posterior to P. crucibulus (Rud.), especially P. crucib- end of anterior testis; sperm cells in proximal ulus japonicus Yamaguti, 1958, but has fewer coils of uterus. Vitelline follicles large, in vitellaria, the testes closer together, and a lateral rows, 13 to 18, usually 15 or 16, on larger cirrus sac. each side; rows usually not meeting anteriorly, In most species of Prosorhynchus, the ovary but forming an arc in two (of 13) specimens; is anterior to the anterior testis but in several posterior extent of vitelline follicles near of them it more or less overlaps, or may be ovarian level; anterior extent variable, rarely directly opposite, this testis. In all six named to base of rhynchus. Uterus extending ante- species of Neidhartia, the ovary is at a level riorly dorsal to pharynx and cecum; anterior

Copyright © 2011, The Helminthological Society of Washington 148 PROCEEDINGS OF THE HELMINTHOLOGICAL SOCIETY limit usually near anterior limit of vitellaria, HAPLOSPLANCHNIDAE Poche, 1925 sometimes anterior to vitellaria to base of Hymenocotta mulli Manter, 1961 rhynchus, in one specimen not anterior to cecum; descending uterus passing between HOSTS AND LOCALITIES: Mullet; Mugilidae; testes and extending postatrially. Eggs 24—29 New Caledonia and Fiji. Mugil cephalus Linn., by 15-17 p.. at Heron. Island, Queensland, Australia. Excretory pore terminal; excretory vesicle LOCATION: Intestine. extending to right of cirrus sac, curving to left FELLODISTOMATIDAE Nicoll, 1913 of posterior testis, ending near anterior end of Tergestia clonacantha Manter, 1963 latter. DISCUSSION: This species agrees well with HOST: Hemirhamphus sp.; halfbeak; Hemi- Nagaty's (1937) "P. crucibuhis (Rud., 1819)" rhamphidae. from the same host from the Red Sea. It can- LOCATION: Intestine. not be P. crucibuhis because of the smaller NUMBER: One in one host. size; smaller, more slender rhynchus; pretestic- DISCUSSION: Tergestia clonacantha Manter, ular ovary; and uterine coils lying between 1983, from Hemirhamphus sp. in Fiji, agrees the testes. A related species is P. tsengi Tsin, in general morphology and measurements with 1933 (syn. Gotonius platycephali Yamaguti, this specimen from New Caledonia. The only 1934) which, however, has a cirrus sac not marked difference is that the two specimens reaching the posterior testis, a longer excretory from Fiji show a long cirrus sac extending vesicle, and shorter rhynchus. around the left side of the acetabulum and Some details of the cirrus sac of species of reaching halfway or more between the ace- Prosorhynchus have not been well delineated tabulum and the ovary. The specimen from in either descriptions or figures. In most, if not New Caledonia has a much shorter cirrus sac all, species the tubular seminal vesicle bends which does not reach the posterior edge of the about 180° and becomes a fairly wide, thin- acetabulum. However, the entire cirrus of this walled tube free of sperm cells. Such a tube specimen is protruded from the body which is is present in every specimen of 14 species of somewhat contracted; this may account for the genus in the Manter collection. Since it the difference in this character. connects the seminal vesicle with the "pars prostatica" (actually in Prosorhynchus an MICROSCAPHIDIIDAE Travassos, 1922 elongate prostatic vesicle), it must at times Synonym: Angiodictyidae Looss, 1902 contain sperm cells, probably only temporarily. Hexangium sigani Goto and Ozaki, 1929 The true condition in the type species, P. SYNONYMS (according to Razarihelisoa, squamatus (Rud.) Odhner, 1905, is question- 1960): H. affiniim Tubangui and Masilungan, able. There is evidence that more than one 1944; H. secundum Annereaux, 1947; H. loossi species have been considered to be P. squa- (Nagaty, 1954) Yamaguti, 1958; Arthwioossia matus. loossi Nagaty, 1954. HOSTS AND LOCALITIES: Siganus sp.; Sigan- Prosorhynchus freitasi Nagaty, 1937 idae; New Caledonia. Siganus sp. (six species); HOSTS AND LOCALITIES: Epinephelus sp.; Heron Island and Green Island; Queensland, Serranidae; New Caledonia. Plectropomus Australia. Lutjanus vaigiensis (Quoy and maculatus (Bloch); Serranidae; Heron Island; Gaimard); Lutjanidae; New Caledonia. Queensland, Australia. LOCATION: Intestine. LOCATION: Ceca and intestine. NUMBER: Numerous in Siganus sp.; one NUMBER: Twelve in one Epinephelus-., one from Lutjanus. in one Plectropomus. DISCUSSION: This species is known, usually DISCUSSION: P. freitasi was first reported from Siganus spp., in Japan (Goto and Ozaki, from Serranus guttatus Peters from the Red 1929); Celebes (Yamaguti, 1953); Madagas- Sea. The present specimens agree in general car (Razarihelisoa, 1960); Philippines (Velas- morphology and measurements. New hosts quez, 1961); Borneo (Fischthal and Kuntz, and new geographical localities are thus re- 1965). The Australian specimens show con- corded. siderable variation in the arrangement of the

Figures 9-16. Bivesiculoides posterotestis. Holotype. Ventral view. 10. Lasiotocus longitestis. Holotype. Ventral view. 11. Hysterorchis vitellosus. Holotype. Ventral view. 12. Same. Frontal section through cirrus sac showing double spination. 13. Same. Spines from base of cirrus sac. 14. Same. Egg. 15. Diphthcrostomum tropicum. Holotype. Ventral view. 16. Same. Terminal genital organs of paratype.

Copyright © 2011, The Helminthological Society of Washington 150 PROCEEDINGS OF THE HELMINTHOLOG1CAL SOCIETY testes (usually diagonal, sometimes tandem, orientation of the cirrus sac which lies posterior rarely symmetrical), but these variations were rather than anterior to the genital pore; (2) noted in Madagascaran specimens by Razari- more posterior testis; and (3) vitellaria not helisoa. overlapping the testis. It is most like B. atherinae, especially in egg size and shape of BIVESICULIDAE Yamaguti, 1939 the oral sucker. Bivesiculoides posterotestis n. sp. (Fig. 9) MONORCHIIDAE Odhner, 1911 HOST: Myctophidae; shiner; "pretre." The Genera Proctotrema Odhner, 1911, LOCATION: Intestine. and Lasiotocus Looss, 1907 in Odhner, 1911 NUMBER: One specimen in one host. Bartoli and Prevot (1966) restudied the HOLOTYPE: USNM Helm. Coll. No. 63301. type species of Proctotrema, P. bacilliovatum DESCRIPTION: Body ovoid, thin, widest near Odhner, 1911, and decided this genus should midbody; length 0.992; width 0.557. Cuticula be considered monospecific, all other species with traces of spines mostly lost by maceration. named in it to be moved to Lasiotocus. The Scattered eye-spot pigment near oral sucker. chief generic characters of Proctotrema would Mouth terminal; oral sucker elongate, funnel- then be as follows: shaped, 0.189 long by 0.103 greatest width. Seminal vesicle a coiled tube rather than Ceca obscured by vitellaria. saccular; "vagina" or terminal organ a simple Genital pore median, near midbody, more enlargement of end of metraterm, armed with or less muscular. Testis large, transversely a single cluster of large spines; genital atrium ovoid, at posterior end of body; 0.181 long by unarmed; eggs long and narrow (length about 0.316 wide. External seminal vesicle ovoid, three times width); ovary three- or four-lobed; between testis and cirrus sac. Cirrus sac thin- acetabulum funnel-shaped. Type and only walled, about 0.256 long by 0.144 wide, species: P. bacilliovatum Odhner, 1911. oriented with base at posterior end, extending The most distinctive of these generic charac- 0.064 anterior to genital pore; containing ters is the tubular, rather than saccular, seminal transversely ovoid internal seminal vesicle, very vesicle. large prostatic gland, large prostatic vesicle, Bartoli and Prevot also studied the type and short cirrus. species of Lasiotocus, L. mulli (Stossich, 1883) Ovary ovoid; to left of middle of cirrus sac; Looss, 1907 in Odhner, 1911. The chief generic 0.078 long by 0.098 wide. Mehlis' gland just characters would be: seminal vesicle saccular; anterior to ovary; seminal receptacle just pos- uterus entering side of terminal organ; atrium terior to ovary; uterus entirely pretesticular, unspined, without atrial sac. between ovary and testis. Metraterm thin- Bartoli and Prevot did not attempt to sub- walled, extending slightly anterior to genital divide the remaining species of Lasiotocus, pore. Vitellaria follicular, filling sides of body now containing several species formerly in from posterior end of oral sucker to anterior Proctotrema. The result is a rather large genus end of testis, almost confluent anteriorly. Eggs (at least 23 species). Manter and Pritchard yellowish, mostly collapsed, 54-64 by 42-48 /A; (1961) proposed separating these species on some almost round; embryos undeveloped. the basis of a distinctly lobed ovary in contrast Excretory vesicle seen only as a trace be- with an unlobed or indistinctly lobed ovary. tween testis and posterior end of body. Such a division can be made without difficulty The name posterotestis refers to the far if based on published descriptions and figures. posterior location of the testis. However, these species are often based on a DISCUSSION: The genus Bivesiculoides Yama- small number of specimens so that individual guti, 1938, differs from Bivesicula Yamaguti, variations are not certainly known. Some 1934, in having the uterus entirely pretestic- species of the related genus, Genolopa Linton, ular. Only two species have been named 1910, show great variation in indentations of hitherto: Bivesiculoides atherinae Yamaguti, the ovary. Pending more information on the 1938, from Japan and B. otagoensis Manter, various species involved, the genus Lasiotocus 1954, from New Zealand. Bivesiculoides in the sense of Bartoli and Prevot (1966) is posterotestis differs from both of these in (1) accepted here.

Lasiotocus longitestis n. sp. a much longer esophagus, less elongate oral (Fig. 10) sucker, and longer posttesticular space. In HOST: Plectorhynchits sp.; Lutjanidae; "loche L. macrorchis (Yamaguti, 1934) Yamaguti, castex." 1954, the posttesticular space is almost half LOCATION: Intestine. body length. NUMBER: One. Hysterorchis vitellosus n. gen., n. sp. HOLOTYPE: USNM Helm. Coll. No. 63306. (Figs. 11-14) DESCRIPTION: Body truncate at anterior end; HOSTS AND LOCALITIES: Plectorhijnchus sp.; broadly rounded posteriorly. Length 1.253; Lutjanidae; Noumea, New Caledonia. Plecto- width 0.365. Oral sucker funnel-shaped, 0.246 rhijnchus pictus (Thunberg); Heron Island, long by 0.176 wide. Forebody 0.348 long, Queensland, Australia; parasite collected by contracted. Acetabulum 0.115 wide (0.070 Dr. John C. Pearson, University of Queensland. long); sucker ratio 1 : 0.65. Prepharynx wide; LOCATION: Intestine. pharynx about 0.056 long by 0.040 wide; NUMBER: Four from New Caledonia; one esophagus probably about same length as from Australia. pharynx; ceca extending to 0.214 from pos- HOLOTYPE: USNM Helm. Coll. No. 63307. terior end of body. Testis elongate, 0.342 by DESCRIPTION (based on four specimens; one 0.128 greatest width, in posterior half of body, sectioned): Body entirely and strongly spined; toward right side of body, more or less pointed 0.988-1.064 long; 0.362-0.442 greatest width; at posterior end; posterior end near level of tapered and broadly rounded at each end. cecal ends. Cirrus sac large, bending around Remains of pigment spots in forebody. Oral right side of acetabulum, base near midbody; sucker rounded; 0.080-0.093 in diameter; containing saclike seminal vesicle, prostatic acetabulum near midbody, 0.080-0.093; sucker vesicle, and cirrus armed with broad-based ratio 1 : 0.86-1.16. Forebody 0.435-0.523 spines. Genital atrium unspined; genital pore long. Prepharynx short; pharynx 0.061-0.080 median. long by 0.032-0.038 wide; esophagus very Ovary deeply four-lobed; lobes spherical and long, thick-walled, muscular, 0.130-0.192 long; connected only by narrow stalks; to right of bifurcation about one-third body length from midline; immediately anterior to testis. Seminal anterior end; ceca inconspicuous, hidden by receptacle not observed. Vitellaria lateral at vitellaria, ending slightly posterior to acetab- ovarian level, cecal and extracecal; seven to ulum. nine follicles on each side. Mehlis' gland and Testes two, elongate, symmetrical or slightly yolk reservoir posterior to ovary; uterus coiling diagonal, in posterior third of body, extending to left of testis, extending nearly to posterior to posterior end of body. Cirrus sac (Fig. 12) end of body, entering terminal organ near large, curved, arcuate to U-shaped, base middle. Terminal organ 0.182 long by 0.080 slightly posterior to midbody; with thick wall greatest width, curving around left side of of longitudinal (outer) and circular (inner) acetabulum; basal half unspined, filled with muscles; length 0.272-0.448; width 0.054- eggs; anterior half armed with slender spines. 0.070; partly dorsal to acetabulum, or curving Eggs thin-shelled, collapsed, 24-28 by 14-15 ja. around either side of acetabulum; extending Excretory pore ventrotenninal, glandular; ex- anterior to genital pore, then recurving to cretory vesicle apparently short and wide. genital pore. Genital pore slightly postero- DISCUSSION: This species is most similar to sinistral to anterior edge of acetabulum. Cirrus L. plectorhynchi (Yamaguti, 1934) Yamaguti, sac containing bipartite seminal vesicle, ante- 1954 from a related host in Japan. Lasiotocus rior chamber narrowing to tube, then entering longitestis differs in its longer, more narrow a small sac armed with long spines with swol- testis reaching to within less than one-fourth len bases (Fig. 13), followed by tube with body length of posterior end of body, as do circular muscles and cirrus armed with short the ceca; its cirrus sac relatively considerably spines. Prostatic cells around seminal vesicle longer; and four- rather than three-lobed ovary. and cirrus except for region of thin-walled, Lasiotocus himezi Yamaguti, 1951, has a four- transparent cells adjacent to tube between lobecl ovary and a fairly long testis but it has seminal vesicle and cirrus.

Ovary deeply three-lobed or (in holotype) genital pore median. The double spination of four-lobed; to right of cirrus sac; sometimes the cirrus in Hysterorchis suggests the con- overlapping acetabulum dorsally; seminal re- dition found by Bartoli and Prevot (1966) in ceptacle lacking; sperm in uterus. Vitelline Lasiotociis mulli (Stossich, 1883) Looss, 1907, follicles large, numerous, from midesophagus in Odhner, 1911. level to testes level, surrounding ceca, confluent Hysterorchis has a number of characters anterior to acetabulum, slightly overlapping which might be considered as primitive (or uterus and testes dorsally. Uterus with a single less specialized) in the family Monorchiidae; large coil extending posteriorly between, or for example, the numerous vitelline follicles, ventral to, testes to near posterior end of body. simple oral sucker, and unspined metraterm. Metraterm thick-walled, about 0.240-0.256 long by 0.048 wide, extending along left side ZOOGONIDAE Odhner, 1911 of acetabulum, posterior part usually bent Diphtherostomum tropicum n. sp. toward midline and sometimes crossing cirrus (Figs. 15-16) sac; unspined; uterus entering posterior end of HOSTS AND LOCALITIES: Lethrinus sp.; Leth- metraterm (i.e., terminal organ not differen- rinidae; "bee de cane"; New Caledonia. Leth- tiated). Eggs (Fig. 14) yellow, 26-32 by rinus glyphodon Giinther; Green Island, 16-19 p., with long filament at one end. Queensland, Australia. Excretory vesicle extending between testes NUMBER: Five in one of four hosts in New or flattened posterior to them, not distinctly Caledonia; six in two of six hosts in Australia. Y-shaped but spreading at anterior end. HOLOTYPE: USNM Helm. Coll. No. 63308. The name hysterorchis refers to the far DESCRIPTION: Body rather plump, entirely posterior location of the testes. The name spined but spines more sparse posteriorly; vitellosus refers to the vitellaria which are tapered toward each end; length 0.783-1.190; much better developed than in other monor- width 0.199-0.300. Oral sucker 0.099-0.123 chids. wide; acetabulum just posterior to midbody, DIAGNOSIS OF Hysterorchis: Monorchiiclae. 0.193-0.221 wide, usually somewhat wider Testes two, near posterior end of body. Oral than long. Sucker ratio 1 : 1.80-1.95. No sucker round. Cirrus sac with bipartite, sac- prepharynx; pharynx 0.033—0.045 long by cular seminal vesicle and spined cirrus. Ovary 0.033-0.045 wide; esophagus 0.164-0.185 deeply three- or four-lobed. Vitellaria exten- long, extending almost halfway between suck- sive in middle third of body. Eggs fila- ers; bifurcation somewhat nearer to oral sucker; mented. Uterus entering base of metraterm; ceca short, extending to anterior border of metraterm muscular but unspined; atrium un- acetabulum. spined. Type species: Hysterorchis vitel- Genital pore sinistral, variously lateral, losus. slightly dorsal, or slightly ventral, near level DISCUSSION: This genus is unusual in its of bifurcation. Testes lateral, far apart, at bipartite seminal vesicle, extensive vitellaria, posterior border of acetabulum. Cirrus sac and unspined metraterm. The nearest related (Fig. 16) large, arcuate, containing bipartite genus is Diplohurleytrema Nahhas and Cable, seminal vesicle, prostatic vesicle with length 1964, from Echidna catenate, (Bloch), a moray more than twice width, thick-walled cirrus, and eel, from the Caribbean. Diplohurleytrema has prostatic cells. Cirrus with fine papillae (or fairly extensive vitellaria, a bipartite seminal microvilli ?) which sometimes appear spine- vesicle, two testes, and filamented eggs. How- like. ever, it differs in its unlobed ovary, presence Ovary globular, median, between testes, of a seminal receptacle, spined metraterm, partially overlapping acetabulum. Seminal median genital pore, and more anterior testes. receptacle between ovary and left lestis, rarely Diplolasiotocus Yamaguti, 1952, from Chae- postovarian. Two globular vitelline masses, todon Linn, in the Celebes is somewhat similar juxtaposed, immediately posterior to seminal in its long esophagus, bipartite seminal vesicle, receptacle. Uterus filling most of hinclbody; two testes, and filamented eggs; but its vitel- short, thick-walled portion of uterus at anterior laria are reduced to a few tubular glands, the edge of acetabulum, entering well-developed ovary is unlobed, the metraterm spined, the metraterm. Metraterm bent dorsallv once,

Copyright © 2011, The Helminthological Society of Washington OF WASHINGTON, VOLUME 35, NUMBER 2, JULY 1968 153 appearing somewhat shorter than cirrus sac but Literature Cited if straightened would be about same length Bartoli, Pierre, and Georges Prevot. 1966. (Fig. 16); less wide than cirrus sac; thick- Contribution a 1'etude des Monorchiidae (T. walled; inner surface with fine, delicate papil- Oclhner, 1911) parasites de poissons du genre lae. Eggs with thin shells, containing devel- Mull-us en Mediterranee. Ann. Parasit. 41(5): oped embryos; 37-53 by 16—25 p. Excretory 397-412. vesicle small sac at posterior end of body. Fischthal, J. H., and R. K. Kuntz. 1965. Di- DISCUSSION: Diphtherostomum tropicum is genetic trematodes of fishes from North Bor- most similar to D. brusinae (Stossich, 1889) neo (Malaysia). Proc. Helm. Soc. Wash. 32 Stoss., 1904, but differs in (1) the shape of (1): 63-71. the prostatic vesicle which is much longer than Goto, S., and Y. Ozaki. 1929. Brief notes on wide; (2) the absence of a transparent flap new trematodes II. Jap. J. Zool. 2(3): 369- or lip on the acetabulum; (3) its more anterior 381. seminal receptacle; (4) the dorsally curved Manter, H. W., and Mary Hanson Pritchard. metraterm. The fine spines of the metraterm 1961. Studies on digenetic trematodes of Hawaiian fishes: Families Monorchiidae and described for D. brusinae might be microvilli. Haploporidae. J. Parasit. 47(3): 483-492. Diphtherostomum brusinae is known from Nagaty, H. F. 1937. Trematodes of fishes from Labridae in the Mediterranean and from Leth- the Red Sea. Part I. Studies on the Family rinus Cuvier in Japan. Bucephalidae Poche, 1907. Fac. Med. Egypt. Summary Univ. Pub. 12, 172 pp. Razarihelisoa, M. 1960. Stir quelques trema- Twelve species of Digenea, in seven families, todes digenes de poissons de Nossibe (Mada- are reported from fishes of New Caledonia. gascar). Bull. Soc. Zool. France 84: 421-434. The following new genera (2) and new species 1959. (8) are described: New genera: Myorhyn- Velasquez, C. C. 1961. Some digenetic trema- chus (Bucephalidae); Hysterorchis (Monor- todes of Philippine food fishes. J. Parasit. 47 chiidae). New species: Neidhartia coronata; (3): 521-526. Myorhynchus pritchardae; Prosorhi/nchus Yamaguti, S. 1939. Studies on the helminth longisaccatus; P. serrani; Bivesiculoides fauna of Japan. Part 26. Trematodes of posterotestis; Lasiotocus longitestis; Hyster- fishes. VI. Jap. J. Zool. 8(2): 211-230. orchis vitellosus; Diphtherostomum tropi- . 1953. Parasitic worms mainly from cum. The Prosorhynchus crucibulus (Rud.) Celebes. Part 3. Digenetic trematodes of of Nagaty, 1937, is considered a synonym of fishes. II. Acta Med. Okayama 8(3): 257- P. serrani. 295.

Observations on Hexatylus viviparus and Neotylenchus abulbosus (Neotylenchidae: Nematoda)

W. R. NlCKLE1

T. Goodey (1926a) described Hexatylus Holland, and Germany; from carrots in Eng- viviparus from potato in England as a new land; from sugarbeets in California; from iris in genus and species. He also found it in dis- Holland; from turnip in Wales; from poppy in eased gladiolus conns, and later, T. Goodey England; and from plantain in Oregon. D. J. (1926b) expanded the description. In 1931 Hooper of the Rothamsted Experiment Station Steiner proposed Neotylenchus abulbosus as kindly loaned me mounted syntype specimens a new genus and species for nematode speci- of the original Hexatylus viviparus and Neo- mens found associated with strawberry plants tylenchus consobrinus (de Man, 1907) Filipjev, in California. He also stated that he collected 1936, and a vial containing a mass collection it from strawberry plants in Wisconsin and of H. viviparus from potato in Ireland grown Germany; from potatoes in ships stores from on a Botrytis culture. Hexatylus viviparus and England, Holland, Norway, and New Bruns- other neotylenchids from the Thorne Collection wick, Canada; and from carrots in England and also were thoroughly studied for relevant rela- Sweden. In 1932 Steiner and Buhrer described tionships. Paratype slides of Hexatylus mulveyi a single male specimen from carrots in Sweden Das, 1964, were sent to me for study by R. H. as that of N. abulbosus. Both T. Goodey and Mulvey. Steiner accurately identified these nematodes, Steiner's slides of Neotylenchus abulbosus which lacked a valved median esophageal bulb, from strawberry in California were flooded as not being the bulb and stem nematode, with warm glycerine and remounted. Some Ditylenchus dipsaci (Kiihn), that required specimens from the mass collection of Hexa- quarantine restrictions at that time. tylus viviparus were stained by the cottonblue- Tylenchicls lacking a valved median bulb lactophenol technique, while others were have been known and studied for 42 years, and stained in picric acid and mounted in dehy- as a result of this study, the genus Neotylen- drated glycerine. chus has been designated as the type genus of the family Neotylenchidae. This family now Results and Discussion includes about 75 nominal species. Studies of the original syntype material of A study of the esophageal morphology of the Neotylenchus abulbosus revealed that the sty- original specimens of Steiner's A7, abulbosus, let was surrounded by strengthening rings and H. viviparus of T. Goodey is basic to (Fig. 2C), similar in arrangement to those understanding the taxonomy of this large group illustrated for Hexatylus viviparus. The stylet of nematodes because various esophageal inter- knobs of N. abulbosus were not in the form of pretations now appear in the literature (Fig. 1). outward-pointing curved processes as described by Steiner, 1931, but appeared slightly bifid Materials and Methods (Fig. 2C). The posterior part of the esophagus Female specimens of Neotylenchus abulbo- of N. abulbosus did not have the definitely sus, the original collection from strawberry in set-off posterior bulb, as expected from pre- California, were obtained from the USDA vious descriptions, but had dorsally overlapping Nematode Collection. The male from carrot glands. Syntype specimens and a mass col- in Sweden and some females from the same lection of H. viviparus from Rothamsted also collection \veve also available. Other original were studied and other similarities between the two species were noted. There was a specimens, labeled by Steiner and others as prominent junction of the esophageal and in- Neotylenchus abulbosus, were studied. These testinal lumena (Fig. 2A), just anterior to the included nematodes from potato in England, nerve ring in both N. abulbosus and H. vivip- 1 Nematologist, Crops Research Division, Agricultural Re- arus. This eliminates the possibility of a search Service, U. S. Department of Agriculture, Beltsville, Maryland. definitely set-off posterior bulb. Previous

± 00 .oy «o>

o IO o to

CO -3

4) T3 O tf)

O IO

o IO oo o—>

-o CVJ

Copyright © 2011, The Helminthological Society of Washington 156 PROCEEDINGS OF THE HELMINTHOLOGICAL SOCIETY workers considered the junction of the esoph- five nematologists in the last 42 years (Fig. agus and intestine to be at the widening of 1). Because this area is difficult to see, we the intestinal lumen at the level of the base are confronted by illustrations that depict the of the overlapping esophageal glands (Fig. 1). esophagus as being fused with the intestine, The suggestion that the esophagus and intes- as having a definitely set-off posterior bulb, tine are fused, as stated in the diagnosis of and now as having overlapping glands. T. the genus Hexatylus, was difficult to accept Goodey's original drawing (Fig. 1A) showed since fusion of two organ systems is not known a fusion of the esophagus with the intestine to occur anywhere in the animal kingdom, and and no esophageal gland orifices. Steiner's it would be unlikely to occur only in this nema- drawing (Fig. IB) may have been a dorsal tode genus. or ventral view of the overlapping gland which Both stained and glycerine mounted speci- made it appear as a definitely set-off posterior mens of H. viviparus revealed that the esoph- bulb. In 1933 T. Goodey redrew this nema- ageal glands overlap the intestine and lie tode (Fig. 1C), and though he maintained the dorsally in a diverticulum. Three esophageal fusion of the esophagus with the intestine, he gland nuclei and two large coelomocytes were added the dorsal and subventral gland orifices. present in the area (Fig. 2A). These coelomo- He redrew the nematode again in 1938 (Fig. cytes may have been confused by earlier work- ID) and added the important junction of the ers with the gland nuclei (Fig. 1). The over- esophageal and intestinal lumena. He also still lapping esophageal glands were also observed considered the esophagus and intestine to be in the original N. abulbosus and H. viviparus fused, but he changed the pattern on the material. The posterior part of the esophagus ventral side of the esophagus to appear more did not vary from a posterior bulb to over- like an intestine. Thome made a drawing lapping glands. Other differences previously (Fig. IE) of this nematode in 1941 and de- thought to exist between N. abulbosus and cided that the esophagus was fused with the H. viviparus were not found. Both nematodes intestine. He pointed out the strengthening were found to have the cephalic framework rings around the stylet but did not differentiate in eight sectors (Fig. 2B) as Steiner (1931) between the subventral gland orifices and the described and four lateral lines. The subven- junction of the esophageal and intestinal lu- tral gland orifices opened into the area where mena. T. Goodey redrew the nematode with the median bulb would have been if it were minor changes for his textbook of 1951 (Fig. present. The intestinal lumen widened broadly IF). Thorne redrew the front end of this at a level just posterior to the base of the nematode for his textbook in 1961 (Fig. 1G). overlapping gland. It was similar to his earlier drawing, and he The male specimen from carrot in Sweden, still considered the esophagus to fuse with the described as the male of N. abulbosus, and intestine. In 1963 J. B. Goodey rounded off some female nematodes from the same collec- the base of the overlapping esophageal gland tion, were studied. These female nematodes (Fig. 1H) and mentioned that further work were the same as Hexatylus viviparus as de- might show that H. viviparus and N. abulbosus scribed in this paper. The male was not in are indistinguishable. Figure II depicts the good condition. It looked like it may have present author's interpretation of the esoph- had a stylet, but the specimen gave the appear- ageal intestinal area. ance of having decomposed before fixation No differences could be found between syn- rather than that of lacking a stylet as de- type specimens of H. viviparus, the type spe- scribed by Steiner. N. abulbosus was described cies of Hexatylus, and N. abulbosus, the type from female specimens taken from strawberry species of Neotylenchus. They thus become plants in California, and it appears unwise that conspecific, requiring the synonymy of the the description of this male specimen from genus Neotylenchus under the genus Hexa- carrot in Sweden be given much credence in tylus. This synonymy has been proposed in the taxonomy of N. abulbosus. the past, or suggested by several workers (T. The esophageal area of Hexatylus viviparus Goodey, 1933, 1938; Christie, 1938; Andrassy, has been illustrated differently nine times by 1952; Meyl, 1954; J. B. Goodey, 1963). How-

20u

Figure 2. Drawings from Hexatylus viviparus and the original Neotylenchus abulbosus material. A. Anterior end of H. viviparus from gladiolus corm. B. En face view, from original N. abulbosus slide. C. Head and stylet of original N. abulbosus specimen, showing bifid knobs and strengthening rings. D. Female from an original specimen of N. abulbosus.

Copyright © 2011, The Helminthological Society of Washington 158 PROCEEDINGS OF THE HELMINTHOLOGICAL SOCIETY ever, this action seems justified by the present the genus Neotylenchus, is placed as a syn- studies of the original material of Sterner and onym of Hexatylus. Scytaleum viglssi (Skar- T. Goodey. bilovich, 1952) Andrassy, 1961, the type spe- The results of this study of syntype material cies of the genus Scytaleum, with a shorter of N. abulbosus and H. viviparus require cer- stylet and the presence of males, is also contain adjustments in the Neotylenchidae, some sidered a valid species of Hexatylus, as orig- of which are beyond the scope of this paper. inally proposed by Skarbilovich. The three As the type species of the genus Neotylenchus remaining species in the genus Scytaleum: becomes a junior synonym of the type species S. italicum (Meyl, 1954) Andrassy, 1961; S. of the genus Hexatylus, the former generic skarbilowiezae (Atakhanov, 1959) Andrassy, name can no longer be used. The family name 1961; and S. thornei (Meyl, 1954) Andrassy, Neotylenchidae and the subfamily name Neo- 1961, are placed in species inquirendae. Other tylenchinae are retained. Studies of the avail- species inquirendae include: Hexatylus boett- able specimens and literature of the remaining geri Meyl, 1954; H. brevicaudatus Meyl, 1954; species of the old genus Neotylenchus do not and H. dipapillatus Meyl, 1954. Future work, reveal a suitable type species to represent them especially on the esophageal area, will un- as a group. My observations require the trans- doubtedly reveal their exact status. fer of N. consobrinus (de Man, 1907) Filipjev, The following changes are therefore pro- 1936, to another genus in the Neotylenchidae, posed at this time: namely, Paurodontus Thome, 1941, because of the presence in T. Goodey's specimens of Hexatylus T. Goodey, 1926. a long stemlike projection of esophageal tissue Syn. Neotylenchus Steiner, 1931. into the intestine. N. linfordi Hechler, 1962, Scytaleum, Andrassy, 1961. has a short stemlike projection at the base of Hexatylus viviparus T. Goodey, 1926. the esophagus and is also placed in the genus Syn. Neotylenchus abulbosus Steiner, Paurodontus. In making these changes, I agree 1931. with J. B. Goodey (1963) in reducing the Hexatylus vigissi Skarbilovich, 1952. taxonomic value placed on the number of Syn. Scytaleum vigissi (Skarbilovich, sectors in the cephalic framework of members 1952) Andrassy, 1961. of this family. After a study of excellently pre- Paurodontus linfordi (Hechler, 1962) served syntype material, I have transferred n. comb. Neotylenchus arcuatus Thome, 1941, to the Syn. Neotylenchus linfordi Hechler, genus Nothanguina Whitehead, 1959. Neo- 1962. tylenchus velatus (Btitschli, 1873) Skarbilo- Paurodontus consobrinus (de Man, 1907) vich, 1952, N. obtusicaudus (Stek., 1941) n. comb. Skarbilovich, 1952, and N. zeae (Stek., 1941) Syn. Neotylenchus consobrinus Skarbilovich, 1952, are placed in species (de Man, 1907) Filipjev, 1936. inquirendae. N. acutus Thome, 1941; N. bel- Nothanguina arcuatus (Thorne, 1941) jaevae (Atakhanov, 1959) Andrassy, 1961; n. comb. N. coprophagus (T. Goodey, 1938) Thorne, Syn. Neotylenchus arcuatus Thorne, 1941; N. dendrophilus Riihm, 1956; N. inter- 1941. medius (Christie, 1938) Thorne, 1941; N. Species inquirendae latus Thorne, 1935; N. obesus Thorne, 1934, The following species seem to belong in and N. serpens Andrassy, 1961, require more the Neotylenchidae but the details in their study before they can be placed properly descriptions are inadequate for recognition by within the family Neotylenchidae. They are present-day standards. These species require also considered as species inquirendae. more study, in the light of the basic changes Hexatylus mulveyi Das, 1964, with six lines imposed upon this group by the information in the lateral field is retained as a valid species presented in this paper. of the genus Hexatylus. The genus Scytaleum Neotylenchus acutus Thorne, 1941. Andrassy, 1961, which he considered to be Neotylenchus beljaevae (Atakhanov, intermediate between the genus Hexatylus and 1959) Andrassy, 1961.

Neotylenchus coprophagus (T. Goodey, Neotylenchus abulbosus Steiner, 1931. 1938) Thome, 1941. Hexatylus abulbosus (Steiner, 1931) Neotylenchus dendrophilus Riihm, 1956. T. Goodey, 1933. Neotylenchus intermedius (Christie, Anguillulina (Neotylenchus) abulbosa 1938) Thorne, 1941. (Steiner, 1931) W. Schneider, 1939. Neotylenchus lotus Thorne, 1935. lotonchium abulbosum (Steiner, 1931) Neotylenchus obesus Thorne, 1934. Filipjev and Stek., 1941. Neotylenchus obtusicaudus (Stek., 1941) MALE: Unknown. Skarbilovich, 1952. FEMALE: Length 0.67-1.5 mm; width Neotylenchus serpens Andrassy, 1961. 0.037-0.057 mm; a = 15-35; b = 13-16; c = Neotylenchus velatus (Biitschli, 1873) 10-20; V% = 81-89; stylet = 9-11 /jt. Skarbilovich, 1952. Females with great variation in length and Neotylenchus xeae (Stek., 1941) width between those not yet producing eggs Skarbilovich, 1952. and senile forms. Cuticle striated. Lateral Hexatylus boettgeri Meyl, 1954. field with four lines. Phasmids and deirids Hexatylus brevicaiidatus Meyl, 1954. not observed. Cephalic framework slightly Hexatylus dipapttlatus Meyl, 1954. sclerotized. Stylet 9-11 p. Excretory pore Scytaleum italicum (Meyl, 1954) posterior to nerve ring. Hemizonid just an- Andrassy, 1961. terior to excretory pore. Overlapping esoph- Scytaleum skarbilowiezae (Atakhanov, ageal glands located dorsally in a diverticulum, 1959) Andrassy, 1961. later more degenerate, and pushed to side by Scytaleum. thornei (Meyl, 1954) developing ovary. Andrassy, 1961. Bionomics. This species has worldwide dis- Genus: Hexatylus T. Goodey, 1926. tribution and has been found in various situa- Syn. Neotylenchus Sterner, 1931. tions in which rotting plant material was pres- Scytaleum Andrassy, 1961. ent. It has been successfully cultured on fungal DIAGNOSIS (Emended): Neotylenchinae. Ce- plates. phalic framework octagonal. Stylet knobs Other species: slightly bifid. Pharynx slightly sclerotized, Hexatylus mulveyi Das, 1964. with several strengthening rings surrounding Hexatylus vigissi Skarbilovich, 1952. stylet. Dorsal esophageal gland orifice near Syn. Scytaleum vigissi (Skarbilovich, base of stylet. Subventral gland orifices open 1952) Andrassy, 1961. in area where median bulb would have been I believe that members of the genus Hexa- if it were present. Lumen of esophagus joins tylus should be limited to those nematocles lumen of intestine near base of corpus, in having the strengthening rings surrounding the region of nerve ring. Lumen becomes wider stylet, bifid stylet knobs, the prominent junc- at this junction and widens again markedly tion of the esophageal and intestinal lumena, at level of base of overlapping esophageal and overlapping esophageal glands. glands. Esophageal glands overlap intestine dorsally in a diverticulum. Intestinal lumen Summary with prominent villae. Gonad prodelphic, well- Studies of syntype specimens of Neotylen- developed, extending into esophageal area; chus abulbosus Steiner, 1931, and Hexatylus vulva posteriorly located; postuterine sac ab- viviparus T. Goodey, 1926, revealed that they sent. Tail short, more or less acuminated. were conspecific. Because N. abulbosus, the Males rare or absent. type species of Neotylenchus, is now a junior Type species: Hexatylus viviparus T. Goodey, synonym of the type species of the older genus 1926. Hexatylus, the genus Neotylenchus becomes Syn. Anguillulina (Hexatylus) vivipara unavailable. (T. Goodey, 1926) W. Schneider, The esophagus of Hexatylus viviparus was 1939. found to have dorsally overlapping esophageal lotonchiumviviparum (T. Goodey, glands instead of a fusion of the esophagus 1926) Filipjev and Stek., 1941. with the intestine. The descriptions of the

Copyright © 2011, The Helminthological Society of Washington 160 PROCEEDINGS OF THE HELMINTHOLOGICAL SOCIETY genus Hexatylus and its type species H. vivip- urales Musei Nationalis Hungarici. Budapest. arus are emended. Illustrations of H. vivip- (Series Nova) Vol. II, pp. 13-65. arus are presented. Christie, J. R. 1938. Two nematodes associated Neotylenchus consobrinus (de Man, 1907) with decaying citrus fruit. Proc. Helm. Soc. Filipjev, 1936, and Neotylenchus linfordi Wash. 5(1): 29-33. Hechler, 1962, are transferred to the genus Goodey, J. B. 1963. Soil and Freshwater Nem- Paurodontus Thome, 1941. Neotylenchus atodes. John Wiley and Sons, Inc., New York. arcuatus Thorne, 1941, is transferred to the 544 pp. Goodey, T. 1926a. Hexatylus viviparus gen. et genus Nothanguina Whitehead, 1959, and the sp. nov., a nematocle found in a diseased po- genera Scytaleum Andrassy, 1961, and Neo- tato tuber. J. Helminth. 4(1): 27-30. tylenchus Sterner, 1931, are considered to be . 1926b. A further note on Hexatylus vi- synonyms of the genus Hexatylus T. Goodey, viparus Goodey, 1926. J. Helminth. 4(4/5): 1926. The following species are placed in 183-184. species inquirendae: Neotylenchus acutus . 1933. Plant Parasitic Nematodes. Meth- Thorne, 1941; IV. beljaevae (Atakhanov, 1959) uen and Co., London. 306 pp. Andrassy, 1961; IV. coprophagus (T. Goodey, . 1938. Some observations on the nem- 1938) Thorne, 1941; N. dendrophilus Riihm, atode Hexatylus viviparus Goodey, 1926. J. 1956; N. intermedius (Christie, 1938) Thorne, Helminth. 16(2): 109-116. 1941; N. latus Thorne, 1935; N. obesus . 1951. Soil and Freshwater Nematodes. Thorne, 1934; N. obtusicaudus (Stek., 1941) John Wiley and Sons, Inc., New York. 390 pp. Skarbilovich, 1952; N. serpens Andrassy, 1961; Meyl, A. H. 1954. Die Nematodenfauna Hoherer N. velatus (Biitsehli, 1873) Skarbilovich, Pilze in Laubund Nadelwaldern zwischen 1952; N. zeae (Stek., 1941) Skarbilovich, Braunschweig und dem Harz. Mycopath. et 1952; Hexatylus boettgeri Meyl, 1954; H. Mycol. Applic. 7(1-2): 1-80. brevicaudatus Meyl, 1954; H. dipapillatus Steiner, G. 1931. Neotylenchus abulbosus n. g., n. sp. (Tylenchiclae, Nematoda) the causal Meyl, 1954; Scytaleum italicum (Meyl, 1954) agent of a new nematosis of various crop Andrassy, 1961; S. skarbiloioiezae (Atakhanov, plants. J. Wash. Acad. Sci. 21(21): 536- 1959) Andrassy, 1961; S. thornei (Meyl, 1954) 538. Andrassy, 1961. , and E. M. Buhrer. 1932. The male of the nematode species, Neotylenchus abulbosus Ackno wledgmen t Steiner, and its sexual dimorphism. T. Wash. The author wishes to express thanks to Miss Acad. Sci. 22(16/17): 482-484. Patricia A. Mclntosh for providing technical Thorne, G. 1941. Some nematodes of the fam- assistance on this study. ily Tylenchidae which do not possess a val- vular median esophageal bulb. Great Basin Literature Cited Naturalist 2(2): 37-85. Andrassy, I. 1952. Freilebende Nematodes aus . 1961. Principles of Nematology. Mc- clem Biikk-Gebirge. Annals Historico-Nat- Graw-Hill Book Co., New York. 553 pp.

Report of the Bray ton H. Ransom Memorial Trust Fund

FUNDS ON HAND, 1 January 1967 $2,500.49 RECEIPTS: Interest received in 1967 155.47 DISBURSEMENTS: Grant to Helminthological Society of Washington 10.00 BALANCE ON HAND, 31 December 1967 $2,645.96 A. O. FOSTER Secretary-Treasurer

Parasitic Development of Filipjevimermis leipsandra Poinar and Welch (Mermithidae) in Diabrotica u. undecimpunctata (Chry somelidae) GEORGE O. POINAR, JR. Department of Entomology and Parasitology, University of California, Berkeley, California

Introduction D. balteata showed that the parasitic develop- In July 1967 Dr. F. P. Cuthbert sent the ment was similar in both insects. Diabrotica author some living mermithids which had u. undecimpunctata was grown on corn seed- developed in larvae of Diabrotica balteata and lings using a modification of the method pre- were subsequently described as a new species, sented by Rimando et al. (1965). Larval de- Filipjevimermis leipsandra Poinar and Welch velopment ranged from 18-25 days, with three (in press). The life history of this species will instars. be discussed by Cuthbert (in press). One-clay-old Diabrotica larvae were placed F. leipsandra is unusual in several respects. between two layers of moist filter paper in a Males are very rare and not necessary for the small petri dish for laboratory infections. In- propagation of the species. Another interesting fective stage nematodes were placed directly character of this nematode is its association on the inner side of both pieces of filter paper. with the central nervous system of the host The edges of the paper were held together during the early stages of parasitism. with a ring and the infection chamber left After penetrating through the cuticle and at 70 C for 80 minutes. The insects were entering the hemocoel of the host larva, the then removed and kept on roots of germinating infective stage nematode seeks out and enters corn seedlings during the remainder of the one of the ganglia of the central nervous sys- experiment. tem. The protocerebral lobes and subesoph- At regular intervals over a 4-week period, an ageal ganglion are most frequently attacked, infected host was dissected in Ringer's solution but other ganglia may also be entered. The and nematode development noted. All obser- nematodes do not initiate development with- vations, including photographs and drawings, out first entering a ganglion, and those which were made by the author with fresh material lightly stained with cotton blue. remain in the hemolymph are encapsulated and eventually killed. Results Further studies on the host range of this nematode and selection of ganglia will be The juvenile of F. leipsandra molted once presented by the author and Dr. Gotz, a visitor within the egg before hatching. After hatch- in this laboratory for three months during the ing, the preparasitic stages were extremely autumn of 1967. active and entered the host by direct penetra- The present paper discusses the parasitic tion through the cuticle. Observations indi- development of F. leipsandra and its associa- cated that both glandular secretions and stylet tion to the central nervous system of D. u. action aided penetration. When contact with undecimpunctata. the host was made, the nematode forced its head against the cuticle and began a succes- Materials and Methods sion of stylet movements. Occasionally, a quick movement of the host brushed off the Adult mermithids oviposited in dishes of nematode and viscous material could be seen water. After the eggs hatched, the infective coming from the mouth of the latter. In one juveniles were held at 12 C for future infec- instance after entering the hemocoel, the tion experiments. Since D. balteata does not nematode migrated to the head end of the occur in Northern California, a local species, host and explored the periphery of the sub- D. undecimpunctata undecimpunctata was esophageal ganglion for several seconds. It successfully infected and served as a host for then forced its head through the neural lamella the nematode. Comparisons of this host with and perineurium and slowly entered the gan-

-• 5

Figures 1-6. 1. Two juveniles of F. leipsandra 2 days after infection in one of the protocerebral lobes of a first instar Diabrotica larva, n — nematode. 2. Close-up of nematodes in figure 1 showing the gland cells or stichocytes (g) of 1 specimen and the stylet(s) of another. 3. F. leipsandra in a protocerebral lobe of a first instar Diabrotica larva 4 days after infection, n — nematode. 4. F. leipsandra

Copyright © 2011, The Helminthological Society of Washington OF WASHINGTON, VOLUME 35, NUMBER 2, JULY 1968 163 glion, following the inner contour of the neural in the early parasitic stages, but gradually dis- lamella and eventually coming to rest in the appear and are absent in the postparasitic neuropile. It appears that the cells and fibers juvenile. The cellular structure of the gut is in the ganglion are not broken, but just pushed still obscure, but a distinct rectum and anus aside as the nematode enters. Once inside the are present. The gonad rudiment lies on the ganglion, the parasite remains still, only alter- ventral side of the body just behind the junc- ing its position through subsequent growth. tion of the pharynx and intestine. The nematode may remain within the host Figure 1 shows two nematodes in one of the from 12 to 22 days, depending on the state protocerebral lobes of the host 2 days after of the host and number of parasites present. infection. Although still no significant increase After 22 days, the parasite may reach a size in length, the width was now 0.025 mm and of 4.70 cm and a width of 0.26 mm. This is the stichocytes were prominent (Fig. 2). an enormous change from the average length Little noticeable change occurred after 3 days (0.54 mm) and width (0.018 mm) of the in the host except for an increase in width infective juvenile. (0.034 mm). After 4 days, however, a sig- The size of the parasite after 24 hours in nificant increase in length (0.712 mm) as the host did not differ significantly from that well as width (0.053 mm) produced a notice- of the infective stage (Fig. 7). The well- able enlargement of the infested protocerebral developed stylet leads directly into the phar- lobe of the host (Fig. 3). yngeal tube which runs the length of the After 5 days within the protocerebral lobe pharynx. The anterior portion of the pharynx of a first instar host, the nematode reached a is narrow and consists of a cuticle lined tube length of 0.741 mm (Fig. 4, 8). The stylet surrounded by a layer of nucleated sheath and pharyngeal tube were distinct and the tissue. The prominent nerve ring encircles stichocytes increased greatly in size. Remnants this anterior portion of the pharynx, which of the paired gland reservoirs were still visible. gradually widens to its full width at the level Movement of the crystals within the intestinal of the atrophied gland reservoirs. These latter lumen suggested that the lumen was filled paired oval shaped bodies are what remains with liquid. The cellular structure of the of the two long gland reservoirs of the infec- intestine proper is now evident and the rectum tive juvenile. and anus are still distinct. Cells of the hypo- In the infective stage, these reservoirs are dermal cords are prominent. Increase in filled with globules which are probably en- growth of the parasite after 6 days (L = 1.36 zymes used in softening the host cuticle during mm, W = 0.08 mm) forced it against the penetration. The basal two-thirds of the neural lamella of the protocerebral lobe in a pharynx, including the stichosome (Steiner, second instar host (Fig. 5). 1933) contains 16 gland cells or stichocytes. Most of the parasites broke out of the These cells are arranged in two rows forming ganglion proper on the 7th day, but were still the two subdorsal portions of the pharynx. contained by the neural lamella which pro- The pharynx is attached to the intestine by a tected them from direct contact with the hemo- thin layer of connective tissue, however, this lymph (Fig. 9). The length increased greatly is only a physical, not a functional union and to 2.39 mm and the width to 0.11 mm. The does not persist for long. The intestine proper structure of the stylet was obscure and the is filled with globules, but a lumen in the stichocytes began to elongate. The junction anterior portion contains large crystals of un- between the pharynx and intestine was severed known origin. These crystals are very distinct and the gut grew forward. The gonad rudi- in a protocerebral lobe of a first instar Diabrotica larva 5 days after infection, n — nematode. 5. Greatly enlarged protocerebral lobe of a second instar Diabrotica larva containing a developing juvenile of F. leipsandra 6 days after infection. 11 = nematode. 6. Two specimens of F. leipsandra breaking out of a protocerebral lobe and the subesophageal ganglion, respectively, of a second instar Diabrotica larva 8 days after infection. Note that both specimens are still contained by the neural lamella which sur- rounds the ganglia, n — nematode; p — uninfected protocerebral lobe.

Figure 7. F. leipsandra in the first thoracic ganglion of a first instar Diabrotica 1 day after infection, s — stichosoinal portion of pharynx; g — gland cell or stichocyte; r — gland reservoir; t = pharyngeal tube; n — nerve ring; 1 — stylet; d = gonad; c = crystals in anterior lumen of intestine. merit was still an undifferentiated mass of chocytes. The intestine reached its definitive cells. After 8 days in the ganglia of a second location in relation to the rest of the body and stage host, the parasite reached a length of the gonad rudiment began to differentiate into 2.7 mm (Fig. 6). The neural lamella continued vaginal and ovarial portions (Fig. 10). to stretch, and 011 the 9th day still contained The nematode remained within the pouch the parasite, which had reached a length of for two more days while the neural lamella 5.00 mm and a width of 0.11 mm. The stretched up to 3 mm in length. The mem- pharynx had now assumed a distinct tripartate brane broke on the 12th day after infection structure with the ventral portion containing and the parasites moved directly into the host the pharyngeal tube, and the two subdorsal hemocoel. The period the nematode remained portions each with a row of eight large sti- inside the ganglia proper and the membrane,

Copyright © 2011, The Helminthological Society of Washington OF WASHINGTON, VOLUME 35, NUMBER 2, JULY 1968 165 respectively, varied depending on the rate of may jointly be regarded as the stichosome infection and age of the host. Growth in (Steiner, 1933). The stichocytes are large length continued at a phenomenal rate and clear cells with granular nuclei and are most on the 14th day, the nematode was 34.6 mm prominent during the early stages of para- long and 0.20 mm wide. sitism. They probably play an important role When the nematode emerged from the host, in the nutrition of the nematode, but their in this case on the 22nd day, it had reached exact nature is unknown. In later parasitic a length of 47.0 mm and a width of 0.26 mm. and postparasitic stages the stichosome tissue Although the intake of nourishment probably breaks down and the posterior part of the ceased even before the nematode made its pharynx consists of isolated stichocytes at- exit from the host, sexual differentiation con- tached to the slender pharyngeal tube. tinued up to the postparasitic molts and ovarial The number of stichocytes in F. leipsandra development continued until the nematode was constantly 16 which may be basic for expired. mermithids since Christie (1936) also found 16 in developing juveniles of Agamermis Discussion decaudata and Gotz (1964) recorded 16 in The parasitic development of Filipjevimer- juveniles of Gastromermis rosea. However, mis leipsandru, as with mermithids in general, other workers have found variable numbers; results in an extreme increase in length rather Johnson (1955) recorded from 15-17 in than width as is found in other entomogenous thjdromermis contorta depending on the age nematocles (Poinar, 1965). In one case, the of the juvenile, and Couturier (1963) found length and width increased 87 and 14 fold up to 10 in Tunicamermis melolonthae. A respectively over a 22-day period in the host. bulblike enlargement of the pharynx as Christie No sign of a molt was noted during the para- (1936) reported in juveniles of A. decaudata sitic period. One molt occurred within the was lacking in F. leipsandra. egg just before hatching and the postparasitic The intestine, or trophosome as it is called juvenile molted twice before oviposition. The in its modified form in mermithids, appears sensory papillae were obscure during parasitic initially as a tube filled with globular inclu- development, although the nerve ring and sions. Intestinal cells become distinct by the associated cells were distinct at all stages. 5th day and the intestine rapidly begins to An increase in length did not occur until elongate by the 7th day. The significance of the 4th day after infection. This may be typical the lumen in the anterior portion of the of mermithid development since Christie trophosome with its crystal contents is un- (1936) also remarked that with Agamermis known. decaudata in Melanoplus femur-rubrum, there A minute, nonfunctional anus and rectum was little change in length of the parasite are present throughout most of the parasitic during the first three days in the host. How- development, but only a vestigial anus re- ever, internal development began immediately mains in the postparasitic juvenile. The cells after entering the host. This was especially forming the hypodermal chords are distinct evident with the stichocytes of the pharynx. during all stages of development and some Although Hyman (1951) considered the appear to be binucleate or in process of pharynx and stichosome as separate structures division. Although cells of the gonad rudiment in the Mermithidae, it is obvious that they are begin dividing early in the development, sexual part of the same organ and the stichosome is differentiation was completed only after the regarded here as a highly specialized part of parasites left the host. No sign of an excretory the pharynx. During parasitic development, the basal pore was seen during this study. portion of the pharynx differentiates into three Most nematodes which develop exclusively longitudinal divisions. The ventral part con- in invertebrates are not known to invade tains the remainder of the pharyngeal tube specific tissues of the host, in contrast to the which ends blindly at the base of the organ. filarioid and spiruroid nematodes which utilize The two subclorsal portions each contain a row invertebrates as intermediate hosts. However, of eight gland cells or stichocytes and they even in the latter two groups, a record of

29p Figure 8. Developing juvenile of F. leipsandra 5 days after infection in a protocerebral lobe of a first instar Diabrotica larva, h — hypodermal cell.

Figure 9. Developing juvenile of F. leipsandra which has just broken out ot an abdominal ganglion of a second instar Diabrotica larva 7 days after infection. It is retained by the thin neural lamella (m). development in the nervous system of the the tachinid Gonia and noctuid larvae. Upon host could not be found, although Lavoipierre entering the body cavity, the parasites were (1958) mentions once finding a juvenile of encapsulated unless they entered the supra- Loa loa partly embedded in the brain of the esophageal ganglion of the host. After remain- adult fly, Chrysops silicea. ing there for a few days, they re-entered the Thus, it was surprising to discover a mermi- body cavity and were not attacked by blood thicl which showed an affinity to the ganglia cells. Other mermithids occurred in a sheath of an insect host. Still, the affinity of mermi- within their hosts, but whether this protected thid nematodes to specific tissues of insects them from an encapsulation reaction is not may be more widespread than imagined since known (Rennie, 1925; Couturier, 1963). Hagan and Hoopingarner (in press) found the The "normal" host of an entomogenous early stages of an undetermined mermithid nematode is usually considered one in which in the brain lobes of larvae of Aedes stimulant. the association supposedly has been of such One of the most obvious reasons for F. long standing that the parasite is now "ac- leipsandra to enter the ganglia of Diabrotica cepted" with a minimum of host reaction would be to escape encapsulation and sub- (Poinar, in press). The behavior of F. leipsan- sequent death. It is interesting that the para- dra could represent an intermediate step in site is not attacked by blood cells when it adapting to a parasitic development free in the breaks out of the ganglion and neural lamella hemolymph of Diabrotica, or it could represent 9-12 days after infection. Whether the nema- the end of a specialized line of selection, tode is now too large or has acquired some enabling the parasite to avoid the defense attribute which makes it "acceptable" to the reaction of various hosts. host is not known. A similar situation was If the latter is true, and artificial infection reported by Strickland (1930) working with studies with various hosts suggest this, then

Figure 10. F. leipsandra 9 days in the host and held within the neural lamella (m) oi the protocerebral lobe of a second instar Diabrotica larva, i — intestine or trophosome; g = gland cell or stichocyte; t = pharyngeal tube. the nematode may not be hindered by the short period for such a tremendous increase in defense reaction of new "unusual" hosts and length of the nematode. would be able to expand its host range. In fact, Diabrotica may be a relatively "recent" Acknowledgments host since it could easily destroy the parasites The author expresses his sincere thanks to by encapsulation if they did not enter the Dr. Cuthbert for providing the original supply ganglia. Of course, the possibility that the of nematodes, to Mr. R. A. Corey of the Shell ganglia furnish developing juveniles of F. Development Company, Modesto, California, leipsandra some subtle nutritional factor neces- for his help in rearing methods of Diabrotica sary for development is still open for investiga- and to Mr. Carl Reiner who patiently main- tion. tained colonies of D. n. itndecimpunctata for Summary experimental use at the University. Grateful appreciation is also extended to Mr. Gerard First instar larvae of Diabrotica it. undecim- Thomas who assisted in technical aspects of punctata were infected in the laboratory with this study. juveniles of the mermithid nematode, Filip- jevimermis leipsandra. The infective stage Literature Cited nematocles penetrated directly into the hemo- Christie, J. R. 1936. Life history of Agamermis coel and then entered one of the ganglia of decaudata, a nematode parasite of grasshop- the host. The protocerebral lobes and sub- pers and other insects. Jour. Agr. Res. 52: esophageal ganglion were most frequently 161-198. attacked. The parasites developed initially Couturier, A. 1963. Recherches sur des Mer- within the ganglia, then after outgrowing the mithidae, nematodes parasites du hanneton commun (Melolontha melolontha L. Coleopt. latter, were retained by the neural lamella, Scarab.). Ann. Epiphyties 14: 203-267. which stretched a considerable distance before Cuthbert, Jr., F. P. In press. Bionomics of a finally breaking and liberating the nematodes mermithid (Nematoda) parasite of soil-in- in the host hemolymph. habiting larvae of certain chrysomelids (Cole- The growth and internal development of optera). J. Invert. Path. Gotz, P. 1964. Der Einfluss unterschiedlicher the parasite were studied by dissecting infected Befallsbedingungen auf die Mermithogene hosts at regular intervals. Parasitic develop- Intersexualitat von Chironomus (Dipt.) Z. f. ment lasted from 12 to 22 days, a relatively Parasitenkunde 24: 484-545.

Hagan, H., and R. Hoopingarner. In press. vien) (Sphaerulariidae: Aphelenchoidea), a Mermithid nematode parasitism of Aedes nematode parasite of sciarid flies (Sciariclae: stimulans (Walker) (Diptera: Culicidae) Diptera). Parasitology 55: 559-569. from Ingham County, Michigan. Ann. Ent. . In press. Arthropod immunity to worms. Soc. Amer. In: Jackson, G., and R. Herman, Immunity Hyinan, L. H. 1951. The Invertebrates: Acan- to parasitic animals. Appleton-Century-Crofts, thocephala, Aschelminthes, and Entoprocta, New York. The pseudocoelomate bilateria. Vol. III. Mc- -, and H. E. Welch. In press. A new Graw-Hill, New York. 572 pp. nematode, Filipjevimermis leipsandra sp. nov. Johnson, A. A. 1955. Life history studies on (Mermithidae) parasitic in chrysomelid larvae Hydromermis contorta (Kohn), a nematode (Coleoptera). J. Invert. Path. parasite of Chironomiis plumostis (L.). Ph.D. Rennie, J. 1925. A mermithid parasite of Tipula thesis Univ. 111. 92 pp. paludosa Meigen. Proc. Royal Phys. Soc. Lavoipierre, M. J. J. 1958. Studies on the host- Edinb., 21: 1-3. parasite relationships of filarial nematodes and Rimando, L. C., R. A. Corey, and Yun-Pei Sun. their arthropod hosts. 1. The sites of develop- 1965. Mass rearing of the western spotted ment and the migration of Loa loa in Chnj- cucumber beetle. J. Econ. Ent. 59: 230-231. sops silacea, the escape of the infective forms Steiner, G. 1933. Some morphological and phys- from the head of the fly, and the effect of the iological characters of the mermithids in their worm on its insect host. Ann. Trop. Med. relationship to parasitism. J. Parasit. 19: Parasit. 52: 103-121. 249-250. Poinar, Jr., G. O. 1965. The bionomics and Strickland, E. H. 1930. Phagocytosis of inter- parasitic development of Tripius sciarae (Bo- nal insect parasites. Nature. London, 126: 95.

A Revision of the Genus Rotylenchulus Liiiford and Oliveira, 1940 (Nematoda: Tylenchiclae)1 D. R. DASGUPTA, D. J. RASKI, AND S. A. SHER Department of Nematology, University of California

Species of the genus Rotylenchulus have The genus Rotylenchulus was proposed by been one of the most misidentified groups of Linford and Oliveira in 1940 when they de- all the tylenchs. As evidence, nematodes scribed R. reniformis. Yokoo and Tanaka belonging to this genus have been described (1954) described Tetylenchus nicotiana from in at least four different genera. Also this Japan which was subsequently transferred to genus has been variously assigned to three the genus Rotylenchulus by Baker (1962). families: Tylenchidae (Linford and Oliveira, Three other species (Helicotylenchus eli- 1940; Thome, 1949, 1961; Allen and Sher, sensis Carvalho, 1957, 1959; Spirotylenchus 1967); Heteroderidae (Chitwood and Chit- queirozi Lordello and Cesnik, 1958, and Heli- wood, 1950; Skarbilovich, 1960); Hoplolaim- cotylenchus parvus Williams, 1960) were idae (Hopper and Cairns, 1959; Goodey, transferred to the genus Rotylenchulus by Sher 1963; Husain and Khan, 1967) and five dif- (1961). In 1960 Das proposed a new genus, ferent subfamilies: Nacobbinae (Hopper and Leiperotylenchus, which he considered to be Cairns, 1959; Goodey, 1963); Pratylenchinae closely related to Tylenchus and Ditylenchus. (Thome, 1949, 1961; Baker, 1962); Tylen- However, the position of the dorsal gland chulinae (Skarbilovich, 1960); Hoplolaiminae orifice and characters of male tail indicated a (Loof and Oostenbrink, 1962); Rotylenchul- close relationship with Rotylenchulus. Indeed, inae (Husain and Khan, 1967; Allen and Sher, Loof and Oostenbrink (1962) transferred 1967). Leiperotylenchus leiperi to the genus Roty- lenchulus. Goodey (1963) synonymized eli- i A part of the thesis submitted by senior author in par- sensis, parvus, leiperi and queirozi with R. tial fulfillment of the requirements for the Ph.D. degree, University of California, Davis. reniformis. Husain and Khan (1965) described

R. stakmani from India. Swarup, et al. (1967) land (USDANC, Beltsville); Nematology De- considered it a synonym of R. reniformis. partment, Rothamsted Experimental Station, The kidney-shaped mature females from Harpenden, England (NDRES, England); which the common name, reniform nematodes, Plantenziektenkundige Dienst, Wageningen, was derived, are described only in two species, The Netherlands (PD, The Netherlands). R. reniformis and R. borcalis. The descriptions of other species were based on immature Morphology of Rotylenchulus females and males. Rotylenchulus species are characterized by Representatives of the genus Rotylenchulus sexual dimorphism, mature females being have world-wide distribution and their host swollen to kidney shape and males vermiform records are numerous and diverse (Linford with not as well developed stylet and esoph- and Yap, 1940; Peacock, 1956; Martin, 1955). agus. Although detailed morphology of Their general occurrence and economically mature females is obscure, the overall shape great potential as plant pathogens make it and that part of the body posterior to the especially important that the taxonomy of this vulva provide useful taxonomic characters. group be soundly established to assure accu- Fixed specimens of males, immature females rate identification of species. For this purpose and larvae assume a spiral to open C shape this study has been carried out. upon fixation. INCISURES: Immature females, males and Materials and Methods larvae all have four incisures in the lateral Over 3,500 permanently mounted specimens field. Posteriorly on males these incisures of Rotylenchulus from more than thirty coun- extend varying distances from slightly anterior tries covering most parts of the world have to the cloacal opening to about midway on been assembled for this generic revision. Type the tail. The most ventral line always ends at specimens for most of the nominal species of the level of or anterior to the cloaca and on this genus were obtained. the caudal alae except in R. macrodoratus For immature females two measurements of where that ventral line appears to merge with the esophagus, b and b' (Sher, 1963), are the edge of the caudal alae. The lateral field included in the de Man formula. Also included of mature females was not visible. in these measurements are: o (Perry, Darling PHASMIDS: These are porelike and located and Thome, 1959); c' (Sher, 1986) and h slightly anterior to the middle of the tail. (length of hyaline portion of tail in microns). EXCRETORY PORE: The excretory pore is The measurements given in parenthesis in located posterior to the median bulb in all descriptions of the holotype, allotype and neo- species. type refer to population range. HEMIZONID: The hemizonid is two to three An immature female was designated as annules long and immediately anterior to the holotype for R. parvus and mature females as excretory pore. holotype for R. borealis and neotype for R. LIP REGION: The lip region is continuous, nicotiana. The immature female is used as not set off and varies in shape and annulation. the holotype for the new species described Most species have either low and rounded lip here because many morphological characters region (R. parvus) or higher and conoid in are more easily seen in these than in swollen shape (R. macrodoratus n. sp.). Some species females. The swollen female is more difficult appear to be intermediate between these two to collect, indeed is unknown for most species categories (R. reniformis1). despite diligent search for them. Immature females of some species have Type specimens are being deposited in five three or more distinct annules (R. reniformis). permanent institutional collections -which are In other species there is evidence of fine indicated by the following abbreviations: ampliations, but these are difficult to count University of California Nematode Collection, (R. parvus). There is still another group Davis (UCNC, Davis); University of Califor- where annulation is entirely lacking or the nia Nematode Collection, Riverside (UCNC, annules are too fine to resolve (R. leptus n. Riverside); United States Department of Agri- sp.). culture Nematode Collection, Beltsville, Mary- The shape and annulation of the lip region

Copyright © 2011, The Helminthological Society of Washington OF WASHINGTON, VOLUME 35, NUMBER 2, JULY 1968 171 of larvae in all species are similar to the im- agus in males is much reduced with almost mature females except R. anamictus n. sp. no evidence of the median bulb, valve and where the larvae have a conoid lip region in lumen. The posterior part of the male esoph- contrast to the low and rounded lip region of agus overlaps the intestine laterally and ven- immature females. Males in general have a trally, usually more ventrally. rounded lip region. The esophagus of the second-stage larva is The labial framework of immature females not reduced like that of males. Often it is is conspicuous. The cheilorhabdions are obscure in fixed specimens but when seen it thickened anteriorly, extend laterally and in is usually similar to the immature female. some species are thickened at both ends. The However, the posterior part of the esophagus basal ring of the cephalic framework has the of larvae is often asymmetrical. same thickness as that of the cheilorhabdions REPRODUCTIVE SYSTEM: Rotijlenchulus fe- and arches posteriorly in a characteristic man- males have didelphic, amphidelphic ovaries. ner. The cephalic framework in males is The ovaries of immature females are very weaker than that in immature females. small and undeveloped. Each ovary forms a STYLET: Total length of the stylet and size double flexure near the distal end (Fig. 4, B) and shape of the stylet knobs provide taxo- which is unique among tylenchs. The details nomically important characters. Stylet length of mature female reproductive systems could in immature females ranges from 10-26 M. not be traced in their entirety due to the The stylet knobs slope backward except in densely-packed granules and much coiling of R. macrodoratiis n. sp. (Fig. 10, A and D) the ovaries. One of the useful characters in where the knobs are anchor-shaped in im- distinguishing species is the postmedian posi- mature females and in larvae. The male stylet tion of the vulva. The value of V usually falls is smaller and less developed than the cor- between 58-66 except for R. reniformis and responding female stylet and knobs are much R. anamictus n. sp. where it is more posteriorly reduced. located. Lips of the vulva of mature females DORSAL GLAND ORIFICE: One of the impor- usually protrude to varying degrees beyond tant characters of Rotijlenchulus is the location the body contour. The transverse slit of the of the dorsal esophageal gland orifice. Its vulva widens about halfway across the body position varies from 0.56-1.90 times the stylet assuming a funnel shape. The uterus in mature length posterior to the stylet knobs and is females is distinctly convoluted. Nakasono diagnostic for different species. (1966) reported recognizable sperms and ESOPHAGUS: The esophagus of immature spermatheca in egg-laying females in this females is long with a well-developed median genus. These structures have been observed bulb and valve, distinct long and narrow isth- in some species and the presence or absence mus, and lobelike glandular region. The pos- of sperms and spermatheca might prove to terior region of the esophagus is visually hemi- be a useful taxonomic character. Unfortu- spherical in shape but there is occasional intra- nately, these structures usually cannot be population variation in this character. The observed clue to the coiling of the ovaries. glandular region of the esophagus overlaps the The male reproductive system is monorchic intestine laterally and ventrally and is pre- and outstretched. Gubernaculum and spicules dominantly lateral. The lumen of the esoph- are well-developed. Caudal alae are much agus curves ventrally in the form of an open reduced, adanal and annulated. loop at the dorsal gland orifice. The tissue at the junction of esophageal lumen with the TAIL: Generally the tail shape of the larvae intestine appears to be more dense than the is rounded whereas it is more pointed in the surrounding area. No distinct valve could be immature females. There is considerable intra- found at the junction. The esophageal gland specific variation in tail shape of immature nuclei are three in number. females which limits the diagnostic value of The esophagus of mature females is irregular this character. However, length of tail, ratio in shape. The median bulb of mature females of anal body diameter to tail length (c') and is spheroid, more than twice the size of the length of the hyaline portion of the tail (h) median bulb of immature females. The esoph- are important for distinguishing species. The

Figure 1. Rotylenchulus reniformis. A—Immature female, face view; B—Immature female; C—Larva; D—Male; E—Male, tail; F-H—Immature female, tail. body content of the larvae reaches almost to Systematics the terminus and there is virtually no indica- Subfamily Rotylenchulinae Husain tion of a hyaline area. and Khan, 1967 TKe distal portion o£ the tail of mature females in some species is set off into a finger- DIAGNOSIS EMENDED: Tylenchidae. There like appendix. In other species the distal is only one genus in this subfamily and the portion of the tail is in the shape of a short characters of that genus, Rotylenchulus, con- spike. Still in other species the tail is bluntly stitute the characters of the subfamily. rounded with no indication of an appendix or Type genus Rotylenchulus Linford and projection. Oliveira, 1940.

Genus Rotylenchuhts population originally from type locality, reared on cowpea): L — 0.38-0.52 mm; width at Rotylenchulus Linford and Oliveira, 1940, vulva = 0.10-0.14 mm; a = 4-5; V = 68-73; pp. 35-39; Thorne, 1949, pp. 41-42; Thome, length of swollen portion of body plus tail = 1981, pp. 244-245; Goodey, 1983, pp. 87-88. 0.19-0.47 mm; stylet = ?; median esophageal Spirotylenchus Lordello and Cesnik, 1958, bulb diameter = 18-22 M. p. 161; Sher, 1961, p. 159. (26 immature 5 topotypes): L = 0.34- Leiperotylenchus Das, 1960, pp. 560-562. 0.42 mm; a = 22-27; b = 3.6-4.3; b' = 2.4- DIAGNOSIS EMENDED: Rotylenchulinae. Ma- 3.5; c = 14-17; c' = 2.6-3.4; h = 4-8; V = ture female swollen, kidney-shaped, male 68-73; stylet = 16-18 n; o = 81-106. vermiform. Lip region of immature female (10 $ topotypes): L = 0.38-0.43 mm; a = not set off, cephalic framework conspicuous. 24-29; b' = 2.8-4.8; c = 12-17; h = 4-8; Dorsal gland orifice more than one-half stylet T = 35-45; stylet = 12-15 p.; gubemaculum length posterior to base of stylet knobs. = 7-9 fi; spicules = 19-23 A*. Esophagus long with narrow isthmus, glands (10 larvae topotypes): L = 0.35-0.41 mm; overlapping intestine laterally, ventrally, more a = 20-24; b' = 3.5-4.1; c = 12-16; stylet = often laterally. Vulva post-median, ovaries 13-15 n. diclelphic, amphiclelphic with two flexures in IMMATURE FEMALE (neotype): L = 0.40 immature female, highly convoluted in mature mm; a = 24; b = 3.8; b' = 2.8; c = 16; c' = female. Female tail usually more than twice 2.9; h = 6; V = 72; stylet = 16 ^ o - 82. anal body diameter. Larval tail more rounded Lateral field slightly less than quarter of body than tail of immature female. Phasmicl pore- width. Body in open C shape. Lip region like, anterior to middle of tail. Male with weak high, conoid, rounded, not set off from body, stylet and stylet knobs, reduced esophagus, with 5 annules (4-6). Stylet knobs rounded, indistinct median bulb and valve. Caudal alae slope backward. Excretory pore 77 M from adanal. Lateral field of male, immature female anterior end (73-90 M) at the level of posterior and larvae with four incisures, nonaerolatecl. end of isthmus below hemizonid. Metacorpus Eggs deposited in gelatinous matrix. elongate, oval and valve massive 4 A* (4-6 M) Type species Rotylenchulus reniformis Lin- long. Esophageal glands overlap intestine ford and Oliveira, 1940. laterally and ventrally, the longest overlap Rotylenchulus reniformis Linford ventral. Tail 24 ,u (19-26 A*) long, conoid, terminus rounded. and Oliveira MATURE FEMALES: Anterior part of body (Figs. 1, A-H; 2, A-H) contour irregular. Body curved ventrally. syn. Tetylenchus nicotiana Yokoo and Tanaka, Esophagus irregular in shape. Metacorpus 1954. spherical, with large valve. Vulva with raised Rotylenchnhis nicotiana (Yokoo and Tan- lips. Vagina funnel shaped. Spermatheca aka, 1954) Baker, 1962. New Syn- spherical to irregular in shape containing many onymy. sperms. Body swollen almost to terminus then Rotylenchus elisensis Carvalho, 1957. round to grossly hemispherical shape. Spike- Helicotylenchus elisensis (Carvalho, 1957) like process of tail short (5-8 M). Carvalho, 1959. MALES: More slender than immature females. Lip region high, rounded, cephalic Spirotylenchus queirozi Lordello and Ces- sclerotization and stylet weaker than immature nik, 1958. females. Esophagus reduced, lumen usually Rotylenchulus queirozi (Lordello and Ces- difficult to see. Caudal alae adanal and rudi- nik, 1958) Sher, 1961. mentary. Leiperotylenchus leiperi Das, 1960. LARVAE: Resembling immature females. Rotylenchulus leiperi (Das, 1960) Loot Posterior part of esophagus asymmetrical. and Oostenbrink, 1962. Esophageal glands overlap intestine laterally Rotylenchulus stakmani Husain and Khan, and ventrally. Dorsal gland orifice about one 1965. stylet length posterior to stylet knob base. MEASUREMENTS: (16 mature 9, Hawaii, Genital primorclium four-celled, at the same

Figure 2. Rotylenchulus reniformis. A—Immature female, anterior end; B—Male, anterior end; C— Larva, anterior end; D—Larva, tail; E—Mature female; F—Immature female (Ratnapura, Ceylon), tail; G-H—Mature female, tail. position as vulva. Tail more rounded than 928, UCNC, Davis; 3 mature 9, 7 immature immature females. 9, 1 $, slides numbered R1-R3, UCNC, NEOTYPE-. Immature female, collected by Riverside; 2 mature 2, 9 immature ?, 1 $, J. Radewald, 24 February 1961, slide num- slides numbered T587-T590, USDANC, Belts- bered 900, UCNC, Davis. ville; 2 mature 9, 6 immature 9,1 $, slides TOPOTYPES : 23 mature 9, 75 immature $ , numbered 60/1/10-60/1/13, NDRES, En- 20 6', 21 larvae same data as neotype, dis- gland; 2 mature 9, 6 immature 9,1 $, slides tributed as follows: 12 mature 9 , 47 immature numbered b66-69, PD, The Netherlands. 9, 16 $, 21 larvae on slides numbered 901- NEOTYPE HABITAT AND LOCALITY: Soil

Copyright © 2011, The Helminthological Society of Washington OF WASHINGTON, VOLUME 35, NUMBER 2, JULY 1968 175 around pineapple (Anamis sativus), Dole 0.41-0.49 mm; a = 24-28; b = 3.9-4.5; b' = Corporation, Field 4522, Block C-9, County 3.0-3.7; c = 11-14; c' = 4.0-5.0; h = 5-9; of Oahu, Hawaii, USA. V = 68-70; stylet = 18-21 AS o = 75-105. TOPOTYPES OF R. elisensis (4 immature 5 ): 13 immature ? (San Miguel, Philippines): L = 0.38-0.43 mm; a = 27-28; b = 3.7-3.9; L = 0.42-0.44 mm; a = 27-32; b = 3.6-4.9; b' = 2.8-2.9; c = 13-14; c' = 2.9-3.3; h = b' = 2.9-3,5; c = 11-13; c' = 4.1-4.6; h = 8-9; V = 69-70; stylet = 17-18 ^ o = 79-81. 5-10; V = 67-70; stylet = 18-20 AS o = 75- TOPOTYPES OF R. elisensis (4 $ ): L = 0.43- 100. 0.47 mm; a = 28-31; b' = 3.9-4.1; c = 15-17; 4 mature 2 (Florida, USA): L = 0.40- h = 7-8; T = ?; stylet = 12-13 ^ guber- 0.64 mm; width at vulva = 0.08-0.14 mm; naculum = 8-9 AS spicules = 20-22 A*. a = 4-5; V = 66-72; length of the swollen PARATYPES OF R. stakmani (5 5 ): L = portion plus tail — 0.26-0.46 mm; stylet = 0.34-0.39 mm; a = 24-26; b = 3.6-3.7; b' = 14-15 M; median esophageal bulb diameter = 2.7-3.5; c = 14-16; c' = 2.8-3.0; h = 5-7; 19-24 A*. V = 70-72; stylet = 16-18 M; o = 100. 10 mature 5 (Hawaii, USA): L = 0.44- PARATYPES OF R. stakmani (3 5): L = 0.57 mm; width at vulva = 0.11-0.14 mm; 0.38-41 mm; a = 28-29; b' = 3.8-4.2; c = a = 4-5; V = 67-70; length of the swollen 14-15; h = 4-6; T = ?; stylet = 14-15 M; portion plus tail = 0.25-0.40 mm; stylet = ?; gubernaculum = 8-9 AS spicules = 18-22 /*. median esophageal bulb diameter = 19—20 A1; TOPOTYPES OF R. nicotiana (8 immature 5 ): eggs = 60-94 ^ x 37-44 n. L = 0.36-0.44 mm; a = 24-31; b = 3.5-3.9; Mature females of R. reniformis also have b' = 2.7-3.1; c = 12-14; c' = 3.1-3.8; h = been identified from the following habitats and 6-12; V = 69-72; stylet = 17-19 ft; o = 75- localities: pineapple root, Columbia; soya (Gly- 90. cine max] root, Gold Coast; tomato (Lijcoper- TOPOTYPES OF R. leiperi (8 immature 9): sicon esculentum) root, Japan; tomato root, L - 0.35-0.41 mm; a = 23-26; b = 3.6-4.0; Nigeria; Vigna sesquipedales root, Philippines; b' = 2,5-3.5; c = 14-15; c' = 2.8-3.0; h = Yucca gloriosa root, San Bernardino, USA. 5-6; V = 70-73; stylet = 16-18 AS o = 81-98. DIAGNOSIS: R. reniformis can be identified TOPOTYPES OF R. leiperi (4 $ ): L = 0.39- by its high conoid and rounded lip region of 0.43 mm; a = 26-28; b' = 3.6-4.0; c = the immature females, more posterior position 13-14; h = 4-6; T = 30-32; stylet = 12-13 of the vulva in the immature and mature AS gubernaculum = 7-8 AS spicules = 18-21 A*. females, hemispherical shape of body beyond 22 immature $ (Ratnapura, collection no. anus ending in a short spikelike process in 1, Ceylon): L = 0.43-0.53 mm; a = 27-32; mature females. b = 3.6-4.4; b' = 3.0-3.5; c = 11-15; c' = R. reniformis is the most widely distributed 3.9-5.0; h = 8-12; V = 68-71; stylet = 20- species of the genus. Immature females and 22 AS o = 70-100. males have been examined and identified from 8 immature $ (Ratnapura, collection no. 2, the following habitats and localities: tomato Ceylon): L = 0.33-0.46 mm; a = 21-33; soil, Sao Paulo, Brazil; cocoa (Theobroma b = 3.7-3.9; b' = 2.9-3.5; c = 13-15; c' = cacao) soil, Kandy, Ceylon; tea (Thea sinen- 2.9-3.7; h = 5-10; V = 69-72; stylet = 16- sis) soil, Ratnapura, Ceylon; coconut (Cocos 20 AS o = 86-100. nucifera) and tea soil, Ceylon; jungle soil, 6 ci (Ratnapura, collection no. 2, Ceylon): Kahawatta, Ceylon; malanga (Arum sp.) soil, L = 0.35-0.46 mm; a = 25-34; b' = 3.4-4.2; papaya (Carica papaya) soil, sweet potato c = 14-16; h = 5-6; T = 29-40; stylet = (Ipomoea batatas) soil, Vinales, Cuba; banana 12-14,«; gubernaculum = 6-8 AS spicules = (Musa sp.) soil, Kerala, India; rose (Rosa sp.) 18-21 M. soil, Delhi, India; potato (Solanum tuberosum) 15 immature 2 (Sherwood forest, Jamaica): soil, Aligarh, India; Sorghum, sp. soil, Hydera- L = 0.43-0.53 mm; a = 27-32; b = 3.6-4.7; bad, India; soil around unknown plant, Java, b' = 2.8-3.5; c = 13-17; c' =3.8-4.6; h = Indonesia; tobacco (Nicotiana sp.) soil, Java, 4-6; V = 67-72; stylet = 17-21 AS o = 76- Indonesia; tea soil, Java, Indonesia; coffee 105. (Coffea sp.) soil, Indonesia; avocado (Persea 6 immature $ (Kingston, Jamaica): L = americana) soil, Iraq; sugar cane (Saccharum

Copyright © 2011, The Helminthological Society of Washington 176 PROCEEDINGS OF THE HELMINTHOLOGICAL SOCIETY officinarum) soil, Basrah, Iraq; tobacco soil, The illustration and description of Leipero- Kogoshima, Japan; burdock (Arctium lappa] tylenchus leiperi published by Das (I960) soil, Saitama, Japan; banana soil, Darlingford shows a monodelphic ovary, posterior bulb in and Kingston, Jamaica; Citrus sp. soil, Kings- the esophagus and six lines in the lateral field ton, Jamaica; fumigation plot soil, Spanish which do not conform with the generic charac- Town, Jamaica; sugar cane soil, Jamaica; terization of Rotylenchulus. However, the papaya soil, Ganta, Liberia; tomato soil, Cul- male and the position of the dorsal gland ican, Mexico; banana soil, Benin Province, orifice as well as other measurements of the Nigeria; banana soil, Abeokuta Province, Ni- female indicate a relationship with R. reni- geria; general bush soil, Ondo Province, formis. Jones (as mentioned by Goodey, Nigeria; garden eggplant (Solanum melon- 1963) as well as Raski (personal communica- gena) plant soil, Nigeria; unknown grass soil, tion) observed type specimens in Hyderabad. Ibadan Province, Nigeria; gravel field soil, They considered the specimens to be immature Ibaclan Province, Nigeria; corn (Zea mays] females of Rotylenchulus. Five collections soil, Adamawa Province, Nigeria; corn soil, from soil taken in 1963 from the type locality Benin Province, Nigeria; corn soil, Enugu also were examined. None of these samples Province, Nigeria; lettuce (Lactuca sativa) contained nematodes as described by Das. soil, Ondo Province, Nigeria; Hyparrhenia rufa Two of these samples contained immature soil, Ibadan Province, Nigeria; unknown tree females and males of R. reniformis. Goodey soil, Ibaclan Province, Nigeria; water leaf (1963) synonymized L. leiperi with R. reni- (Talinum triangulate) soil, Adamawa Province, formis. The additional evidence above sup- Nigeria; soil around unknown plant, Dacca, ports the synonymy. Pakistan; soil around unknown plant, San The differences judged by Husain and Khan Miguel, Philippines; Vigna sesquipedales soil, (1965) to represent a different species, R. Laguna, Philippines; unknown weed, soil, stakmani, are: dorsal gland orifice less than Somaliland; C rot on sp. soil, Florida, USA; one stylet length behind the base of stylet cotton soil (originally from a field population knob, absence of caudal alae in males and of Puerto Rico), North Carolina, USA; Yucca shorter tail of males. Examination of paratype gloriosa, San Bernardino, USA; Texas, USA; specimens sent by these authors revealed all tomato soil, Venezuela; naranja (Citrus sp.) the males have distinct but rudimentary, soil, Venezuela; and weed soil, Western adanal alae which is typical of all known Panama. species of Rotijlenchulus. The dorsal gland Sher (1961) considered H. elisensis a syn- orifice and male tail show no differences when onym of R. reniformis. The opportunity to compared with specimens of R. reniformis study the specimen supplied by Carvalho from the type locality. Therefore, the place- indicated that they are conspecific with im- ment of R. stakmani in synonymy with R. mature females of R. reniformis and the syn- reniformis by Swarup et al. (1967) is con- onymy is confirmed. firmed. Sher (1961) synonymized the genus Spiroty- Specimens from Ceylon, Jamaica, and Japan lenchus Lordello and Cesnik, 1958, with Roty- present a complex of morphological characters lenchulus and by that action S. queirozi be- which are especially difficult to resolve. These came Rotylenchulus queirozi (Lordello and widely variable characters include the mea- Cesnik, 1958) Sher, 1961. Goodey (1963) surements of L, stylet length, c', h and o. In synonymized R. queirozi with R. reniformis. addition the presence or absence of males also The only character of R. queirozi which falls must be considered. outside the range of R. reniformis is the short At one extreme is a population from Ceylon male tail of queirozi (c = 18.0-18.2). This (Ratnapura, collection no. 1) in which im- variance is not considered to be great enough mature females have values for L, stylet length, to represent a distinct species. In all other c' and h which are distinctly greater and do respects R. queirozi closely resembles R. reni- not overlap the measurements of specimens formis in dimensions and descriptions. There- from the type locality. Also no males could fore, the synonymy of Goodey is considered be found in the Ceylonese collection. How- valid. ever, a second collection from the same localitv

Copyright © 2011, The Helminthological Society of Washington OF WASHINGTON, VOLUME 35, NUMBER 2, JULY 1968 177 and habitat in Ceylon (Ratnapura, collection species. Further research should give a better no. 2) had males in equal frequency to the understanding of the importance of males in females. Also the immature females had the this complex group of nematodes and also values of L, stylet length and h which were serve as a more reliable basis for distinguishing intermediate and bridged the measurements different species that may exist in the complex. of Ratnapura, collection no. 1 and type speci- In the meantime on the basis of the additional mens of R. reniformis. Furthermore popula- evidence presented here it is judged wiser to tions from Jamaica and Philippines give further consider these collections as representing a evidence of values which bridge and overlap single species, the most variable in the genus. the differences in L, stylet length and h. Recently Nakasono and Ichinohe (1967) Rotylenchulus macrosomus n. sp. reclescribed R. nicotiana diagnosing it from (Fig. 3, A-G) R. reniformis on (1) absence of males, (2) MEASUREMENTS (21 immature 2 para- lower value of o (81.3 ± 2.3% as against 91.1 types): L = 0.52-0.64 mm; a = 30-38; b = ± 3.5% tor reniformis) and (3) a higher value 3.8-5.7; b' = 2.9-4.3; c = 12-16; c' = 3.7- of h (8.5 as against 6 for reniformis). There 5.0; h = 13-18; V = 63-68; stylet = 18-22 ^ is also evidence for a difference in the host o = 139-188. preference between reniformis and nicotiana. (21 $ paratypes): L = 0.50-0.68 mm; However, the range of o (60.0-94.2) and h a = 30-41; b' = 3.7-5.7; c = 12-16; h = 15- (6-12) in nicotiana overlaps considerably the 23; T = 20-33; gubernaculum = 8-10 ^ values of each in reniformis. Also there are a spicules = 21-24 M; stylet = 13-16 n. number of collections from other parts of the (7 larvae paratypes): L — 0.49-0.68 mm; world in which the o and h values for the a = 25-35; b' = 4.3-5.3; c = 13-17; stylet = immature females are similar to reniformis and 14-16 M. nicotiana and males are frequent. These col- IMMATURE FEMALE (Holotype): L = 0.64 lections include San Bernardino, California mm; a = 37; b = 4.7; b' = 3.7; c = 14; c' = (h = 5-9, o = 77-94); North Carolina (h = 3.9; h = 17; V = 64; stylet = 18 /*; o = 143. 4-6, o = 75-88); Western Panama (h = 6- Lip region bluntly conoid, finely annulated. 10, o = 80-92); Mexico (h = 5-10, o = Stylet knobs sloping backwards. Excretory 83-100); and Brazil (h = 8-9, o = 79-81). pore posterior to hemizonid, 106 n from ante- These data are suggestive of a polymorphic rior end (90-106 M). Metacorpus oblong, species with many local variations. The same valve 5 M (5-6 M) long. Esophageal glands could be true of host preferences for which overlapping the intestine laterally and ven- there are many examples of variations between trally, more ventrally. Tail 39 // (35-43 M) populations of the same species. Furthermore long, annulation around the terminus prom- the immature females in these many collections inent; terminus bluntly rounded. resemble each other very closely in almost all MALE (Allotype): L = 0,56 mm; a = 37; other specific characters. The close relation- b' = 4.4; c = 14; h = 15; T = 22; stylet = ship of all these specimens is still further 13 n; gubernaculum = 9 /j.; spicules = 22 M. emphasized by the close similarity of the adult Sclerotization of the lip region weak. Stylet females. Indeed where adult females are and esophagus reduced. Metacorpus and valve known for other species of Rotylcnchnlus they indistinct. show marked differences in size and shape LARVAE: Similar to immature females. Tail (Fig. 12, A-J). more rounded than immature females. There remains the significance of the per- HOLOTYPE: Immature female, collected by G. Minz, July 4, 1960, slide numbered 1010, plexing presence or absence of males and their UCNC, Davis. role in reproduction. Unfortunately there is ALLOTYPE: Male, same data as holotype, not sufficient information on which to base slide numbered 1011, UCNC, Davis. firm conclusions. When judged from the PARATYPES: 48 immature ? , 36 $ ,7 larvae, generic level the similarities are more persua- same data as holotype, distributed as follows: sive for a variable species or a group of sibling 44 immature 9, 31 $,7 larvae, slides num- species rather than a multiplicity of distinct bered 1012-1026, UCNC, Davis; 1 immature

Figure 3. Rotylenchulus macrosomus n. sp. A—Immature female, anterior end; B—Larva, anterior end; C—Larva, tail; D—Male, anterior end; E—Immature female; F—Immature female, tail; G—Male, tail.

9,1 $, slide numbered R8, UCNC, Riverside; guished by its larger size of males (0.50-0.68 1 immature 9,2 $, slides numbered T-5S7p- mm vs. 0.40-0.49 for borealis) and immature T-588p, USDANC, Beltsville; 1 immature 9, females (0.52-0.64 mm vs. 0.37-0.46); also 1 5, slide numbered 40/6/1, NDRES, En- larger stylet (18—22 iJ. vs. 13—16 /j.) and longer gland; 1 immature 9,1 c , slides numbered hyaline portion of immature female tail (h = WT 1025-1026, PD, The Netherlands. 13-18 vs. 9-13 for borealis). TYPE HABITAT AND LOCALITY: Olive (Olea Additional specimens of R. macrosomus have eiiropaea) soil, Hulda, Israel. been identified from the following habitats DIAGNOSIS: R. macrosomus is elosely related and localities: peanut (Arachis hi/pogaea), to R. borealis from which it can be distin- Beit Dagan, Israel; bean (Phaseolus vulgaris)

Figure 4. Rotylenchulus clavicaudatus i\. sp. A—Male, anterior end; B—Immature female; C—Imma- ture female, anterior end; D—Male, tail; E—Immature female, tail. soil, Naven Yarr, Israel; banana soil, Karkur, region high, conoid, without any visible annu- Israel. lation. Metacorpus spheroid, valve 4 //. long. Rotylenchulus clavicaudatus n. sp. Esophageal glands overlapping intestine later- (Fig. 4, A-E) ally. Tail 43 n (35-50 /*) long, gradually tapering from the anus, terminus bluntly MEASUREMENTS (4 immature 5 ): L = 0.46-0.59 mm; a = 28-30; b = 4-5; b' = 2.9- rounded to hemispherical. 3.9; c = 12-16; c' = 3.8-5.3; h = 16-23; V MALE (Allotype): L = 0.54 mm; a — 29; = 57-59; stylet = 17-20 M; o = 75-85. b' = 4.0; c = 15; h = 18; T = 30; stylet = IMMATURE FEMALE (Holotype): L = 0.51 18 AS gubernaculum = 9 v; spicules — 20 M. mm; a = 30; b = 4; b' = 3.0; c = 12; c' = Similar to immature females. Esophagus re- 4.4; h = 23; V = 58; stylet = 19 M; o = 82. duced, metacorpus and valve indistinct. Cheilorhabdions thickened at both ends. Lip HOLOTYPE: Immature female, collected bv

J. Heyns, January, 1964, slide numbered 970, IMMATURE FEMALES: Following is an addi- UCNC, Davis. tion to the original descriptions. Esophageal ALLOTYPE: Male, same data as holotype, glands overlapping intestine laterally and slide numbered 970, UCNC, Davis. ventrally, usually more laterally. Each ovary PARATYPES: 3 immature females, same data with two flexures. Dorsal gland orifice more as the holotype, slide numbered 971, UCNC, than one stylet length posterior to stylet knob Davis. base. TYPE HABITAT AND LOCALITY: Associated LARVAE: Resembling immature females. with Strelitzia sp., virgin soil, on beach front Lip region conoid, finely annulated, not set at Port St. John's, Transkei, South Africa. off. Esophageal glands overlapping intestine DIAGNOSIS: R. clavicaudatus is closely re- laterally and ventrally. Tail bluntly rounded. lated to R. macrosomus from which it can be Specimens of R. borealis have been iden- distinguished by lower value of o; lack of tified from the following habitats and localities: annulation on lip region, more anterior position grass and weed soil, 2 miles below Venice, of the vulva and broader clavate shape of the France; grass and weed under Pine Grand tail. It can be distinguished from R. borealis Cornice above Monaco, France; grape vine by lack of annulation on lip region and lower soil (Vitis sp.) Castelnaudary, France; orchard value of o. nursery soil, Pieve del Cairo, Italy; vine soil, South Colombano, Italy; vine soil, Bresciano, Rotylenchidus borealis Loof and Italy. Oostenbriiik, 1962 (Fig. 11,A-C) Rotylenchulus leptus n. sp. MEASUREMENTS (9 immature 5 paratypes): (Fig. 5, A-G) L = 0.38-0.44 mm; a = 30-31; b = 3.5-4.2; MEASUREMENTS (23 immature 2 para- b' = 2.8-3.4; c = 11-13; c' = 3.7-4.2; h = types): L = 0.31-0.37 mm; a = 28-33; b = 8-13; V = 59-63; stylet = 14-16 M; o = 140. 3.4_4.0; b' = 2.6-3.3; c = 11-14; c' = 3.4- 5 $ paratypes: L = 0.43-0.49; a = 34-39; 4.5; h = 3-7; V = 58-64; stylet = 12-14 ^ b' = 3.6-4.2; c = 13-14; h = 9-15; T = 26- o = 140-160. 32; stylet = 12-14 «.; gubernaculum = 7-8 M; (10 larvae, paratypes): L = 0.30-0.36 spicules = 21-22 /x. mm; a = 24-26; b' = 3.5-3.9; c = 13-15; 8 immature 2 (Monaco, France): L = stylet = 12-14 /*. 0,39-0.44; a = 27-29; b = 3.7-4.2; b' = 2.3- IMMATURE FEMALE (Holotype): L = 0.36; 3.4; c = 13-15; c' =: 2.9-3.4; h = 9-13; V = a = 29; b = 3.4; b' = 2.8; c = 13; c' = 4.0; 60-64; stylet = 13-15 M; o = 147-185. h = 5; V = 62; stylet = 12 /x; o = 162. Body 8 $ (Monaco, France): L = 0.40-0.48 in open C shape, slender. Lip region high, mm; a = 29-37; b' = 3.5-4.9; c = 12-14; conoid, annules not visible. Stylet knobs small, h = 9-15; T = 28-41; gubernaculum = 7-8 sloping backwards. Excretory pore 71 M from AS spicules = 19-20 M; stylet = 12-14 /*. the anterior end (65-76 /x), below hemizonid. 6 larvae (Monaco, France): L = 0.39-0.43 Metacorpus oblong, esophageal glands over- mm; a = 25-29; b' = 3.5-4.1; c = 13-15; lapping the intestine laterally (some paratypes stylet = 12-13 /*. more ventrally). Tail 27 ^ (22—28 M) long, 2 immature 2 (South Colombano, Italy): slender, conoid, terminus rounded. L = 0.34-0.44 mm; a = 25-29; b = 3.7-4.2; MALE : Unknown. b' = 3.0-3.5; c = 13-14; c' = 3.0-3.5; h = LARVAE: Resembling female, tail more 13; V = 60-63; stylet = 14 M; o = 142-150. rounded than the immature female. 3 $ (South Colombano, Italy): L = 0.44- HOLOTYPE: Immature female, collected by 0.47 mm; a = 28-34; b' = 3.6-4.2; c = 14- G. Martin, 1957, slide numbered 972, UCNC, 16; h = 10-15; T = ?; stylet = 12-13 /x; Davis. gubernaculum = 8 M; spicules = 20—22 /x. PARATYPES: 30 immature 2, 15 larvae, 1 $ (Pieve del Cairo, Italy): L = 0.44 same data as holotype, distributed as follows: mm; a = 27; b' = 3.7; c = 13; h = 13; T = 26 immature 2, 15 larvae, slides numbered ?; stylet = 12 M; gubernaculum = 7 /x; spicules 973-981, 986-987, UCNC, Davis; 1 immature = 20 n. 2, slide numbered R6, UCNC, Riverside; 1 im-

Figure 5. Rotylenchulus leptus n. sp. A—Immature female, anterior end; B—Immature female; C— Larva, anterior end; D-F—Immature female, tail; G—Larva, tail. mature 5, slide numbered T-589p, USDANC, Rotylenchulus variabilis n. sp. Beltsville; 1 immature $, slide numbered 40/ (Figs. 6, A-H; 7, A-C) 4/1, NDRES, England; 1 immature 5, slide MEASUREMENTS (16 mature 9 paratypes): numbered WT 1022, PD, The Netherlands. L = 0.38-0.53 mm; width at the vulva = TYPE HABITAT AND LOCALITY: Soil around 0.07-0.12 mm; a = 4-6; V = 58-64; swollen bamboo (Bamboos vulgaris), unknown grass, portion of the body plus tail = 0.20-0.35 mm; and weed soil, Gwelo, Southern Rhodesia. stylet = ?; median esophageal bulb diameter ADDITIONAL COLLECTIONS: 20 immature ? = 17-22 M; eggs = 50-52 /* X 22 /* (within (Chipinga, Southern Rhodesia): L = 0.29- uterus). 0.33 mm; a = 27-31; b = 3.2-3.8; b' = 2.3- (22 immature ? paratypes): L = 0.30- 3.1; c = 11-13; c' = 3.6-4.7; h = 3-7; V = 0.37 mm; a = 22-26; b = 3.3-3.9; b' = 2.4- 57-63; stylet = 11-13 M; o = 123.* 3.1; c = 13-16; c' = 2.6-3.2; h = 3-6; V = DIAGNOSIS: R. leptus is most closely related 59-66; stylet = 13-15 M; o = 120-138. to R. parvus from which it can be distin- (21 $ paratypes): L = 0.34-0.41 mm; guished by its high conoid lip region, lack of a = 22-33; b' = 3.1-4.1; c = 14-20; h = 3-7; annulation in the lip region, longer and more T = 29-51; stylet = 10-12 M; gubernaculum slender tail, and larger hyaline portion of = 7-9 AI; spicules = 19-23 /*. immature female tail (h = 3-7 vs. less than 3 (2 larvae paratypes): L = 0.36-0.37 mm; for parvus). a = 22-23; b' = 3.6-3.7; c = 14-15; stylet = 12-13 M. Measurement of one specimen. IMMATURE FEMALE (Holotype): L = 0.36

Figure 6. Rotylenchulus variabilis n. sp. A—Immature female, anterior end; B—Immature female; C-D—Immature female, tail; E—Larva, anterior end; F—Larva, tail; G—Male, anterior end; H—Male, tail. mm; a = 26; b = 3.7; V = 2.8; c = 16; c' = terior to anus most specimens have rounded 2.6; h = 3; V = 63; stylet = 13 /x; o = 122. shape more strongly curved on ventral side Body in open C shape. Lip region hemi- setting off small projection (4—6 /j. long) more spherical with 5 annules (4-6). Cheilorhab- gross and cylindrical than in R. reniformis. dion thickened at both ends. Stylet knobs Metacorpus spheroid, valve large. Vulva with sloping backwards. Excretory pore 76 /* from prominent raised lips. the anterior end (72-82 M). Metacorpus ob- LARVAE: Resembling immature females. long, valve 4 n (4-6 M) long. Esophageal Lip region bluntly conoid. Tail more rounded glands overlapping intestine laterally. Tail than immature females. 21 n (20-24 n) long, terminus bluntly HOLOTYPE: Immature female, collected by rounded, annulations around the terminus G. Martin, March 1962, slide numbered 988, prominent. UCNC, Davis. MALE (Allotype): L = 0.37 mm; a = 28; ALLOTYPE: Male, same data as holotype, b' = 3.5; c = 17; h = 6; T = 44; guber- slide numbered 989, UCNC, Davis. naculum = 8 /u; spicules = 21 M. Stylet and PARATYPES: 17 mature 9, 33 immature 9, esophagus much reduced. Metacorpus and 31 $,2 larvae same data as holotype, dis- valve indistinct. Tail rounded. Stylet 10 M. tributed as follows: 17 mature 9, 26 imma- MATURE FEMALE: Body strongly curved, ture 9, 26 $, 2 larvae, slides numbered 990- tail often crosses the neck. Body shape pos- 999, 1007-1009, UCNC, Davis; 3 immature terior to vulva intermediate between R. reni- 9,1 $, slide numbered R7, UCNC, Riverside; formis and R. pawns, narrows to anus. Pos- 2 immature 9,25, slides numbered T-590p-

Figure 7. Rotylenchulus variabilis n. sp. A—Mature female; B-C—Mature female, tail.

T-591p, USDANC, Beltsville; 1 immature 9, 1 been identified from the following habitats and $, slides numbered 40/5/1-40/5/2, NDRES, localities: bean (Phaseohis vulgaris) soil, corn England; 1 immature 9 , 1 <5 , slides numbered soil, Machakos district, Kenya; bean soil, sweet WT 1023-1024, PD, The Netherlands. potato soil, Kiambu Hill, Kenya; corn soil, TYPE HABITAT AND LOCALITY: Soil around banana hill, Kenya; cowpea soil (Vigna Rumex sp., Inyanga Orchard area, Southern sinensis), Ondo Province, Nigeria; spear grass Rhodesia. (Imperata cylindrical soil, oil palm (Elaeif; DIAGNOSIS: R. variabilis is most closely guineensis) soil, Oyo Province, Nigeria; soil related to R. parvus and distinguished by the around the roots of unknown vegetables, Niger distinct annulation of the lip region, longer Province, Nigeria. hyaline tail portion (h = 3-6 vs. less than 3) R. variabilis shows variation in certain char- and longer valve in metacorpus of immature acters to a remarkable degree within the same females, and by large mature female with population. Most of the immature females terminal projection. It can be distinguished of paratypes and other collections have a long from R. leptiis by the presence of annulation esophagus measuring up to 80 /x from the in lip region. valve to the end of the esophageal glands. In Additional specimens of R. variabilis have some specimens this distance may be as low

Figure 8. Rotylenchulus parvus. A—Male, anterior end; B—Larva, anterior end; C—Immature female, anterior end; D—Male, tail; E—Larva, tail; F—Immature female; G—Mature female; H-I—Immature female, tail; J—Mature female, tail. as 56 M. In most of the immature females the ture females collected from sugarcane soil in annulation is pronounced around the tail termi- Mauritius. Goodey (1963) judged it to be a nus but there are some individuals in which synonym of R. reniformis. Type specimens the tail terminus is smooth. Some male speci- sent by Williams were studied together with mens from corn soil, Kiambu Hill, Kenya a population originally from soil of a cotton have a very pointed tail. field in Imperial Valley, California. That field population was maintained in the greenhouse Rotylenchulus parvus (Williams, 1960) at Davis and used for life history and host Sher, 1961 range studies. Large numbers of specimens Helicotylenchus parvus Williams, 1960 were available from roots of barley in those (rig. s, A-j) greenhouse cultures. No males were seen in Williams described this species from imma- these Davis cultures. The 3 males described

Copyright © 2011, The Helminthological Society of Washington OF WASHINGTON, VOLUME 35, NUMBER 2, JULY 1968 185 below were provided by Dr. D. E. Konicek anus 10—16 At. Vulva with prominent raised who collected 2 from the field population and lips. Vagina funnel-shaped extending half one from his greenhouse culture. It was found across the body. Tail without spikelike process both immature and mature females have mor- or projection. phological characters distinctly different from MALES: Lip region high, hemispherical. R. reniformis which made possible a diagnosis Cephalic sclerotization weak. Prorhabdion not clearly indicating its status as a separate pronounced, difficult to see. Esophagus re- species. duced. Lumen of the esophagus, metacorpus MEASUREMENTS (10 immature 2 paratypes): and valve difficult to resolve. Tail rounded. L = 0.23-0.27 mm; a = 18-25; b = 3.5-3.8; LARVAE: Resembling immature females. b' = 2.4-3.1; c = 16-19; c' = 2.1-2.7; h = Tail bluntly rounded, without projections. less than 3; V = 61-65; stylet = 12-13 M; Specimens identified as R. parvus have been o = ?. examined from the following habitats and lo- 40 immature 5 (Imperial Valley, California): calities: papaya soil, Nairobi, Kenya; corn soil, L = 0.25-0.34 mm; a = 20-26; b = 3.1-3.7; Southern Rhodesia; fallow soil, Penhalonga, b' = 2.1-3.0; c = 16-20; c' = 2.0-2.7; h = Southern Rhodesia; munga (Pennisetum ty- less than 3; V = 60-66; stylet = 12-14 /*; phoides) soil, Salisbury, Southern Rhodesia; o = 90-107. sunhemp (Crotalaria juncea) soil, Southern 15 mature 2 (Greenhouse culture, Davis; Rhodesia; Turkish tobacco soil, Natoba farm, originally from Imperial Valley, California): Pimba, Northern Rhodesia. L = 0.25-0.36 mm; width at vulva = 0.04- The original description of R. parvus shows 0.08 mm; a = 4-7; V = 61-66; swollen por- the esophageal gland as overlapping the intes- tion of the body plus tail = 0.15-0.25 mm; tine more dorsally and the ovaries as out- stylet = 12-15 M; median esophageal bulb stretched. The paratype specimens show that diameter = 12-15 /*; eggs = 56-69 /* X 30- the esophageal glands overlap the intestine 38 M. ventrally and laterally and the ovaries typical 2 $ (Imperial Valley, California): L = with two flexures. The stylet could be seen 0.38-0.46 mm; a = 28-32; b' = 3.6-4.0; c = only in one specimen of three males examined. 18-23; h = less than 3; T = 32-34; stylet = 10 M; gubernaculum = 5 M; spicules = 16 A*. Rotylenchiihis anamictus n. sp. 1 $ (Greenhouse culture, Sacramento; (Fig. 9, A-H) originally from Imperial Valley, California): MEASUREMENTS (31 immature 5 paratypes): L = 0.39 mm; a = 29; b' = 4.0; c = 14; h = 3; L = 0.27-0.33 mm; a = 22-26; b = 3.0-3.9; T = 41; stylet = ?; gubernaculum = 7 M; spic- b' = 2.1-3.0; c = 13-17; c' = 2.3-2.9; h = ules = 16 /j.. 4-7; V = 67-72; stylet = 12-14 M; o = 107- 10 larvae (Greenhouse culture, Davis; orig- 141. inally from Imperial Valley, California): L = (12 $ paratypes): L = 0.33-0.40 mm; a = 0.25-0.34 mm; a = 21-25; b' = 2,5-3.5; c = 26-30; b' = 3.3-4.3; c = 14-17; h = 3-6; 14-18; stylet = 12-13 /*. stylet = 10—12 n; gubernaculum = 6-7 M; IMMATURE FEMALES: Body in an open C to spicules = 18-20 /*. loose spiral. Lip region low rounded, finely (5 larvae paratypes): L = 0.31-0.36 mm; annulated; stylet knobs sloping backwards. a = 23-26; b' = 3.1-3.5; c = 13-15; stylet = Excretory pore 60—75 A* from anterior end. 12 M. Metacorpus spheroid, valve length 3 M. Esoph- IMMATURE FEMALE (Holotype): L = 0.30; ageal glands overlapping intestine laterally and a = 23; b = 3.4; b' = 2.6; c = 15; c' = 3.0; ventrally, frequently laterally. Tail short, 13— h = 4; V = 70; stylet = 13 AS o = 140. Body 18 M; terminus in form of a short ventral in open C shape. Lip region low, rounded, projection. with 3 annules (3—4). Stylet knobs rounded, MATURE FEMALES: Posterior part of the sloping backwards. Excretory pore posterior body often crosses the neck. Body contour to hemizonid, 69 /* from the anterior end (69- beyond the vulva tapers abruptly, postanal 83 A1). Metacorpus spheroid, valve length 3 M part slender, not hemispherical, similar to im- (3—4 M). Esophageal glands overlapping the mature females but wider. Body width at the intestine laterally and ventrally, slightly more

Figure 9. Rotylenchulus anamictus n. sp. A—Immature female, anterior end; B—Male, anterior end; C—Larva, anterior end; D-E—Immature female, tail; F—Male, tail; G—Larva, tail; H—Immature female.

ventrally. Tail conoid, 21 M (18-23 M) long, esophagus not distinct. Tail more rounded terminus rounded. than immature females. MALE (Allotype): L = 0.34 mm; a = 26; HOLOTYPE: Immature female, collected by b' = 3.4; c = 17; h - 3; T = 29; gubernac- F. Lamberti, July 1964, slide numbered 944, ulum = 7 ft; spicules = 19 M. Lip region hemi- UCNC, Davis. spherical, annulated. Cephalic sclerotization ALLOTYPE: Male, same data as the holo- weaker than immature females. Stylet reduced, type, slide numbered 945, UCNC, Davis. 10 M. Excretory pore 74 M from anterior end PARATYPES: 38 immature 2, 12 6 , 10 lar- (69—75 ,"). Lumen of esophagus, metacorpus vae, same data as holotype, distributed as and valve difficult to resolve. Tail conoid, follows: 30 immature 2,8 6 , 10 larvae, slides terminus rounded. numbered 946-962, UCNC, Davis; 3 immature LARVAE: Lip region conoid. Lumen of 2, 1 $, slides numbered R4-R5, UCNC,

Figure 10. Rotylenchuhis macrodoratus n. sp. A—Immature female, anterior end; B—Male, anterior end; C—Male, tail; D—Larva, anterior end; E-G—Immature female, tail; H—Immature female; I—Larva, tail.

Riverside; 3 immature ?, 1 $, slide numbered portion of immature female tail (h = 4—7 vs. T-592p, USDANC, Beltsville; 1 immature 9, 1 less than 3), different shape of the immature $, slides numbered 40/3/1-40/3/2, NDRES, female tail, conoid shape of larval head and England; 1 immature ?, 1 $, slides numbered usually longer tail of immature female. It can WT 1020-1021, PD, The Netherlands. be distinguished from R. reniformis by the TYPE HABITAT AND LOCALITY: Soil around short body length, smaller stylet, and low Acacia sp. about 2 miles south of Merca, rounded lip region. Somaliland. DIAGNOSIS: R. anamictus is most closely re- Rotylenchuhis macrodoratus n. sp. lated to R. parvus and can be distinguished on (Figs. 10, A-I; 11, D-F) the more posterior position of the vulva (V = MEASUREMENTS (25 immature 5 paratypes): 67-72 vs. 60-66 for parvus), larger hyaline L = 0.40-0.49 mm; a = 22-28; b = 3.2-4.7;

Figure 11. Rotylenchulus borealis. A—Larva, anterior end; B—Larva, tail; C—Male, tail. Rotylench- ulus macrodoratiis n. sp. D-E—Mature female, tail; F—Mature female. b' = 2.8-3.6; c = 16-24; c' = 1.8-2.4; h = a = 21-28; b' = 3.0-4.7; c = 15-20; stylet = 6-12; V = 64-72; stylet = 22-26 ^, o = 56- 20-22 M. 82. IMMATURE FEMALE (Holotype): L = 0.46 (12 $ paratypes): L = 0.45-0.53 mm; a = mm; a = 28; b = 4.2; b' = 3.3; c = 20; c' = 27-32; b' = 3.8-4.6; c = 21-28; h = 6-10; 2.4; h = 6; V = 71; stylet = 25 M; o = 64. stylet = 17-20 /JL; gubernaculum = 7 M; spic- Lip region conoid with fine annulation. Sec- ules = 20 iJ.. ond and third incisures in lateral field lighter (12 mature 9 paratypes): L = 0.41-0.53 than the 2 outer lines, gradually disappearing mm; width at vulva = 0.06-0.19 mm; a = 4-6; near level of flexure of posterior ovary. Ce- V = 64-71; length of the swollen portion plus phalic sclerotization heavy, arches downwards tail = 0.17—0.34 mm; stylet = ?; median esoph- strongly. Spear knobs large, anchor-shaped. ageal bulb diameter — 13—16 n. Excretory pore 100 M (91-101 M) from anterior (10 larvae paratypes): L — 0.40—0.50 mm; end. Metacorpus oblong, valve 3 M (3-5 /u)

Copyright © 2011, The Helminthological Society of Washington OF WASHINGTON, VOLUME 35, NUMBER 2, JULY 1968 189 long. Esophageal glands overlapping intestine in a latero-ventral position. However, these ventrally. Tail 2.1. ^ (20-23 M) long, terminus measurements could be taken accurately by bluntly rounded. mounting the tail end in proper position in MALE (Allotypc): L = 0.46 mm; a = 29; glycerin jelly. b' = 3.9; c = 23; h = 6; T = 42; stylet = 18 /*; Mature females of R. macrodoratiifi also gubernaculum = ?; spicules = ?. Lip region have been identified from the roots of Laurus high, hemispherical. Stylet knobs and esoph- nobilis, Bari, Italy. Additional specimens of agus much reduced, metacorpus and valve in- immature females, males and larvae, of R. distinct. First lateral line from the ventral side macrodoratiis have been identified from the merging with edge of caudal alae. following habitats and localities: grape (Vitis MATURE FEMALES: Body ventrally curved, sp.) soil, Lazzaretto, Italy; almond (Primus tail sometimes crossing the neck. Slope of the amygdalus) soil, Torre Tresca, Italy. body contour anterior and posterior to vulva abrupt. Body contour behind vulva tapers to Key to the Species of Rotylenchulus* a degree intermediate between R. reniformis 1. Stylet knobs anchor-shaped, stylet 22 t*. and R. parviis. Metacorpus spheroid. Vulva or more macrodoratiifi n. sp. with raised, pronounced lips. Tail terminus Stylet knobs spheroid, stylet usually less with spikelike process. than 22 /* 2 LARVAE: Resembling immature females. 2. h = 14 or more; 15 or more in male, L Tail more rounded than immature females. of male 0.5 mm or more 3 HOLOTYPE: Immature female, collected by h — 13 or less; 15 or less in male; L of A. Ciccarone, March 1962, slide numbered male when present 0.49 mm or less „ 4 929, UCNC, Davis. 3. o = < 100 clauicaiidatus n. sp. ALLOTYPE: Male, same data as holotype, o = > 100 macrosomus n. sp. slide numbered 930; UCNC, Davis. 4. V = 66 or less 5 PARATYPES: 15 mature 2, 85 immature 2, V = 67 or more 8 24 $, 28 larvae, same data as holotype, dis- 5. h = > 8; > 9 in male borealis tributed as follows: 15 mature 2, 70 imma- h = < 7; < 7 in males when present ____ 6 ture 2, 19 $, 27 larvae, slides numbered 931- 6. c' = 3.3 or more, lip region conoid 943, 963-969, 982-985, 1027-1032, UCNC, without visible annulation, males un- Davis; 7 immature 2,15,1 larva, slide num- known leptus n. sp. bered R9, UCNC, Riverside; 4 immature 2, c' = 3.2 or less, lip region hemispher- 1 $, slide numbered T-593p, USDANC, Belts- ical or rounded, annulations present, ville; 3 immature 2,2 $, slide numbered males common or rare 7 40/7/1, NDRES, England; 1 immature 2, 7. h = < 3; lip annulation very fine, not 1 $, slides numbered WT 1027-1028, PD, distinct, swollen female tail without The Netherlands. spikelike process parvus TYPE HABITAT AND LOCALITY: Soil around h = > 3; lip annulations distinct; swol- grape (Vttis sp.), Bari, Italy. len female tail with spikelike proc- DIAGNOSIS: R. macrodoratiis is distinguished ess variabilis n. sp. from all other species of Rotylenchulus by its 8. Lip region high; stylet 15 /* or more; robust stylet and knobs with forward directed L = 0.34 mm or more; o — < 106 —. processes, acute conoid lip region, and heavy reniformis sclerotization of the cephalic framework; Lip region low; stylet 14 /*. or less; L = shorter tail which is sometimes less than twice 0.33 mm or less; o usually > 110 the anal body diameter. anamictus n. sp. Esophageal glands in some paratype specimens of immature females are in the form of Discussion hemispherical lobes and overlap intestine lat- Husain and Khan (1967) indicated a close erally. It was not possible to take measure- relationship between Rotylenchulus and the ments of spicules and gubernaculum in toto Hoplolaiminae when they proposed the sub- mounts because all the male tails of the para- * Characters mentioned hereafter refer to immature fe- types as well as from other collections were males unless and otherwise mentioned.

Figure 12. Rotylenchulus spp. Mature females. A-D—R. parvus; E-F—jR. variabilis n. sp.; G-H—R. macrodoratus n. sp.; I-J—R- reniformis. family Rotylenchulinae within the family Hop- tail." Rotylenchulus females all have double lolaimidae. The present study supports this flexures in the ovaries. In most species the conclusion of relationship principally by the female tail is more than twice the anal body immature females of R. macrodoratus which diameter, even in R. macrodoratus where the bear many characters similar to species of female tail is short, many individuals have a Helicotylenchus. tail more than twice the anal body diameter. Some of the characters of Hoplolaiminae as The males of Rotylenchulus have adanal emended by Sher (1961) are: "ovaries out- caudal alae. Considering these differences it stretched, amphidelphic, and didelphic, female is judged most appropriate to place Rotylen- tail short, not more than twice the anal body chulus in Rotylenchulinae. diameter . . . caudal alae enveloping male The characters of immature females which

Copyright © 2011, The Helminthological Society of Washington OF WASHINGTON, VOLUME 35, NUMBER 2, JULY 1968 191 are considered useful in separation of the other groups. The lip region of R. borealis is species are: shape of the lip region, annula- similar to that of reniformis and the lower tion in lip region, stylet length, knob shape, limit of h comes within the range of reniformis. position of the dorsal gland orifice, position of Gollections of mature females are needed for the vulva and the length of the hyaline portion all species for comparative studies with imma- of the tail. Although males are known in all ture females, larvae, and males to gain a better species except R. leptus, the male morphology understanding of the relationship of the does not usually provide useful diagnostic species. characters. Generally the same is true with Acknowledgments the larvae. The authors are indebted to Dr. M. W. Allen Mature females are known in only five of for critically reviewing the manuscript and his the nine species of Rotylenchulus. This stage advice and counsel during the study. has considerable importance in distinguishing The authors also acknowledge the help of species but will be limited in application until Dr. D. E. Konicek, State Department of Agri- mature female specimens are available for all culture, Sacramento, for the loan of specimens species. In the meantime, the relationship be- and to other authors who made available some tween species perhaps is best judged on the of their type speciemns and other collections basis of the lip region in combination with tail of Rotylenchulus. characters of immature females. Such an examination reveals several groupings: Group Summary I—species with lip region low, rounded to The genus Rotylenchulus Linford and Oli- hemispherical, annulated; h less than 8—rep- veira, 1940 is reviewed and the generic diag- resented by parvus, variabilis, and anamictus; nosis is emended. The morphology of Roty- Group II—species with lip region high and lenchulus and key to the species is given. The conoid, without visible annulation; h less than nominal species of Rotijlenchulus now total 8—represented by leptus; Group III—species nine, six of which are described here as new. with lip region high, conoid, rounded with Rotylenchulus nicotiana (Yokoo and Tanaka, annules, h less than 13—represented by reni- 1954) Baker, 1962 and Rotylenchulus stakmani formis; Group IV—species with lip region Husain and Khan, 1965 are synonymized with high, conoid, with or without annulation; h is R. reniformis. Description of neotype and re- 8—24—represented by borealis, macrosomus, descriptions of various stages are made for the and clavicaudatus; Group V—species with lip genotype, R. reniformis. Males and mature region acutely conoid, with annules; h less females of R. parvus and larva of R. borealis than 13—represented by macrodoratus. are described for the first time. Immature females of macrodoratus show the least relationship with any other group. Lip Literature Cited region, stylet size and knob shape of this spe- Allen, M. W., and S. A. Sher. 1967. Taxo- cies show complete departure from all the nomic problems concerning the phytoparasitic nematodes. A. Rev. Phytopath., 5: 247-264. species of this genus. The closer relationships Baker, A. D. 1962. Check lists of the nematode of parvus-group and reniformis is exhibited by superfamilies Dorylaimoidea, Rhabditoidea, anamictus which shares characters of both Tylenchoidea and Aphelenchoidea. E. J. Brill, parvus and reniformis. Its total size, stylet Leiden, Holland. 261 pp. length, lip region are similar to those of parvus Carvalho, J. C. 1957. Rotylenchus elisensis, but the vulva position, tail and frequency of nova especia associado com raizes de soja. occurrences of males bring this species closer Revista Inst. Adolfo Lutz 17: 43-46. to reniformis. R. leptus stands in between . 1959. Helicotylenchus elisensis n. comb. Group I and Group III and comes closer to the (Nematoda: Tylenchidae). Arg. Inst. Biol., parvus-group. Whereas in gross morphology Sao Paulo, 26: 45-48. Chitwood, B. G., and M. B. Chitwood. 1950. this species is similar to the parvus-group its An introduction to nematology. Section I, conoid lip region, lack of annulation in lip Anatomy. (Revised). Monumental Printing region, slender and long tail do not certainly fit Co., Baltimore, Md. 213 pp. this species in Group I. The borealis-group is Das, V. M. 1960. Studies on the nematode more closely related to Group III than any parasites of plants in Hyderabad (Andhra

Pradesh, India). Zeitschr. fur Parasitenkunde koo and Tanaka, 1954) n. comb, in Japan I9(b): 553-605. (Tylenchida: Hoplolaimidae). Jap. J. Appl. Goodey, T. 1963. Soil and freshwater nema- Ent. Zool. 11: 169-176. todes. 2nd Edition by J. B. Goodey, Me- Peacock, F. C. 1956. The reniform nematode thuen & Co., London. 544 pp. in the Gold Coast. Nematologica 1(4): 307- Hopper, B. E., and E. J. Cairns. 1959. Taxo- 310. nomic keys to plant, soil and aquatic nema- Perry, V. G., H. M. Darling, and G. Thome. todes. Ala. Polytech. Inst., Auburn, Ala. 1959. Anatomy, taxonomy and control of (mimeo). 176 pp. certain spiral nematodes attacking blue grass Husain, S. I., and A. M. Khan. 1965. Rottj- in Wisconsin. Bull. Wise. Agr. Exp. Stn. No. lenchulus stakmani n. sp., with a key to the 207: 1-24. species of the genus (Nematoda: Tylenchida). Sher, S. A. 1961. Revision of the Hoplolaiminae Proc. Helm. Soc. Wash. 32(1): 21-23. (Nematoda) I. Classification of nominal gen- , and . 1967. A new subfamily, a era and nominal species. Nematologica 6(2): new subgenus and eight new species of nema- 155-169. todes from India belonging to superfamily . 1963. Revision of the Hoplolaiminae Tylenchoidea. Proc. Helm. Soc. Wash. 34: (Nematoda) II. Hoplolaimus Daday, 1905 and 175-186. Aorolaimus n. gen. Nematologica 9(2): 267- Linford, M. B., and J. M. Oliveira. 1940. 295. Rotylenchulus reniformis, nov. gen., n. sp., a 1966. Revision of the Hoplolaiminae nematode parasite of roots. Proc. Helm. Soc. (Nematoda) VI. Helicotylenchus Steiner, Wash. 7(1): 35-42. 1945. Nematologica 12(1): 1-56. , and F. Yap. 1940. Some host plants of Skarbilovich, T. S. 1960. Sveklowinaya nema- reniform nematode in Hawaii. Proc. Helm. toda i mery borjby s nei. Trudy Vses. Inst. Soc. Wash. 7(1): 42-44. Gel'mint. 8: 5-214. Loof, P. A. A., and M. Oostenbrink. 1962. Swarup, G., R. P. Nath, and C. L. Sethi. 1967. Rotylenchulus borealis n. sp. with a key to The plant parasitic nematode genus Rotylen- the species of Rotylenchulus. Nematologica chulus in India. Indian Phytopath. 20(2): 7(1): 83-90. 118-123. Lordello, L, G. E., and R. Cesnik. 1958. Thorne, G. 1949. On the classification of the Alguns nematodeos do tomateiro. Revista Bra- Tylenchida, new order (Nematoda: Phas- sil. Biol. 18(2): 159-165. mida). Proc. Helm. Soc. Wash. 16(2): 37- Martin, G. C. 1955. Plant and soil nematodes 73. of the Federation of Rhodesia and Nyasaland. . 1961. Principles of Nematology. Mc- Rhodesia Agr. J. 52(4): 346-361. Graw-Hill Book Co., New York. 553 pp. Nakasono, K. 1966. Role of males in repro- Williams, J. R. 1960. Studies on the nematode duction of the reniform nematodes, Rotijlen- soil fauna of sugarcane fields in Mauritius. chulus spp. (Tylenchida: Hoplolaimidae). Tylenchoidea (Partim). Mauritias Sugar Ind. Appl. Ent. Zool. Tokyo, 1: 203-205. Res. Inst. 4: 1-30. , and M. Ichinohe. 1967. On the reni- Yokoo, T., and I. Tanaka. 1954. Bull Kago- form nematode Rotylenchulus nicotiana (Yo- shima Tob. Exp. Stn. No. 9: 59-63.

Isoglaridacris hexacotyle comb. n. (Cestoidea: Caryophyllidea) from Catostomid Fishes in Southwestern North America JOHN S. MACKIEWICZ State University of New York at Albany, Albany, New York

Glaridacris hexacotyle (Linton, 1897) is Ulmer, 1967, holotype, USNM Helm. Coll. known only from the superficial description No. 60302; and I. bulbocimis Mackiewicz, of Linton (1897) and the more detailed ac- 1965, topotypes, 136 whole mounts of count of Hunter (1930); both descriptions gravid and immature worms from the author's were based on the same eight specimens from collection. "Catostomus sp." from the Gila and Salt Rivers in Arizona. This paper presents additional Isoglaridacris hexacotyle comb. n. descriptive data and proposes nomenclatorial (Figs. 1-8) changes based upon a restudy of the original SYNONYMY: Monobothrium hexacotyle Lin- material and of new collections from the Salt ton, 1897; Caryophijllaeus hexacotyle (Linton, River. 1897) Woodland, 1923; Glaridacris hexacotyle (Linton, 1897) Hunter, 1927. Materials and Methods HOSTS AND DISTRIBUTION: Catostomus sp., A total of 160 cestodes from 15 of 16 sonora Gila and Salt River, Arizona (Linton, 1897); suckers, Catostomus insignis Baird and Girard, White River, 8 km W. Meeker, Rio Blanco and 17 from 7 of 29 gila suckers, Catostomus Comity, Colorado (collected by G. Schmidt). (Pantosteus) clarki Baird and Girard, was col- Catostomus insignis Baird and Girard, Gila lected in December 1964, at Coons Bluff on River, Tonto National Forest, Mariocopa the Salt River in Tonto National Forest near County, Arizona. Catostomus (Pantosteus) Tempe, Arizona (Mariocopa County). The clarki Baird and Girard, same as C. insignis sonora suckers ranged in total length from as well as Birch Creek, a tributary of the 225-494 mm (mean: 359 mm) whereas the Virgin River, Zion National Park, Washington gila suckers ranged from 213-375 mm (mean: County, Utah. 260 mm). Living cestodes were fixed in HABITAT: Small intestine, weakly attached steaming AFA or 5 per cent formalin and or free. stained with Semichon's carmine. TYPE: Lectotype USNM Helm. Coll. No. Only 108 cestodes (82 gravid) from the 4793, paralectotypes USNM Helm. Coll. No. sonora sucker and 16 (nine gravid) from the 4793. Supplementary material includes USNM gila sucker were stained and mounted; two Helm. Coll. No. 70927, 3 whole mounts from gravid individuals from the sonora sucker C. clarki, USNM Helm. Coll. No. 70928, 10 were sectioned. Measurements were made on whole mounts from C. insignis, and British a selected sample of 30 gravid worms from Museum (Natural History) Helm. Coll. No. 12 sonora suckers and five from three gila 1967.12.29. 1-3, 3 whole mounts from C. suckers. Host nomenclature is according to insignis, the recent treatment of Smith (1966). MORPHOLOGY: There appears to be some Comparative material included: Glaridacris confusion regarding the number of gonopores hexacotyle (Linton, 1897), USNM Helm. Coll. found in this species. According to Hunter No. 49727 (label: USNM Helm. Coll. 49727 (1930: 66), "The cirrus sac opens flush with from 4793) consisting of a vial with three the ventral surface of the parasite, and the posterior regions, two scolexes, and darkly female system empties in a similar fashion just stained fragments of the testicular region; 19 behind that of the male." In his specific of Hunter's original slides of this species (his diagnosis, however, he seems to imply that numbers: 584.1 a-g, .2 to .4 a-b, .5 to .7 a-cl, there is one gonopore since the terms "common and .8 to .9); Isoglaridacris folius Fredrickson genital atrium" are used. Linton (1897), on and Ulmer, 1967, paratype, USNM Helm. the other hand, mentioned a genital aperature Coll. No. 60301; I. longus Fredrickson and and clearly pictured one in his figure 3, plate

8 Figures 1-8. Isoglaridacris hexacotyle. 1. Gravid individual. 2. Scolex. 3. Scolex variations. 4. Varia- tions in ovarian shape. 5. Posterior region. 6. Midsagital section through the gonopore. 7. Operculate egg with ovum and vitelline cells. 8. Immature individuals.

XXVIII. That the cirrus joins the uterovaginal incidence have been presented by Chandra canal that in turn opens on the surface as a (1966) and Amin (1968). single gonopore is borne out by a reexamma- HOST-PARASITE RELATIONSHIPS: There were tion of the fragments in Linton's vial, of striking differences in the incidence and worm Hunter's slides, particularly numbers 584.7c burden between sonora and gila suckers col- and 584.9 and examination of 108 cestodes in lected from the same pool. While a mean of the present study. Sections on Hunter's slide, 10.7 (3-20) cestodes occurred in 15 of 16 that includes the one used for his figure 41, sonora suckers only 2.4 (1-6) were found in have the cirrus partially everted through a 7 of 29 gila suckers. Using the chi square single gonopore. Since Glaridacris Cooper, contingency test, the probability is less than 1920, has two gonopores, it is necessary to 0.005 per cent that the two hosts are infected make the nomenclatorial changes indicated to the same degree. Such a significant dif- above. ference may be associated with the different My other data are in general agreement with feeding habits of the hosts as reflected in the Hunter's (1930) detailed description but in- structure of their mouths. For example, the dicate that there is a greater variation in the sonora sucker has thick, fleshy papillose lips species. Mean measurements (in mm, ranges that aid in sucking material from mud, sand, in parentheses) of a selected sample of 30 or the surface of rocks. The gila sucker, on gravid worms from sonora suckers are: length the other hand, has the same soft, papillose 10.4 (7-15.5), width at gonopore 0.74 (0.5- lips but with a tough cartilaginous inner edge 1.1), and number of testes 162 (132-205). that aids in scraping or tearing material from Measurements of five specimens from gila the surface of rocks. Sonora suckers are, there- suckers are: length 10.1 (8.7-12), width at fore, more apt to ingest the probably inter- gonopore 0.54 (0.5-0.6), and number of testes mediate host of I. hexacotyle, tubificid worms, 174 (143-223). Immature worms resembled which normally live in mud. adults and lacked a cercomer (Fig. 8). The In August 1964, an average of three (1—12) characteristic hexagonal scolex, from which the cestodes were found in four of seven gila species derives its name, is found only on suckers, ranging in size from 8.5-18 cm, from contracted specimens that have the apex of Birch Creek in Zion National Park, Utah. Fish the scolex thrust forward. Normally the scolex were observed scraping and tearing algae and is much like that of Glaridacris catostomi other growths from the surface of large rocks. Cooper, 1920 (Fig. 2); some of the variations SYSTEMATICS: This species is placed in the are shown in figure 3. In all cases (91) the genus Isoglaridacris Mackiewicz, 1965, primar- vitellaria extended more anteriorly than the ily on the basis of its having a single gonopore; testes (Fig. 1) and posteriorly to the cirrus; Glaridacris has two gonopores and, therefore, postovarian vitellaria were always present. cannot receive this species. All of the other Two vitelline ducts connect the postovarian characters, such as vitellaria in lateral rows, vitellaria with the vitelline reservoir; Hunter cuneiform scolex, and type of ovary, are clearly (1930) reported only one. those of Isoglaridacris. There are three other Two types of ovarian variation were ob- species of Isoglaridacris: I. biilbocirrus Mack- served: unequal arm lengths (Figs. 1, 4) and iewicz, 1965, from C. commersoni (Lacepede) fusion of the posterior arms to form an in- (Mackiewicz, 1965), and /. longus Fredrick- verted A-shaped ovary. This latter variation son and Ulmer, 1967, from Moxostoma macro- was found in 8 of the 82 gravid worms from lepidotum (LeSueur) and I. folius Fredrickson only the sonora sucker. and Ulmer, 1967, from M. erythnirum (Raf.) Ten ova dissected from the distal part of (Fredrickson and Ulmer, 1967). the uterus and measured in water were 35 According to Linton (1897: 426) the type (32-40) /x long and 29 (28-30) M wide. The is USMM No. 4793 with the label "From shell is rough, being covered with what ap- sucker (Catostomus sp.) inhabiting the Gila pears to be excess shell-material globules; and Salt River, Arizona; E. Palmer." This there is a small operculum and there are three number, since recatalogued as No. 49727, to five vitelline cells per egg (Fig. 7). consists of a vial with only fragments of Additional data on variation growth and worms; no specimen has been designated as

Copyright © 2011, The Helminthological Society of Washington 196 PROCEEDINGS OF THE HELMINTHOLOGICAL SOCIETY the type. The only other specimens from the nearctic and palearctic Canjophyllaeiis to each original collection are the 19 slides (all sec- other and to Glaridacris. tions) found in Hunter's caryophyllaeid collection. Since there is no type specimen, I Acknowledgments propose to designate slides 584.7a-d of the Thanks are extended to Mr. Richard Koehn, Hunter Collection (now USNM Coll. No. graduate student at Arizona State University, 4793) as lectotype; they consist of serial for his generous help in collecting hosts from saggital sections of a whole worm. All of the the Salt River; to the National Park Service other slides of this species from the same for permission to examine fish from Zion collection are designated as paralectotypes National Park; to Dr. G. W. Hunter III for and are now catalogued as USNM Helm. Coll. loan of the Glaridacris part of his collection; No. 4793. and to Dr. G. Smith for his opinion regarding Discussion the identity of the Birch Creek fish. The Arizona collections were supported by This nomenclatorial change reduces the Grant No. 20-147-A from ASRC of the State number of nearctic Glaridacris species to four; University of New York. Kennedy (1965) has recently considered the palearctic species, G. brachums (Mrazek, Literature Cited 1908) and G. limnodrili Yamaguti, 1934, Amin, O. M. 1968. Helminthic fauna of suck- as synonyms of Archigctes Leuckart, 1878. ers (Catostomidae) of the Gila River system, The nearctic species fall into two dissimilar Arizona. Ph.D. Thesis. Arizona State Univ. groups: one represented by G. catostomi, 202 pp. (No. 68-1644) Univ. Microfilms. Ann which is long and filiform, has a cuneiform Arbor, Mich. (unpublished). scolex, annularly arranged vitellaria and widely Chandra, Shantha. 1966. Morphological varia- separated gonospores; and the other by G. tion of Glaridacris hexacotyle (Linton, 1897) laruei (Lament, 1921), G. oligorchis Haderlie, (Cestoidea: Caryophyllaeidae). M.S. Thesis. 1953 and G. confusa Hunter, 1927, which are State Univ. N. Y. at Albany. 58 pp. (unpub- relatively short forms having a terminal disc lished ). on the scolex, vitellaria in two lateral rows and Fredrickson, L. H., and M. J. Ulmer. 1967. gonopores close together. Caryophyllaeid cestodes from two species of Redhorse (Moxostoma). Proc. Iowa Acad. Such a grouping strongly suggests that the Sci. 72: 444-461. genus Glaridacris should be revised to include Hunter, G. W. III. 1930. Studies on the Caryo- forms more similar to each other. Indeed, the phyllaeidae of North America. III. Biol. type of the genus, G. catostomi, more closely Monog. 11: 1-186. resembles Caryophyllaeits terebrans (Linton, Kennedy, C. R. 1965. The life-history of Archi- 1893) than it does any other species of Glarid- getes limnodrili (Yamaguti) (Cestoda: Caryo- acris. Not only do both species have a similar phyllaeidae) and its development in the general morphology but study of living and invertebrate host. Parasitology 55: 427-437. preserved C. terebrans from Catostomus ardens Linton, E. 1897. Notes on cestode parasites of Jordan and Gilbert in Wyoming clearly con- fishes. Proc. U. S. Nat. Mus. XX: 423-456, firms the presence of loculi on a weakly 8 pi. developed cuneiform scolex. The scolex of Mackiewicz, J. 1965. Isoglaridacris bidbocirrus palearctic Canjophyllaeus Mueller, 1787, is gen. et sp. n. (Cestoidea: Caryophyllaeidae) from Catostomus commersoni in North Amer- basically of another type, however, being ex- ica. J. Parasit. 51: 377-381. panded with apical folds or fimbriae and com- Smith, G. R. 1966. Distribution and Evolution pletely lacking loculi. Comparative studies of of the North American Catostomid fishes of the cirrus complex of G. catostomi and C. the subgenus Pantosteus, genus Catostomus. terebrans are now in progress; hopefully they Misc. Pub. Mus. Zool., Univ. Mich. No. 129, should further elucidate the relationship of the 132 pp.

Echinostome Trematodes from Florida Birds

G. PREMVATI1 Department of Biology, Florida A. & M. University, Tallahassee, Florida

Two new species of trematodes of the sub- Seminal vesicle moderately developed, pars family Echinostomatinae were recovered from prostatica small. Ovary entire, pretesticular, two birds obtained from Leon County in postacctabular, submedian, 100-260 by 100- Florida. Also, 25 worms of an already known 120. Seminal receptacle present. Uterus small, species of Echinochasmus donaldsoni Beaver, pretesticular, has 10 to 15 eggs. Vitellaria 1941 were found in the intestine of two pied follicular; extends from anterior border of an- billed grebes, Podilymbus podiceps Linnaeus. terior testis, along sides of body, to posterior The worms were fixed in hot AFA under slight end of body; the two lateral fields join pos- coverglass pressure and stained in Semichon's terior to hind testis. Eggs measure 60-70 by carmine. 40-45. Measurements are in microns unless other- DISCUSSION: The genus Euparyphium Dietz, wise indicated. 1909 has a number of species, but only two species, namely, E. inerme (Ftihrmann, 1904) Euparyphium anhingae n. sp. Oclhner, 1910 and £. melis (Schrank, 1788) (Fig. 1) Railliat, 1919 have been reported from mammals from North America. From the host HOST: Anhinga anhinga Linnaeus. Anhinga, one species, E. capitaneiim Dietz, LOCATION: Intestine. 1909, is reported from Brazil and Habana. LOCALITY: Leon County, Florida. Euparyphium anhingae resembles E. capi- NUMBER OF WORMS: Twelve from one host. taneum considerably, but differs from it in the TYPE SPECIMENS: Holotype and two para- following: (1) in having 24 collar spines types in USNM Helm. Coll. Nos. 71148, 71149. (not 27); (2) in the absence of prepharynx; DESCRIPTION: Body large, spiny, measures (3) in having genital pore at level of intestinal 2.54-4.83 mm in length, and 540-600 in width bifurcation (not posterior to intestinal bifurca- in the region of ventral sucker, and 610-640 in tion), and (4) in the shape of testes. the region of testes. Body curved at ventral The shape of testes in E. anhingae resem- sucker in all specimens. Head collar with 24 bles that of E. inerme, but the position of spines; four spines on each ventral angle of testes in the former is in midbody while in the crown and sixteen spines in a continuous single latter it is in anterior half of body. Further, dorsal row. Size of largest spine 90-110 by E. anhingae differs from E. inerme in the 20-25. Oral sucker small, measures 70-85 by number and size of collar spines and in the 70-90. Ventral sucker large, diameter 340- absence of prepharynx. 440, and lies in one-fourth anterior part of body. Distance from anterior end of body to Petasiger floridus n. sp. ventral sucker 540-810. Prepharynx absent, (Figs. 2, 3) pharynx 135-160 by 80-110. Esophagus long, bifurcates anterior to ventral sucker; ceca long, HOST: Pied billed grebe, Podilymbus podi- terminating at posterior end of body. ceps Linnaeus. Testes elongated, lobed, tandem, in midbody LOCATION: Intestine. region. Anterior testis 400-580 by 250-340; LOCALITY: Leon County, Florida. posterior testis 370-580 by 230-310. Distance NUMBER OF WORMS STUDIED: Four from one from posterior border of posterior testis to pos- host. terior end of body 0.870-1.81 mm. Cirrus sac- TYPE SPECIMENS: Holotype and one para- elongated, posterior border near the middle of type in USNM Helm. Coll. Nos. 71150, 71151. ventral sucker; anteriorly cirrus sac extends to DESCRIPTION: Body fusiform, broadest at intestinal bifurcation and opens at genital pore. level of ventral sucker; spiny in preacetabular region. Body measures 1.58-2.08 mm in length 1 At present on leave from the Zoology Department, Uni- versity of Lucknow, Lucknow, India. and 530 to 640 in maximum width. Width of

Figures 1-3. 1. Euparyphiurn anhingae from Anhinga anhinga. Holotype: Dorsal view. 2. Petasiger floridus from Podilyrnbus podiceps. Holotype: Ventral view. 3. Petasiger floridus. Egg.

Copyright © 2011, The Helminthological Society of Washington OF WASHINGTON, VOLUME 35, NUMBER 2, JULY 1968 199 body at oral crown region is 270-330 and at with 27 collar spines as P. exaeretus Dietz, neck region 240-250. Preacetabular length 1909, and the other with 19-21 spines from varies from 700-810 and postacetabular from grebes as P. pungens (Von Linstow, 1894) 770-940. Fiihrmann, 1928 or P. megacanthus Kotlan, Oral sucker small, diameter 100; ventral 1922. Since then, P. inopinatum Baer, 1959 sucker large, diameter 370-400; ratio of two has been reported with 33 collar spines from suckers 1:3-1:4. Oral crown has nineteen Hagedashia hagedash Latham, 1790. This spines; four spines on each ventral angle and species does not fall into either of the groups four on lateral sides are larger than the three formed by Bisseru. spines on the dorsal side. Largest collar spine Of the 33 species of the genus Petasiger measures 135 in length, and the smallest 90. so far described from different parts of the Pharynx oval, measures 90 by 50. Esophagus world, only two, namely, P. nitidus Lin ton, long; ceca long and terminate at posterior end 1928 [described in detail by Beaver (1939)], of body. and P. chandleri Abdel-Malek, 1953, have been Testes oval, oblique in position, may be con- described from North America, both from tiguous or slightly apart; anterior one more grebes. Both the species are included by toward the right. Both testes measure 200- Skrjabin under the subgenus (Neopetasiger). 270 by 170-200. Cirrus sac large, slightly Petasiger floridus differs from both these in dextral, completely preacetabular, and crowds the following: (1) in having cirrus sac en- the intestinal bifurcation. Seminal vesicle di- tirely preacetabular; (2) pars prostatica very vided into a larger posterior portion and a long; (3) genital pore far behind the intestinal smaller anterior one; pars prostatica very bifurcation and immediately preacetabular; long; prostate glands very massive. Genital (4) presence of seminal receptacle, and (5) pore preacetabular, far behind the intestinal in the size and shape of eggs. bifurcation. Ovary submedian, postacetabular and pre- Summary testicular, measures 130 by 120. Seminal re- Two new and one known species of Echino- ceptacle and Laurer's canal present. Shell stome trematodes are described from Florida gland dorsal and lies either above or posterior birds. to anterior testis. Uterus short, dextral, with Echinochasmus donaldsoni Beaver, 1941 ascending limbs only. Vitellaria follicular, ex- were obtained from the intestine of two piecl tend from intestinal bifurcation or level of billed grebes, Podilymbus podiceps Linnaeus. cirrus sac into two lateral fields which join Euparyphium anhingae n. sp. from intestine posterior to hind testis, and fill entire posterior of Anhinga anhinga Linnaeus is characterized body space. Eggs few in number, and measure by the number of collar spines, absence of pre- 60-70 by 40-45. pharynx, shape of testes, and position of genital DISCUSSION: Bashkirova (1941) divided the pore. genus Petasiger Dietz, 1909 into two sub- Petasiger floridus n. sp. from the intestine genera: (Petasiger), type Petasiger (Peta- of Podilymbus podiceps is characterized by the siger) exaeretus Dietz, 1909 for species having position of cirrus sac and genital pore, presence tandem testes; and (Neopetasiger), type Peta- of seminal receptacle, long pars prostatica, and siger (Neopetasiger) skrjabini Bashkirova, by the size and shape of eggs. 1941 for species having oblique, or symmet- Acknowledgment rical or nearly symmetrical testes. Skrjabin (1956) followed the classification proposed by I express my sincere thanks to Professor Bashkirova and included all the species of the Robert B. Short for the use of his personal genus Petasiger under either of the two sub- library. genera. Literature Cited Bisseru (1957) does not mention anything Bashkirova, E. I. 1941. Echinostomatids of regarding the classification given by Bash- birds of the SSSR, and a review of their life- kirova. On the other hand, he has divided cycles. Trudy Bashkirsh Vet. Stants. 3: 243- the genus into two groups on the basis of 300. (In Russian: English summary 436- number of collar spines. He named the group 440.)

Beaver, P. C. 1939. The morphology and life Tijphlocoelium, Panjphostomum, and Peta- history of Petasiger nitidus Linton (Trema- siger. J. Helminth. 31: 173-186. toda: Echinostomidae). J. Parasit. 25: 269- Skrjabin, K. I. 1956. Trematodes of Animals 276. and Man. Principles of trematology. Izdatel- Bisseru, B. 1957. On three known trematodes stvo Akademii Nauk SSSR, Moscow. 12: from African birds, with notes on the genera 504-545. (In Russian.)

Onchobdella n. gen. New Genus of Monogenetic Trematodes (Dactylogyridae, Bychowski 1933) from Cichlid Fish from West Africa ILAN PAPERNA National Institutes of Health and Medical Research (Ghana Academy of Sciences), Accra, Ghana:;

Introduction and an accessory piece, consisting of two In the course of parasitological examination elongate flattened pieces, attached to each of fresh water fishes from different habitats other along one or two of their edges and in Ghana (Paperna, 1965), fish of the genus between these two planks the distal part of Hemichromis were found parasitized with the cirrus is protruded. Seminal vesicles and several allied forms of dactylogyrid monogenea 2—3 prostate glands follow the distal part of whose common morphological affinities sepa- the male genital system. Vagina present. rate them from all hitherto known genera of Anchors unequal in size and shape, one, the family Dactylogyridae. Fish caught by usually the larger pair, is followed by one hand net or seine net were placed alive into common solid bar, frequently curved, while vials containing 0.5-1% solution of formalin. each anchor of the second pair, which is After a few hours, when the fish had died, usually the smaller and hook-shaped, is fol- the concentration of the formalin solution was lowed by one club-shaped bar. Each of the increased to about 4% by the addition of a anchors is accompanied by complicated scle- few drops of formaldehyde. rotized filaments which are attached to the The worms were collected from the skin and anchor's shaft as well as lining the outer the gills of the fish as well as from the bottom margins of the aperture in the skin through of the vials. Worms so treated were usually which the anchors are drawn out. fully relaxed, and were immediately mounted Hooklets, 5-6 pairs, each consists of rudi- in Glycerine Gelatin for examination. Only mentary base, delicate long needle-shaped mounted specimens were measured. All mea- shaft and solid spike with delicate posterior surements given are in millimeters. projecting process. The large anchors in the opisthaptor are arranged distal-laterally with Onchobdella n. gen. their tips pointed inward, one against the other. Such position is relatively rare among GENERIC DIAGNOSIS: Body elongated or Dactylogyridae. More often anchors are located stout. Prohaptoral anterior lobes are poorly with their tips toward the lateroventral side of demarcated and head organs are present. the opisthaptor. Eyes, two or four; if four, they may be TYPE SPECIES: Onchobdella voltensis n. sp. arranged in one transverse row in front of the TYPE HOST: Hemichromis fasciatus. round pharynx. Intestinal crura are united LOCALITY: Afram sector, Volta Lake. posteriorly. The single testis is in the posterior REMARKS: Anchors located in similar posi- position within the intestinal loop, while the tion as in Onchobdella n. gen., i.e. lateral- single ovary is located anteriorly to the testis. clistally with tips pointed inward are charac- Copulatory organ consists of tubiform cirrus teristic also of few other "ancyrocephalid" genera such as Bifurcohaptor Jain 1958, Hama- * Present address: Embassy of Israel, P.O. Box 3275, Accra, Ghana. topeduncularia Yamaguti, 1953, Hargitrema

Figure 1: Onchobdella voltensis. a: Opisthaptor, anchors and bars, b: Additional sclerites associated with the erection mechanism of the anchors, c: Hooklets. d: Copulatory organ.

(Hargis, 1955) and a few species of the genus tached together form an ellipsoidal shape Dactylogyrus. Each of these forms, however, through which the cirrus is protruded. differs from Onchobdella in the details of the HOST AND LOCALITIES: Gills of Hemichro- morphological structures of the opisthaptor mis fasciatus, Afram sector, Volta Lake. Gills and its anchors, bars and hooklets. of Hemichromi-s bimaculatus, Afram sector and Kete Krachi, Volta Lake and Adutor Lagoon Onchobdella voltensis n. sp. in the Volta river "delta." (Figures 1 and 3b) SPECIMENS STUDIED: Six, from H. fasciatus DESCRIPTION: Small stout worms 0.20-0.35 and one from H. bimaculatus. long, 0.10-0.15 wide. Two to four eyes TYPE SPECIMENS: Type and paratype in the are present. Opisthaptor occupies the whole British Museum, London; other paratypes in posterior end of the body, 0.05-0.12 wide, the author's collection. 0.02-0.04 long. The pair of the larger anchors are 0.06-0.09 long with 0.01-0.015 long inner Onchobdella spirocirra n. sp. i-oot. The bar associated with these anchors (Figures 2 and 3a) is horseshoe-shaped, 0.07-0.10 long and 0.002- DESCRIPTION: Body stout, 0.28-0.50 long, 0.005 thick. The second pair of anchors are 0.15—0.25 wide. Only two eyes are present. 0.015-0.020 long and have well demarcated Opisthaptor is narrower than the entire body, inner and outer roots. 0.09-0.11 long, 0.08-0.09 wide. The attached bars are 0.02-0.04 long, then- The large anchors are 0.13-0.15 long with proximal thin end is not filiform as in the other species (see further). In the copulatory 0.02-0.25 long inner root. The horseshoe- organs the cirrus is a heavy short tube 0.03- shaped bar of the large anchors is 0.11-0.16 0.05 long with wide thick walled basal funnel. long, 0.010-0.012 wide. The anchors of the The accessory piece consists of two solid small pair are 0.02-0.03 long with well- 0.025-0.030 elongate sclerotized pieces and demarcated inner and outer roots. an additional median filamentous piece, at- Their attached club-shaped individual bars

15r

Figure 2. Onchobdella spirocirra. a: Anchors and bars, b: Copulatory organ, c: Hooklcts. d: Addi- tional sclerites of the large anchors. end proximally with a filiform process. Their filiform spirally coiled cirrus, originating from solid part is 0.05-0.07 long. Hooklets are a thin walled, round funnel and two elongated, 0.017-0.019 long with 0.008-0.009 long spike. triangular accessory pieces, 0.007-0.008 long, Copulatory organ consists of 0.05-0.07 long located at the tip of the cirrus.

.0 K) 3 3 T A

Figure 3. a: Onchobdella spirocirra (Dorsal view), b: O. voltensis (Dorsal view), c: O. pleri-gyalis (Dorsal view, O—ovary, T—testes, V—vaginal pore), d: O. krachii. e: Terminal male genital system of O. ptcrigyalis (A—Accessory piece, E—Cirrus, P—Prostate glands, S—Seminal vesicle).

HOST AND LOCALITIES: Gills of Hemichromis Onchobdella pterigyalis n. sp. bimaculatiis, from Mamahuma stream, east of (Figures 3c, e, 4a, b) Accra, and Afram sector, Volta Lake. DESCRIPTION: Large worms 0.50-0.75 long, SPECIMENS STUDIED: Three. 0.20-0.25 wide. Opisthaptor rudimentary, TYPE SPECIMENS: Type in the British Mu- inserted into the posterior end of the body, seum, London. Paratype in author's collection. 0.02-0.04 long, 0.03-0.04 wide. Four eyes

2OM.

Figure 4. Onchobdella pterigyalis: a: Anchors, bars and booklets, b: Copulatory organ. O. krachii: c: Anchors and bars, d: Hooklet. e: Copulatory organ. O. aframae: f: Anchors and bars, g: Hooklets. h: Copulatory organ. are present, located in one transverse row in is short. Hooklets are about 0.02 long with front of the pharynx. The "large" pair of 0.005 long spike. anchors is greatly reduced, 0.035-0.045 long In the copulatory organ the cirrus is 0.08- with. 0,005-0.008 long inner roots, only slightly 0.09 long, elongated tube with wide, thick- larger than the second "small" pair. walled funnel. The rim of the funnel is Their attached horseshoe-shaped bar is 0.05- strengthened by solid sclerotized processes. 0.06 long. The anchors of the second pair are The 0.02-0.03 long accessory piece consists 0.025-0.030 long, their inner and outer roots of two plates meeting at both ends, enclosing are well demarcated. Their individual bars are an ellipsoidal space. At one edge the accessory 0.032-0.040 long, their filiform prominal end piece is attached to the funnel. Vagina, with-

Copyright © 2011, The Helminthological Society of Washington OF WASHINGTON, VOLUME 35, NUMBER 2, JULY 1968 205 out sclerotized walls, opens on the left side of SPECIMENS STUDIED: Two. the body (when worm is placed on its dorsal TYPE SPECIMENS: Type in the British Mu- side). seum, London. Paratype in author's collection. HOST AND LOCALITY: Fins of young Hemi- chromis bimaculatus, Collected in Nungua Onchobdella aframae n. sp. dam and Mamahuma stream, east of Accra. (Figures 4f, g, h) The parasites were found only after the fish DESCRIPTION: Body elongated, 0.30-0.35 host were kept one month in the laboratory long, 0.05-0.10 wide. Only two eyes are pres- aquarium. ent. Opisthaptor is 0.05-0.06 wide and SPECIMENS STUDIED: Five. 0.01-0.02 long. The large anchors are 0.06- TYPE SPECIMENS: Syntypes in the British 0.07 long with 0.012-0.015 long inner roots. Museum, London. Paratypes in author's col- The associated straight bar is 0.03-0.04 long. lection. The small anchors are 0.02-0.03 long with The following two species differ from the well demarcated inner and outer roots. Their ones described above in having straight and individual bars are 0.02-0.03 long, their prox- not bent bar, attached to the large anchors. imal filiform processes are short. Hooklets are It is not certain whether this type of bar can about 0.01 long. be accepted as a specific character in these In the copulatory organ, the cirrus is a heavy species as they may be rather an immature straight tube, 0.05-0.06 long with a large form of the normal horseshoe-shaped bar. In round funnel. The 0.03-0.05 long, accessory the collected worms all the genital organs piece consists of two parallel rodlike sclerites were fully developed. However, only the pres- usually not jointed distally. ence of fully developed egg in the worms HOST AND LOCALITY: Gills of Hemichromis could safely indicate a fully mature stage of fasciatus, Afram sector, Volta Lake. development.* These forms are, however, de- SPECIMENS STUDIED: Three. scribed as separate species in view of the TYPE SPECIMENS: Type, in the British Mu- specific morphology of the copulatory organs. seum, London. Paratype in author's collection. REMARKS: The divergence of the species Onchobdella krachii n. sp. in this genus as evidenced by the morpho- (Figures 3d, 4c, d, e) logical variability is apparently not only asso- DESCRIPTION: Small, elongated worm, 0.3 ciated with adaptation to the different hosts long, 0.09 wide. Opisthaptor 0.06 wide, 0.04 but is rather more frequently associated with long, simple, poorly demarcated from the rest the adaptation to the different micro habitats of the body. The anchors of the "large" pair occupied within the organs of a single or few are 0.045-0.060 long, while their inner root is closely related host species. O. pterigyalis, 0.015-0.020 long. Their attached bar, is parasitizing the fins can serve as a significant straight 0.032 long, 0.004 wide. The anchors example, showing reduction in the size of the of the small pair are 0.025-0.030 long, their Opisthaptor and its sclerotinoid armature com- individual bars are 0.03-0.04 long, with only pared with those of the gill parasites. short filiform proximal processes. Hooklets are As can be seen from the table of ratios be- 0.01-0.012 long. In the copulatory organ, the low, the ratio between the lengths of the cirrus is a long and delicate tube, 0.08 long, two groups of anchors, the ratio between the arising from round, thin-walled funnel. Its total length of the large anchor and the length accessory piece, 0.02 long, consists of two of its inner root, as well as the ratio between sclerotinoid pieces attached to the funnel and the size of the worm and the size of the opis- another filamentous piece. All the three pieces thaptor are of importance in the diagnosis are jointed distally enclosing a triangular space. distinguishing the different species of this HOST AND LOCALITY: Gills of Pelmato- genus. Of these ratios, the large anchors to chromls guentheri, Kpandu and Kete Krachi, small anchors ratio is the most variable. With Volta Lake. the reduction in the size of the large anchors, * Recently mature specimens of O. aframae were col- there is also a parallel decrease in the size of lected containing egg in the uterus, but still having a straight bar as in the juvenile specimens. the opisthaptor, which becomes very reduced

Table 1. The ratio between the size of the different organs in species of Onchobdclla.

Ratio: in: O. voltensis O. spirocirra O. pterigt/alis O. krachii O. aframae 9-10 3-4.5 20-25 7.5 15 Body length to opisthaptor length 1 1 1 1 1 1-2 2-2.5 6-7 1,33 0.8-1.7 Body width to opisthaptor width 1 1 1 1 1 3-3.5 5—7 1.5 1.8-2 2-3 Large anchors to small anchors 1 1 1 1 1 1 1 1 1 1 Small anchor's bar to large anchors 2-3.5 3.5-5 1.1-1.2 1.9 3-4.9 1 1 1 1 1 Root of large anchors to large anchors 7-8 6 6-7 3 4-5 1.2-1.6 6-7 3-4 4 1.2-1,5 Cirrus to accessory piece 1 1 1 1 1 in O. pterigyalis. In addition, as in most of O. spirocirra, O. pterigyalis, O. krachii, O, the groups of Dactylogyridae, the morphology aframae). The hosts are cichlid fish of the of the copulatory organ is also characteristic genera Hemichromis and Pelmatochromis col- of each species. Here, the relative size of the lected in Ghana in small water reservoirs in cirrus decreases from O. spirocirra, through the coastal plain and in several sectors of the O. pterigyalis and O. krachii to O. voltensis Volta Lake. and O. aframae (Table 1). Literature Cited Summary Paperna, I. 1965. Monogenetic trematodes col- Five new species of monogenetic trema- lected from fresh water fish in Southern todes (Dactylogyridae) included into a new Ghana. Bamidgeh (Bull. Fish Cult. Israel) genus Onchobdella are described (O. voltensis, 17(4): 107-111.

New Host Records of Intestinal Nematodes of Maryland Rodents and Suppression of Capillaria bonnevillei Grimdmann and Frandsen, 1960 as a Synonym of C. americana Read, 19491

J. RALPH LiCHTENFELS2 AND A. JAMES HALEY*

Introduction North Carolina by Harkema (1936), Minne- Major surveys of nematode parasites of sota by Erickson (1938), and the north cen- rodents of the United States have been con- tral states by Rausch and Tiner (1948, 1949). ducted in the southwest by Hall (1916), More recently, studies have been done in Utah by Grundmann and Frandsen (1960a) and 1 Adapted from a thesis presented by the senior author Grundmann et al. (1961), Colorado and Idaho to the Department of Zoology, University of Maryland, in partial fulfillment of the requirements for the degree of by Leiby (1961, 1962), Alaska by Rausch Master of Science. This work was supported, in part, by the Natural Re- (1952, 1957, and others), and Nevada by sources Institute, University of Maryland, and by a National Science Foundation Summer Fellowship. Babero and Matthias (1967). In Maryland, - Present address: Beltsville Parasitological Laboratory, the nematode parasites of rodents have not Animal Disease and Parasite Research Division, ARS, USDA, Beltsville, Maryland. been extensively surveyed. Hall et al. (1955) :l Department of Zoology, University of Maryland, Col- lege Park, Maryland. Present address: Institute of Inter- examined six species of Maryland rodents for national Medicine, University of Maryland School of Medicine, Baltimore, Maryland. helminths and Price (1960) studied the filar-

Table 1. Intestinal ncmatodcs found in a survey of some rodents of Maryland (1963-1964). New host records are indicated by an asterisk (*).

Host No. examined No. infected Nematodes collected

CRICETIDAE Peroin yscus leucoptia 73 2 Capillaria americana 23 Rictnlaria coloradensis 1 Syphacta sp. 1 S. peromysci 1 Aspiculiiris tetraptera 1 A. americana

Ncotoma mogister 3 2 -•'Capillaria americana 2 ••'•Trichuris sp.

Clethrionomys gup peri 17 1 * Capilla ria a m ericana 6 HcUgmosomum carolinensis 1 Syphacia sp.

Microtus pennsylvanicus 15 6 Trichuris opaca 7 Longistriala dalrymplei 4 Syphacia sp. ZAPODIDAE Zapus hudsonius 5 3 ''•'Longistriata dtilrymplei 2 Citellinoides zapodis

SCIUIUDAE Tamias striatus 3 1 '"•Rictularia coloradensis Tamiasciurus li udsoii icii.v 15 3 '•'•'• Strongt/loides robustus 11 ••'Capillaria americana 7 Citellincma bifurcatum 1 Syphacia thompsoni MullIDAE Rattus norvcuicus 3 ] Mastophortis muris 3 Heteraki.i spnmosa Mtis musculiis 35 3 Mastophorus muris 22 Syphacia. obvclatn 23 Aspiculuris tetra ptcra ioid nematodes of some small mammals of stored in a deep freeze. The digestive tracts Maryland. In the present study, nine species of the rodents were opened in water, scraped of rodents from nine localities in five counties with a curved glass rod, and examined with of Maryland were examined for intestinal transmitted light under a dissection micro- nematodes. Seven of the nine rodent species scope. The nematodes were fixed in hot were collected entirely from three western alcohol-glycerine and cleared for study in a counties where apparently no previous surveys solution of 80% phenol and 20% absolute had been conducted. alcohol. Measurements are presented only when they provide new information and are Materials and Methods given in microns unless otherwise indicated. Collections were made with snap and box traps in the summers of 1963 and 1964. Red Results and Discussion squirrels were shot in March of 1964. The Sixteen species of nematodes representing identification of rodents was aided by museum six superfamilies were collected from 100 of specimens of the Inland Resources Division, 169 rodents and seven new host records are Natural Resources Institute, University of reported (Table 1). Brief discussions are given Maryland. Two-thirds of the rodents were of new data that supplement or correct descrip- examined the day of capture; the others were tions of species or establish new host records.

Superf amily Rhabditoidea errors in both descriptions. The spicule sheath Strongyloides robustus Chandler, 1942 of C. americana was described as smooth and HOST: Tamiasciurus hudsoniciis (Erxleben), the vulva as dividing the body 1:2.4 to 1:2.7. red squirrel. However, an examination of the cotypes dis- HABITAT: Small intestine. closed that the spicule sheath is spined and LOCALITY: Northeastern Garrett County, the vulva divides the body 1:3.4. The speci- Maryland. mens from the present collection agree with INTENSITY OF INFECTION: One host harbored the characters found in the cotypes of C. americana. 60 nematodes, two others harbored one each. SPECIMENS: USNM Helm. Coll. No. 65585. Grundmann and Frandsen (196()b) described C. bonnevillei from Dipodomys ordii MEASUREMENTS (based on 13 specimens): As from Utah and separated it from C. americana published by Chandler (1942) except: tail by the presence of spines on the spicule sheath, length 77-103 instead of 60-80; uterus to anus bilobate papillae on the caudal lobes of the 240-473 instead of 100-260. male, and the apparent absence of bacillary DISCUSSION: Strongyloides robustus was de- bands. However, bacillary bands are present scribed from Sciunis niger (type host) and in the holotype and allotype of C. bonnevillei S. carolinensi-s from southeast Texas. Dozier and, as pointed out above, the spicule sheath and Hall (1944) found it in S. niger from of C. americana is spined. The papillae on the Dorchester County, Maryland. The present caudal lobes of the males do appear to be report from Tamiasciurus hudsoniciis is the bilobate but are very similar in the two species. third of this species and a new host record. Therefore, a comparison of the measurements Superfamily Trichuroidea and morphological characteristics of C. amer- Capillaria americana Read, 1949 icana and C. bonnevillei, supplemented with data obtained from the present collection HOSTS: Tamiasciurus hudsoniciis, Red squir- (Table 2), leaves no doubt that C. bonnevillei rel; Neotoma magister Baird, woodrat; Pero- is a synonym of C. americana. mysciis leiicopus (Rafinesque), whitefoot mouse; Clethrionomys gapperi (Vigors), red- Trichuris sp. back vole. HABITAT: Posterior half, small intestine. HOST: Neotoma magister, woodrat. LOCALITY: Northeastern Garrett County, HABITAT: Caecum. Maryland. LOCALITY: Northwestern Allegany County, INTENSITY OF INFECTION: One to six speci- Maryland. mens per host. INTENSITY OF INFECTION: Fragments of SPECIMENS: USNM Helm. Coll. Nos. 65586 three females from one rat; one male from (red squirrel), 65587 (wood rat), 65588 another. (whitefoot mouse), 65589 (redback vole). SPECIMENS: USNM Helm. Coll. No. 66277. MEASUREMENTS: Table 2. DISCUSSION: This is a new host record for DISCUSSION: Capillaria americana was de- Trichuris sp. The limited material and its scribed from Glaucomys volans (type host), fragmented condition made determination Sciurus carolinensis, Peromyscus maniculatiis, difficult. The single male specimen differs in and P. leiicopus from the north central USA several characters from its most similar forms, (Read, 1949). The present report of C. amer- T. minis (Schrank, 1788) and T. madisonensis icana from the red squirrel, wood rat, and Tiner, 1950. Additional specimens are neces- redback vole constitutes three new host sary for a proper description. records. In the course of identification the senior Superfamily Strongyloidea author studied the type specimens of both Longistriata dalrymplei Dikmans, 1935 Capillaria americana and Capillaria bonne - HOSTS: Microtus pennsylvanicus (Orel), villei Grundmann and Frandsen, 1960. It meadow vole; Zapus hudsonius (Zimmerman), became apparent that differences between the meadow jumping mouse. descriptions of the two species were due to HABITAT: Small intestine.

Table 2. A comparison of (lapillaria americana Read, 1949 and C. bonnevillei Grundmann and Frandsen, 1960 including data from the original descriptions of the species, a restudy of the type specimens (data in parentheses), and specimens of C. americana from Maryland rodents. All measurements are in microns unless otherwise indicated.

C. americana C. bonnevillei C. americana Read, 1949 Grundmann and from Maryland rodents Frandsen, 1960 ( 12 specimens of each sex) Sex male female male female male female

Body length 12.2- 23.0- 12.3- 19.8 mm 9.3- 24.0- 15.4 mm 28.4 mm 16.0 mm 12.7 mm 32.5 mm Esophagus length 5.4—6.5 mm 6.8—7.5 mm 4.3—4.7 mm 4.6 mm 3.7—5.0 mm 4.7—7.8 mm Number of paraesophageal nuclei 41-45 36-39 36 36 38-47 37-50 Anterior end to first paraesophageal nucleus (418) (486) (370) (360) 300-384 328-520 Body diameter at base of esophagus (76) (100) (89) (129) 56-77 72-117 Maximum body diameter near middle of body 84-103 121-144 (113) (149) 65-110 115-144 Spicule length 209-258 240-300 182-234 Spicule width at base 11-14 (14) 12-17 Vulva to posterior end of esophagus (113) (50) 48-132 Egg length X width 46-52 X 41-59 X 50-62 X 23-27 20-29 24-30 Spined spicule sheath ( yes ) yes yes Baeillary band yes (yes) yes

LOCALITY: Northeastern Garrett County, bored 15 specimens; the other five harbored Maryland. one to three specimens each. INTENSITY OF INFECTION: Voles—30, 23, 7, SPECIMENS: USNM Helm. Coll. No. 65593. 1 or 2 in each of four others; Jumping mice— MEASUREMENTS: Esophageal length 600- 5, 1 in each of two others. 653; body diameter at base of esophagus 72- SPECIMENS: USNM Helm. Coll. Nos. 65591 91; greatest diameter of esophagus 34-52. (Microtus) and 65592 (Zapus). Male: Body length 5.02-5.43 mm; spicule DISCUSSION: This species was described length 1.90-2.01 mm; body diameter in pre- from Ondatra zibethica and Microtus penn- bursal region, 91-120. sylvanicus. Rausch and Tiner (1949) found Female: Vulva to posterior end body 137- it in voles from a single locality—a marsh in 192; tail length, excluding spine, 62; tail spine Wisconsin. In the present study the hosts length 10; ovejector length 173-259; body were trapped in the edge of a meadow bor- diameter, region of ovejector, 107-166; egg dering a swamp. The present report of L. length 60-65; egg width 36-47; ovejector to dalnjmplei from Zapus hudsonius is a new posterior end of body 657—735. host record. DISCUSSION: The present collection is only the third record of H. carolinensis, and it pro- Heligmosomum carolinensis vides supplemental data on the characteristics (Dikmans, 1940) of the species. The original collection by HOST: Clethrionomys gapperi, reclback vole. Dikmans (1940) from Clethrionomys gapperi HABITAT: Small intestine. in North Carolina included a single entire male LOCALITY: Western and northeastern Gar- and no entire females. An examination of the rett County, Maryland. cotypes revealed the entire male to be a INTENSITY OF INFECTION: One host har- fourth-stage larva. It is not surprising, there-

Copyright © 2011, The Helminthological Society of Washington 210 PROCEEDINGS OF THE HELMINTHOLOG1CAL SOCIETY fore, that males of this species collected by HABITAT: Anterior region, small intestine. Schad (1954) from the redback vole in Canada LOCALITY: Montgomery, Washington, Alle- were longer with normally (instead of weakly) gany, and Garrett Counties, Maryland. sclerotized spicules and longer bursal branches INTENSITY OF INFECTION: One to six speci- than the type material. The present collection mens per whitefoot mouse (including two contained a fourth-stage larva and other speci- male worms) and one female from the single mens intermediate in size between the type chipmunk host. material and that collected in Canada by SPECIMENS: USNM Helm. Coll. Nos. 65598 Schad. (P. leucopus) and 65599 (T. striatus). DISCUSSION: The single female Rictularia Superfamily Oxyuroidea coloradensis from an eastern chipmunk repre- Aspiculuris americana Erickson, 1938 sents a new host record. All specimens obtained in this collection conformed with the HOSTS: Peromyscus leucopus, whitefoot redescription of R. coloradensis given by Tiner mouse. (1948). However, the status of Tiner's re- HABITAT: Large intestine. description is unsettled. It was based on males LOCALITY: Allegany County, Maryland. with precloacal fans, but the type male of the INTENSITY OF INFECTION: Nine females and species (Hall, 1916) has no precloacal fans one male from the single host. and has much larger spicules than Tiner's SPECIMENS: USNM Helm. Coll. No. 65603. males. In this connection, McPherson and MEASUREMENTS: Male: Body length 2.73 Tiner (1952) reported both kinds of males mm; esophagus length 341; esophageal bulb from a single whitefoot mouse and Oswald length 115; esophageal bulb width 55; body (1958) reported both types of males in ex- diameter, base of esophagus 89; termination perimental infections in Mus musculus. Ex- of cervical alae to base of esophagus 144; tail periments using eggs from a single female length 233; most posterior papillae to cloacal worm may be necessary to solve this perplexing opening 72. problem of two types of males for one nema- Female: Body length 4.00-4.30 mm; esoph- tode species. Efforts are underway to provide ageal length 415-456; esophageal bulb length additional information. 139-154; esophageal bulb width 65-72; body diameter, base of esophagus 106-113; ce- Other Helminths phalic expansion length 84-91; termination of cervical alae to base of esophagus 194-238; Very few Platyhelminthes were encountered tail length 676-784; egg length 79-86; egg in this study. One unidentified trematode was width 29-36; vulva to anterior end of body recovered in a deteriorated condition from a 1.56-1.68 mm; excretory pore to anterior end whitefoot mouse. Cestodes were found in one of body 800. of three Norway rats, Rattus norvegicus DISCUSSION: This is the third report of (Berkenhout), from Allegany County and in Aspiculuris americana. It was described from two of 15 meadow voles from Garrett County. both Peromyscus leucopus and P. maniculatus Those from the Norway rat were identified as in Minnesota by Erickson (1938). The next Plymenolepis diminuta (Rudolphi, 1819) and those from the vole as Aprostatandnja macro- report was from P. floridanus in Florida by cephala (Douthitt, 1915). Layne (1963) who included no description of his specimens. The present report, therefore, Summary provides additional information about the range and measurements of A. americana. Sixteen species of intestinal nematodes were found in nine species of rodents collected in Superfamily Spiruroidea five counties of Maryland. Seven new host Rictularia coloradensis Hall, 1916 records are reported. Capillaria bonnevillei Grundmann and Frandsen, 1960 is placed in HOSTS: Peromyscus leucopus, whitefoot synonymy with C. americana Read, 1949 and mouse; Tamias striatus (Linnaeus), eastern errors in the description of C. americana are chipmunk. corrected.

Acknowledgments Hyracoidea. Proc. U. S. Nat. Mus. 50: 1- 258. The writers wish to thank Mrs. M. B. Harkema, R. 1936. The parasites of some North Chitwood for confirming the identifications of Carolina rodents. Ecol. Monogr. 6: 153-232. the nematodes; Dr. T. A. Bookhout, Mr. T. Layne, J. N. 1963. A study of the parasites of Theorig, and Mrs. C. Lichtenfels for help in the Florida mouse, Peromyscus floridanus, in collecting rodents; Mr. W. W. Becklund for relation to host and environmental factors. making arrangements for the study of speci- Tulane Stud. Zool. 11: 1-27. mens from the USNM Helminthological Col- Leiby, P. D. 1961. Intestinal helminths of some lection; and Professor L. A. Jachowski for Colorado mammals. J. Parasit. 47: 311. reviewing the manuscript. . 1962. Helminth parasites recovered from some rodents in southeastern Idaho. Amer. Literature Cited Midi. Nat. 67: 250. Babero, B. B., and D. Matthias. 1967. Proto- McPherson, S. E., and J. D. Tiner. 1952. A spirura peromysci n. sp. (Nematoda: Spiru- new nematode (Rictiilaria microti) from a ridea) and other helminths from Peromyscus vole on St. Lawrence Island, Alaska. Nat. spp. in Nevada. Proc. Helm. Soc. Wash. 34: Hist. Misc., Chicago Acad. Sci. (106): 1-7. 255-261. Oswald, V. H. 1958. Studies on Rictularia Chandler, A. C. 1942. Helminths of tree squir- coloradensis Hall, 1916 (Nematoda: Thela- rels in southeast Texas. T. Parasit. 28: 135- ziidae). II. Development in the definitive 140. host. Trans. Amer. Micr. Soc. 77: 413-422. Dikmans, G. 1940. A note on the genera Ne- Price, D. L. 1960. Epizootiological studies on matospiroides Baylis, 1926, and Sincosta Roe, some filaroid parasites of the family Dipeta- 1929 (Nematoda, Heligmosomidae), with de- lonematidae (Nematoda: Filarioiclea) found scriptions of two new speeies of Nemato- in certain small mammals. Diss. Abstr. 20: spiroides. Proc. Helm. Soe. Wash. 7: 79-82. 4222-4223. Dozier, H. L., and H. E. Hall. 1944. Obser- Rausch, R. L. 1952. Studies on the helminth vations on the Bryant fox squirrel, Sciiirus fauna of Alaska. XI. Helminth parasites of niger bryanti Bailey. Maryland Conserv. 21: microtine rodents—taxonomic considerations. 2-6. J. Parasit. 38: 415-444. Erickson, A. B. 1938. Parasites of some Min- . 1957. Distribution and specificity of nesota Cricetidae and Zapodiclae, and a host helminths in microtine rodents: Evolutionary catalogue of helminth parasites of native implications. Evolution 11: 361-368. American mice. Amer. Midi. Nat. 20: 575- , and J. D. Tiner. 1948. Studies on the 589. parasitic helminths of the North Central Grundmann, A. W., and J. C. Frandsen. 1960a. States. I. Helminths of Sciuridae. Amer. Definitive host relationships of the helminth Midi. Nat. 39: 728-747. parasites of the deer mouse, Peromyscus -, and . 1949. Studies on the para- maniculatus, in the Bonneville Basin of Utah. sitic helminths of the North Central States. II. J. Parasit. 46: 673-677. Helminths of voles (Microtus spp.). Prelim- , and . 1960b. Capillaria bonne- inary report. Amer. Midi. Nat. 41: 665-694. villei n. sp., (Nematoda: Trichuridae) from Read, C. P. 1949. Studies on North American the Ord kangaroo rat in Utah. Proc. Helm. Soc. Wash. 27: 206-208. helminths of the genus Capillaria Zeder, 1800 (Nematoda): II. Additional capillarids from -, and R. G. Warnock. 1961. A mammals with keys to the North American study of geographical isolation of parasitic nematodes on Stansbury Island, Great Salt mammalian species. J. Parasit. 35: 231-239. Lake. Proc. Utah Acad. Sci. 38: 74-77. Schad, G. A. 1954. Helminth parasites of mice Hall, J. E., B. Sonnenberg, and J. R. Hodes. in northeastern Quebec and the coast of 1955. Some helminth parasites of rodents Labrador. Canad. J. Zool. 32: 215-224. from localities in Maryland and Kentucky. J. Tiner, J. D. 1948. Observations on the Ric- Parasit. 41: 640-641. tularia (Nematoda: Thelaziidae) of North Hall, M. C. 1916. Nematode parasites of mam- America. Trans. Amer. Micr. Soc. 67: 192— mals of the orders Rodentia, Lagomorpha, and 200.

Ultrastructure of the Metacercarial Cyst of Ascocotyle chandleri Lumsden, 1963 (Trematoda: Heterophyidae)1 RICHARD D. LUMSDEN Laboratory of Parasitology, Department of Biology, Tulane University, New Orleans, Louisiana

Heterophyid trematodes of the genus Asco- obtained when the cysts were mechanically cotyle Looss, 1899, utilize various brackish ruptured during fixation, thus allowing sim- water fishes and, in some instances, larval ultaneous access of the monomer mixture to anurans as second intermediate hosts. The the inner and outer surfaces of the cyst wall. trematode larvae exhibit marked specificity The material illustrated in the present paper for a particular organ, and are nearly always was fixed in 3 per cent glutaraldehyde buffered found in close proximity to the circulatory at pH 7.3 in 0.12 M monobasic sodium phos- system of the host. The cercariae penetrate phate-sodium hydroxide, saturated with cal- the subendothelial tissues where they encyst cium chloride, and containing 3 per cent and undergo development to the preadult sucrose. After fixation for 2-4 hours at ice (metacercaria). water temperatures, the tissue blocks were Light microscopy studies on the encystment washed in cold phosphate buffer, postfixecl of Ascocotyle spp. in their second intermediate 1 hour with 1 per cent osmium tetroxide hosts have been described by Sogandares and (Millonig, 1961), dehydrated with ethanol and Lumsden (1963, 1964), Timon-David (1961), propylene oxide, and embedded in epon (Luft, and Schroeder and Leigh (1965). During the 1961). Thin sections (displaying silver inter- present investigation, the ultrastructure of the ference colors) were cut on diamond knives cyst wall of Ascocotyle chandleri and surround- with a Sorvall MT-2 ultramicrotome, collected ing host tissues has been examined by elec- on uncoated copper grids, double stained with tron microscopy as an initial step in determin- uranyl acetate and lead citrate, and examined ing the mechanism of the encystment process in a Siemens Elmiskop 1A electron microscope and the nature of the mature cyst. Previous operated at an accelerating voltage of 80 kv. electron microscope studies on metacercarial Thick (1 ^) sections for light microscopy were cyst structure and formation include those of mounted on microslides and stained with azure Dixon and Mercer (1964, 1967) and Mercer B. and Dixon (1967) on Fasciola hepatica Linn., 1758, and that of Macy et al. (1968) on Results and Discussion Sphaeridiotrema globulns (Rud., 1814). Encystment of Ascocotijle chandleri in the Methods and Materials liver of Cyprinodon variegatus is, as noted by Lumsden (1963), immediately subadjacent to Livers of naturally infected Cyprinodon the endothelium of the hepatic sinusoids (Figs. variegatus Lacepede were scanned with a 1-5). The metacercaria is surrounded by a dissecting microscope to locate the metacer- cyst wall approximately 15 ^ thick, consisting carial cysts. Areas of tissue containing the of a finely granular matrix of moderate electron cysts of A. chandleri were excised and pre- density (Figs. 5 and 9). This primary cyst pared for microscopic study by a variety of wall is bordered on its inner surface by a methods, including fixation with buffered layer of more coarsely filamentous material glutaraldehyde, acrolein, and osmic acid, and with adherent membrane-limited vesicles (Fig. embedment in epoxy resins or methacrylates. 7). The periphery of the cyst is marked by Considerable difficulty was encountered in- an electron opaque, bilamellate cortex (Fig. 6) filtrating intact cysts with sufficient plastic in direct contact with a lamina consisting of adequate to obtain thin sections of the material an amorphous matrix of variable thickness for electron microscopy; best results were containing occasional collagen fibers (Figs. 6 and 8). Adjacent to this lamina are one or 1 Supported by grants from NIH (GM 13330) and NSF more layers of endothelial cells (Figs. 1—6) (GB 7276), and with the technical assistance of Miss D. Umlauf. joined to one another by desmosomal junctions

Figures 1-2. Survey images of Ascocotyle chandleri metacercarial cyst and adjacent hepatic tissues. 1. Photomicrograph illustrating the cyst wall (CW) lined by a layer of endothelial cells (*). These endothelial cells border sinusoidal spaces (S) surrounding the hepatocytes (Hp). The sinusoids contain numerous ovoid erythrocytes with densely stained nuclei, x 2,000. 2. Corresponding electron micrograph of the tissue shown in Figure 1, illustrating part of the cyst wall (CW), endothelium (*), some erythrocytes (Er) and cellular debris in the sinusoidal space, x 5,4(10.

Figures 3-4. Electron micrographs illustrating the relationship of host tissues and the metacercarial cyst of A. chandleri. 3. Cells peripheral to the cyst wall (CW) frequently contain significant quantities of glycogen (*). The upper half of this picture is occupied largely by a hepatocyte, whose tangentially sectioned nucleus appears at N. An erythrocyte can be seen in the upper right corner of this micrograph. X 6,100. 4. The junction of two endothelial cells lining the periphery of the cyst wall is marked by a prominent desmosome (D). Part of the fusiform nucleus of the cell on the right is illustrated at N. X 24,000.

- t ,;*••;*>, - -,; *^?^'*ai •* Zi.'-.i'^i ^•:^W¥

*~ ..".>*

Figure 5. A section through the metacercarial cyst of A. chandleri. Illustrated are the inner marginal layer (IML) which borders the cavity occupied by the metacercaria, the outer marginal layer (OML), and bordering hepatic tissue, x 7,200.

•\\.~vr*i is**:-*",~-f--' -• 'a.** ».—-i •- .. • •'•••. • . •• t,**Y r«f--.'.».- ., .: -1

;r^;-, •:'. , '- ^ Ji.'.. .-' •* '"-':•• .'7- :,:'^- "•'''.•'.'•-"•'* liB^^/v ^jftf'.f*^-*.•;&. ^^^^s^^fcSi^'^^&^-ait >:••/', - •, '^ >.-: 'Vv.;:^-, , •>.:.i -I Figures 6-7. Details of the marginal structure of the metacercarial cyst of A. chcmdlerl. 6. The outer marginal layer, defined by a bilamellatc, electron opaque cortex (C and C'), and bordered by a basal lamina (BL) probably of endothelial origin. The plasmalemma of an adjacent endothelial cell is indicated by the arrows, x 80,000. 7. The inner marginal layer, consisting of a meshwork of filamentous material in which membrane bound vesicles (V) are embedded. These vesicles and the blobbed appearance of the marginal matrix (#) suggest that material of parasite origin is being added to the structure of the primary cyst wall. X 24,600.

Figures 8-9. 8. Collagen fibers (Co) in the basal lamina and adjacent tissue space proximal to the metacercarial cyst of A. chandleri. Part of an endothelial cell is visible in the upper right corner of this micrograph, x 66,600. 9. High magnification image of the matrix of the metacercarial cyst wall illustrating its finely granular texture, x 93,300.

(Fig. 4). These cells typically contain signif- Summary icant quantities of participate glycogen (Fig. Metacercariae of Ascocotyle chandleri encyst 3), a few small mitochondria, and a fusiform beneath the endothelium of the hepatic sinus- nucleus (Figs. 2, 4 and 5). The hepatic tissues oids of Cyprinodon variegatus. The primary surrounding these trematode cysts do not con- cyst wall consists of a finely granular matrix, tain an abnormal number of leukocytes or bordered internally by a meshwork of fibers other cell types associated with inflammatory- and vesicles and peripherally by an electron immunological response mechanisms. opaque, bilamellate cortex. The latter is in Collagenlike proteins have been indicated as contact with a lamina of amorphous ground components of metacercarial cysts of certain substance containing occasional collagen fibers. Ascocotyle spp. by staining properties of light A layer of endothelial cells line the periphery microscope preparations (Soganclares and of this cyst complex and border the adjacent Lumsden, 1963, 1964), and chromatographic sinusoidal space. The structure of A. chandleri analysis of cyst wall hydrolysates (Lenhoff metacercarial cysts is briefly compared with et al., 1960). Results of the present study, that previously reported for other trematodes. however, indicate that the cyst wall of A. chandleri does not contain morphologically Literature Cited identifiable collagen fibers, though such are sometimes present in the surrounding host Dixon, K., and E. Mercer. 1964. The fine tissue, including the basal lamina adjacent to structure of the cyst wall of the metacercaria the outer surface of the primary cyst wall of Fasciola hepatica. Quart. J. Micr. Sci. 105: 385-389. (Fig. 8). The possibility that the cyst matrix , and . 1967. The formation of contains a tropocollagenlike protein which is the cyst wall of the metacercaria of Fasciola not polymerized into the familiar 1,000 A hepatica. Z. f. Zellforsch. 77: 345-360. thick, cross-striated collagen fibers cannot be Lenhoff, H., R. Schroeder, and W. H. Leigh. evaluated at this time. 1960. The collagen-like nature of metacer- The majority of the host cells immediately carial cysts of a new species of Ascocotyle. adjacent to the metacercarial cyst of A. chand- J. Parasit. 46 ( Suppl.): 36. leri seem to be derived from the hepatic endo- Luft, J. 1961. Improvements in epoxy resin embedding techniques. J. Biophys. Biochem. thelium, though occasional fibroblasts have Cytol. 9: 409-414. also been identified in the tissues peripheral Lumsden, R. D. 1963. A new heterophyid to the primary cyst wall. trematode of the Ascocotyle complex of spe- The structure of the metacercarial cyst of cies encysted in poeciliid and cyprinodont A. chandleri thus differs from that described for fishes of southeastern Texas. Proc. Helm. Soc. cysts of certain other Ascocotyle species, nota- Wash. 30: 293-296. bly A. leighi Burton, 1956, and A. pachycystis Macy, R. W., A. Berntzen, and M. Benz. 1968. Schroeder and Leigh, 1965, which at the level In vitro excystation of Sphaeridiotrema gloh- of light microscopy appear to be multilayered, ulus metacercariae, structure of the cyst, and perhaps in part collagenous, and surrounded the relationship to host specificity. J. Parasit. 54: 28-38. by layers of fibroblasts (Soganclares and Lums- Mercer, E., and K. Dixon. 1967. The fine den, 1964; Schroeder and Leigh, 1965). The structure of the cystogenic cells of the cer- ultrastructure of the metacercarial cyst of A. caria of Fasciola hepatica L. Z. f. Zellforsch. chandleri differs significantly from that pre- 77: 331-344. viously described for other trematodes. The Millonig, G. 1961. Advantages of a phosphate metacercarial cyst of Fasciola hepatica, as de- buffer for osmium tetroxide solutions in fixa- scribed most recently by Dixon and Mercer tions. J. Appl. Phys. 32: 1627. (1967), consists of tanned proteins and a Schroeder, R., and W. H. Leigh. 1965. The keratinized protein, plus a carbohydrate-protein life history of Ascocotyle pachycystis sp. n., a trematode (Digenea: Heterophyidae) from complex, arranged in 4 distinct layers. Meta- the raccoon in south Florida. J. Parasit. 51: cercariae of Sphaeridiolrema globidus are 594-599. similarly enclosed by a cyst wall composed of Sogandares, F., and R. D. Lumsden. 1963. The four distinct layers or envelopes (Macy et al., generic status of the heterophyid trematodes 1968). of the Ascocotyle complex, including notes on

the systematics and biology of Ascocotyle Timon-David, J. 1961. Recherches sur la mor- angrense Travassos, 1916. J. Parasit. 49: phologic, le developpement experimental et la 264-274. systematique d'une metacercaire du genre , and - . 1964. The heterophyid Ascocotyle Looss, 1899 (Trematoda, Digenea, trematode Ascocotyle (A.) leighi Burton, Heterophyidae) parasite des branchies chez 1956, from the hearts of certain poeciliid and les tetards de Rana esculenta L. Ann. Parasit. cyprinodont fishes. Z. f. Parasitenk. 24: 3-12. Hum. Comp. 36: 737-751.

Revision of the Genus Radopholus Thorne, 1949 (Nematoda: Tylenchoidea) S. A. SHER Department of Nematology, University of California, Riverside, 92502

The genus Radopholus Thorne, 1949 was tology, University of California, Riverside, or proposed in order to include those Pratylen- as indicated in this paper. ehinae most closely resembling the genus Pratylenchus Filipjev, 1934 but which have Morphology two ovaries. Radopholus similis (Cobb, 1893) Species of Radopholus are typically cylin- was named the type species, and one other drical in shape and vary in length as adults species R. onjzae (van Breda de Haan, 1902) from 0.4-0.9 mm. In at least one species was included in the genus Radopholus at that (R. similis) some females from the roots of time. In 1955, Allen transferred three species plants appear larger and fatter than specimens from the genus Tylenchorhijnchus Cobb, 1913 collected from soil around these roots. Mor- to the genus Radopholus [R. gracilis (de Man, phological differences are seen between the 1880), R. behningi (Micoletzky, 1927), and females, males, and juveniles of most of the R. zostericola (Allgen, 1934)]. Two species, species. This is most marked in the lip and R. neosimilis Sauer, 1958 and R. inaeqiialis esophageal region of the female and male. Sauer, 1958, were described from a native The latter having a higher, rounder, distinctly habitat in Australia. set off lip region with reduced labial frame- In 1962, Luc and Goocley proposed the work, stylet, stylet knobs, and esophagus. The genus Hirschmannia for the large species in female and juvenile have a hemispherical to Radopholus with long overlapping ventral flattened lip region with the labial framework esophageal glands. This left the type species, well developed, strong stylet and stylet knobs R. similis and the two species of Sauer from with rounded, flattened anteriorly or forward Australia in Radopholus. An additional spe- pointed projections. The glandular posterior cies was proposed, R. ivilliamsi Siddiqi, 1964, portion of the esophagus overlaps the intestine using a population described from the Mauri- dorsally and laterally and is well developed tius by Williams (1960) as R. similis. and conspicuous in the female and juvenile, This study is based on over 5,000 preserved extending almost to the center of the body specimens fixed usually in formalin (2-5%); in some species with the three lobes often FAA (formalin, alcohol, glacial acetic acid, distinctly visible (b' = length of the body and water); or TAP (formalin, triethylamine, divided by the length of esophagus to the and water); then transferred to diluted glyc- end of the esophageal glands). The median erine, allowed to dehydrate slowly, and bulb and posterior part of the esophagus are mounted in dehydrated glycerine. The rapid reduced and often indistinct in the male. In glycerine technique of Baker (1953) was used face view, the female and juvenile are round for some specimens. with six similar lips, the lateral lips set off by All the specimens studied are in the Nema- longitudinal striations in some species. The tode Collection of the Department of Nema- male face view is squarish, the lips are set off

Copyright © 2011, The Helminthological Society of Washington 220 PROCEEDINGS OF THE HELMINTHOLOGICAL SOCIETY and the lateral lips are greatly reduced. In portion of stylet as long or longer than pos- addition to being smaller, the juveniles usually terior portion. Deirids absent. Esophageal differ from the adults in having a shorter glands overlapping intestine dorsally. Gonads hyaline area between the cuticle and the body amphidelphic. Phasmids usually in anterior contents in the tail terminus. This hyaline part of tail. Tail tapering to a rounded or al- area is conspicuous in all the adult specimens most pointed terminus, usually 2-4 times as except one. The length of the hyaline area is long as the body width at the anus or cloaca. given in the measurements of each species as Gubernaculum protruding from cloaca. the letter H (c' = the length of the tail divided TYPE SPECIES: Radopholus similis (Cobb, by the body width at the anus or cloaca). 1893). Juveniles also often have fewer incisures in Key to the Species of Radopholus the lateral field than adult specimens. The number of incisures vary from three to seven 1. Phasmids in posterior part of tail; in this genus; a variation in number of incisures hyaline area in terminus 3 p. or also occurs in some of the species. The inci- less nigeriensis n. sp. sures themselves may vary, being incomplete Phasmids in anterior part of tail; and/or irregular in some species. Incomplete hyaline area in tail terminus 3 ^ or areolation of the posterior part of the lateral longer 2 field has been observed in many of the species. 2. Spermatheea with round sperms The female reproductive system consists of rotiindisememis n. sp. opposed gonads with the vulva located at Spermatheea with rod-shaped sperms 50-70%. The ovaries are usually outstretched or sperms absent 3 but in some large females obtained from roots, 3. Lateral field with 5 or more incisures the anterior ovary extends to the esophageal inaequali'i region, and the posterior ovary to the tail Lateral field with 4 or less incisures ... 4 where they can be reflexed. The ovary oc- 4. Female tail without annules around cupies from 12—50% of the body length. A terminus wttliamsi spermatheea with rodlike sperms is seen in Female tail with annules around ter- most species, one described species has round minus 5 sperms in the spermatheea, and in three spe- 5. Esophageal glands of females termi- cies the spermatheea is inconspicuous or ab- nating in anterior third of the body sent and no sperms are visible. The rodlike (b' = 3.2-5.2) .. 7 sperms can also be observed in the testes. Esophageal glands terminating pos- Dense spherical granules and irregularly teriorly to one-third of the body looping lateral canals are usually seen in the length (b' = 2.2-3.2) 6 intestinal region. They are often overlapping 6. Adult lateral field with three incisures; the rectum into the tail. vulva at 64% or more anterior The excretory pore is opposite the region of trilineatus n. sp. the esophageal intestinal valve and the hemi- Adult lateral field with four or more zonid immediately anterior to the excretory incisures; vulva at 65% or more pos- pore. The hemizonion, cephalids and caudalid terior magniglans n. sp. are inconspicuous or not seen in most 7. Female stylet 20 ^ or longer; 4 or specimens. more annules on lip region nativus n. sp. Genus Radopholus Thorne, 1949 Female stylet 20 /JL or shorter; 4 or less Radopholus Thorne, 1949, p. 53; Luc and annules on lip region 8 Goodey, 1962, p. 199. 8. Female terminus \vith short hyaline DIAGNOSIS EMENDED: Radopholinae Allen area (5 ^ or less) and broadly and Sher, 1967. Labial framework and stylet rounded with distinct annulation ____ well developed in female and juveniles. Sex- neosimilis ual dimorphism marked; the male has higher Female terminus with conspicuous hya- rounder lip region and lateral lips, labial frame- line area (6 /j. or longer) and nar- work, stylet, and esophagus reduced. Anterior rowly rounded or almost pointed

with indistinct and/or irregular an- 5 $ topotypes: L = 0.63 mm (0.59-0.67); nulation 9 a = 35 (31-44); b = 6.4 (6.1-6.6); b' = 4.8 9. Female stylet 17 p. or less; sperms ab- (4.1-4.9); c = 9 (8-10); c' = 5.7 (5.1-6.7); sent in spermatheca H = 9 p. (7-11); stylet = 14 p. (12-17); vertexplanus n. sp. gubernaculum = 9 p, (8-12); spicules = 20 p. Female stylet 17 p, or longer; sperms (19-22). present in spermatheca 10 SUGAR CANE, HAWAII: 132?: L = 0.68 10. Female tail 3.2 times as long as the mm (0.59-0.82); a = 26 (22-35); b = 6.9 body width at anus or longer with (5.7-7.8); b' = 4.6 (3.9-6.5); c = 9.7 (8- hyaline area 9 p, or longer slmilis 11); c' = 3.6 (2.9-4.1); H = 13 ^ (10-17); Female tail 3.1 times as long as the v - 20 (is-50) 56 (52-60) 2(i do-41); stylet = body width at anus or shorter with 19 p. (17-20); O = 20 (15-25). hyaline area 9 p. or shorter 4$ $: L = 0.62 mm (0.60-0.64); a = 35 vangundyi n. sp. (30-43); b = 5.8 (4.7-6.8); b' = 4.6 (3.9- 5.0); c = 9 (8-10); c' = 5,5 (4.8-6.3); H = Radopholus similis (Cobb, 1893) 9 fj, (7-11); stylet = 14 ^ (12-17); guber- Thome, 1949 (Fig. 1 A-N) naculum = 11 p, (9-12); spicules = 21 p. Tylenchus similis Cobb, 1893, p. 301; Cobb, (18-23). 1915 pp. 563-567; Bally and Reydon, 1931, FEMALE: Lip region hemispherical, slightly pp. 80-86; Steiner and Buhrer, 1933, pp. 415- or not set off with three to four annules; six 416. similar lips, lateral lips slightly set off (face Tylenchus graniilosus Cobb, 1893, pp. 300- view). Stylet knobs rounded to slight pro- 301, new synonymy. jections anteriorly. Spermatheca with rodlike Tylenchus acutocaudatus Zimmermann, sperms. Lateral field with four incisures, 1898, pp. 42-43. coalescing to three incisures near middle of Tylenchus brformis Cobb, 1909, pp. 63-66. tail. Lateral field sometimes incompletely Anguillulina similis: Goodey, 1932, pp. areolated on tail. Intestine indistinctly over- 38-40. lapping rectum into tail. Tail tapering, Anguillulina granulosa: Goodey, 1932, p. rounded variable terminus with or without 92, new synonymy. annulation. Tylenchus (Chitinotylenchus) similis: Mico- MALE: Lip region high, hemispherical, dis- letzky, 1922, pp. 543-547. tinctly set off, with three to five annules in the Tylenchorhynchus similis: Filipjev, 1934, posterior region; lateral lips distinctly smaller p. 142. (face view). Tail tapering to rounded or Tylenchorhynchus acutocaudatus: Filipjev, almost pointed terminus; caudal alae not en- 1934, p. 142. veloping tail. Distal ends of spicules pointed. Bitylenchiis graniilosus: Filipjev, 1934, p. Gubernaculum with small titillae. 152, new synonymy. JUVENILE : Similar to female except for three Rotijlenchus similis: Filipjev, 1936a, p. 545; incisures in the lateral field (Fig. 1 G—H) and Filipjev and Sch. Stekhoven, 1941, p. 211. shorter hyaline area in tail terminus (2—4 p,). Teti/lenchus graniilosus: Filipjev, 1936b, p. TYPE HOST AND LOCALITY: Soil about dis- 81; Filipjev & Sch. Stekhoven, 1941, p. 258, eased banana plants, Fiji. new synonymy. DIAGNOSIS: R. similis can be identified by Radopholus similis: Thome, 1949, pp. 53— the female with three to four annules on the lip region, stylet 17—20 p., long hyaline area in 54; van Weerdt, 1958, pp. 191-193; Thorne, tail terminus (H = 9-17 yu,), and male with 1961, pp. 226-227. long narrow tail (c' — 4.8-6.7). MEASUREMENTS: 12 2 topotypes: L — The description, illustrations and measure- 0.69 mm (0.52-0.88); a = 27 (22-30); b = ments are based on specimens collected from 6,5 (4.7-7.4); b' = 4.5 (3.5-5.2); c = 10.6 the type host and type locality (topotype) of (8-13); c' = 3.4 (2.9-4.0); H = 12 ,j, (9- R. similis and from the roots and soil of sugar 16); V = ~9 (19^41) 56 (55-61) 27 (17-38). styiet cane, Oahu, Hawaii (type host and near type = 19 n (17-20); O = 18 (12-20). locality of T. biformis).

A-D.G-Q

^ E-F M N ' Figure 1. Radopholus similis: A-B. Female, anterior ends. C-D. Male, anterior ends. E. Female, face view. F. Male, face view. G. Juvenile, cross-section near center of body. H. Juvenile, lateral view near center of body. I. Female, cross-section near center of body. J. Female, lateral view near center of body. K-M. Female, posterior regions. N. Male, posterior region. Radopholus williamsi: O. Female, anterior end. P. Male, anterior end. O. Male, posterior region.

Because of the original descriptions, mea- Roseau Valley, St. Lucia; St. Vincent; Sin- surements, and habitat and after an examina- ceretz Estate and Ashenden, Grenada; Neba tion of specimens from the type host and Valley, Newfoundland and Woodford Hill, locality, there is no doubt that Tijlenchus Dominica; Vinales, Cuba; Tiquisate, Guate- similis Cobb, 1893 and Tijlenchus granulosus mala; Santa Tecla, El Salvador; Perez, Do- Cobb, 1893 are the same taxon. Cobb's 1893 minican Republic; Golfito, Costa Rica; Blue- publication contained a description of Tijlen- field, Nicaragua; Santa Marta, Colombia; Sao chus granulosus, based on female specimens Paulo, Brazil; Los Angeles, California; and from diseased banana roots and adjacent soil, Oslo, Florida. on page 300 and 301 (without an illustration); Additional specimens have been examined on page 301, the name T. similis was proposed from the following habitats and localities: on male specimens found around diseased citrus (Citrus sp.) roots, Lake Alfred, Orlando banana plants (without a description but with and Alturas, Florida; avocado (Persea amer- illustrations). On the basis of page priority, icana] roots, Lake Placid and Avon Park, T. granulosus is the older name for this taxon. Florida; anthurium (Anthuriiim sp.) roots, Although T. granulosus has been placed in a Hilo, Pohoa and Manoa Valley, Hawaii; number of genera, there is no evidence that Sterlitzia reginae roots, Hilo, Hawaii; Zingiber this was done on identifying actual specimens officinale roots, Kurtistown, Hawaii; Hedych- with this name, and this taxon is now re- ium flavum roots, Keaau, Hawaii; Shell ginger corded as Tetijlenchus granulosus (Cobb, (Alpinia mutans) roots, Nuuanu Valley, 1893) Filipjev, 1936. To preserve the well- Hawaii, sugar cane (Saccharum officinarum) known name of an important, widely dis- roots, Oahu, Hawaii; and Philodendron haus- tributed economic pest, it is necessary to tatum roots, Puerto Rico. regard T. granulosus as a senior synonym or nomen oblitum (forgotten name). Radopholus Radopholus williamsi Siddiqi, 1964 similis is therefore considered in this paper as (Fig. 1 0-Q) a valid name and T. granulosus a senior Radopholus similis: Williams, 1960, pp. 14- synonym. 15 (in part). Tijlenchus acutocaudatus Zimmermann, 1898 Radopholus icilliamsi Siddiqi, 1964, pp. described from coffee roots in Java was syn- 207-208. onymized with T. similis by Menzel (1929) MEASUREMENTS: 6 9 paratypes: L = 0.42 after studying specimens from tea (Thea mm (0.40-0.44); a = 21 (19-23); b = 5.1 sinensis) in Java. Steiner and Buhrer (1933) (4.6-6,5); b' = 3.0 (2.8-3.6); c = 13 (11-16); confirmed the synonymy of this species after c' = 2.4 (2.2-2.7); H = 7 ^ (5-10); V = an examination of specimens received from 10 (u-18) 64 (62-69) 1(i ("-is). styiet = 15.5 p. Menzel from the same host and locality. Addi- (15-16); O = 15 (11-21). tional reports of R. similis in Indonesia, where 6 $ paratypes: L = 0.41 mm (0,37-0.43); it is the cause of yellows disease of pepper, a = 25 (21-27); b = 6.0 (5.5-6.8); b' = 4.0 have been made by Thome (1949, 1961), (3.9-4.3); c = 12 (11-13); c' = 3.2 (3.0- van der Vecht (1950), and Christie (1957). 3,5); H = 11 ^ (9-13); stylet = 10^ (9-11); Although specimens of R. similis from Indo- gubemaculum = 10 p. (8-11); spicules = 18 ^ nesia have not been available for study, there (17-20). is no reason to doubt the synonymy of T. FEMALE: Lip region hemispherical, not set acutocaudatus. off with three to four annules; similar to type Specimens of R. similis have been examined species in face view. Stylet knobs rounded to from the soil or roots of banana (Musa sp.) slightly flattened anterior surfaces. Sper- from the following localities: Government matheca with rodlike sperms. Lateral field House, Suva and Viti Suva, Fiji; Selangor, with three or four incisures. Intestine indis- Malaya; Malifanua Hill, Western Samoa; Hilo, tinctly overlapping rectum into tail. Tail Hawaii; Bundaberg, Queensland, Australia; tapering, rounded terrain vis without annulation. Quezon City, The Philippines; Ilogbo Village, MALE: Lip region spherical, distinctly set Colony Province, Nigeria; Kade, Ghana; off, with four to five annules; similar to type Kisimaio, Somaliland; Spanish Town, Jamaica; species in face view. Tail narrowing to a

Figure 2. Radopholus neosimilis: A. Female, anterior region. B. Female, longitudinal section at center of body. C. Female, tail region. D. Male, anterior end. E. Male, posterior region. Radopholus inaequalis: F. Female, anterior end. G. Male, anterior end. H. Male, posterior end. I. Female, surface view near center of body. J. Female, cross-section near center of body. K. Female, tail region.

Copyright © 2011, The Helminthological Society of Washington OF WASHINGTON, VOLUME 35, NUMBER 2, JULY 1968 225 slender posterior portion; caudal alae almost anteriorly, distinctly set off, with four or five enveloping tail. Distal ends of spicules needle- annules; similar to type species in face view. like. Titillae not seen on gubernaculum. Tail tapering to a rounded or almost pointed JUVENILE: Similar to female except for terminus. Caudal alae enclosing tail. Distal shorter hyaline area in tail terminus. ends of spicules pointed. Gubernaculum with TYPE HOST AND LOCALITY: Sugar cane small titillae. (Saccharum officinarum) roots, L'Etoile, JUVENILE: Similar to female except for Mauritius. three incisures in lateral field and slightly TYPE SPECIMENS: Female holotype and shorter hyaline area in tail terminus (2-4 //,). male allotype collected by J. R. Williams, TYPE HOST AND LOCALITY: Soil around roots 1959; deposited at Rothamsted Experimental of Codonocarpus cotinifolius in virgin area Station, Harpenden, England. Paratypes (6 near Lake Mournpoul, Hattah, Victoria, Aus- 99, 4$ $ ) are in the personal collection of tralia. R. Siddiqi (1964) and in the collection of the TYPE SPECIMENS: Female holotype slide Department of Nematology, University of Radopholus 3, male allotype slide Radopholus California (99 ?, 5$ $). 4, collection of Commonwealth Research Sta- DIAGNOSIS: R. williamsi can be distin- tion, Merbein, Australia. guished from the type species, R. similis, by DIAGNOSIS: R. neosimilis can be distin- the smaller body size, shorter stylet, shorter guished from closely related R. williamsi by tail, and caudal alae almost enveloping the the more anterior position of the vulva, flat- tail. tened lip region, shorter hyaline area in tail In addition to the paratypes studied (sup- terminus and usually longer stylet and greater plied by J. R. Williams), specimens of R. body length. williamsi have been examined from sugar cane soil in the Fiji Islands. Radopholus inaequalis Sauer, 1958 (Fig. 2 F-K) Radopholus neosimilis Sauer, 1958 Radopholus inaequalis Sauer, 1958, pp. 100- (Fig. 2 A-E) 103. Radopholus neosimilis Sauer, 1958, pp. 103- MEASUREMENTS: 10 9 topotypes: L = 0.64 106. mm (0.53-0.78); a = 25 (20-31); b = 7.5 MEASUREMENTS: 17 9 topotypes: L = (5-10); b' = 5.2 (3.1-6,3); c = 17 (15-18); 0.66 mm (0.60-0.74); a = 30 (26-33); b = c' = 2.3 (1.8-2.9); H = 8 p. (5-10); V = 7.1 (6-8); b' = 4.5 (3.4-4.9); c = 15 (12-17); 20 (18-34) 59 (55_64) 24 (17-28). stylet - £2 p. c' = 2.5 (2.0-3.1); H = 4 p. (3-5); V = (21-23); O = 15 (12-19). 25 (21-28) 58 (55-60) " (20-20). stylet = 17/4 8 $ topotypes: L = 0.59 mm (0.52-0.64); (16-19); O = 16 (14-18). a = 30 (28-33); b = 7.2 (6.1-8.6); b' = 5.1 4 $ topotypes: L = 0.56 mm (0.51-0.65); (3,5-5.7); c = 14 (12-16); c' = 2.9 (2.6- a = 31 (26-35); b = 7.7 (6.4-8.4); b' = 5.0 3.3); H = 8^ (6-10); stylet = 14 p. (13-16); (4.7-5.3); c = 14 (12-15); c' = 3.2 (2.9- gubernaculum = 12 p, (11-13); spicules = 3.7) ;H = 7/* (5-11); stylet = 14 p. (12-16); 23 p. (21-25). gubernaculum = 10 p, (8-12 //,); spicules = FEMALE: Lip region hemispherical, slightly 20 p (19-23). or not set off, with three annules; similar to FEMALE: Lip region flattened anteriorly, type species in face view. Stylet knobs with rounded edges, slightly set off, with three or pointed anterior surfaces, often unequal in four annules; six similar lips without longi- size. Rodlike sperms (not seen in all speci- tudinal striations in face view. Stylet knobs mens) incompletely filling spermatheca. Lat- rounded, often with flattened anterior surfaces. eral field with five to seven incisures, some- Rodlike sperms (not seen in all specimens) times incompletely aerolated on tail. Intestine incompletely filling spermatheca. Lateral field extends into tail. Tail tapering slightly to with four incisures. Intestine extends into tail. rounded, annulated terminus. Tail tapering slightly to rounded, annulated MALE: Lip region oblate, distinctly set off terminus. without annulation; similar to type species MALE: Lip region high, slightly flattened in face view. Lateral field with five or usuallv

'•9\I

i":'.-:-:'.o;;oT tei£S/ill

ily-s.- f>£!Qy

}/««ill

0.02 mm

Figure 3. Radopholus vangundyi n. sp.: A. Female, anterior region. B. Female, longitudinal section at center of body. C. Female, tail region. D. Female, terminus. E. Male, tail region. F. Male, anterior region.

Copyright © 2011, The Helminthological Society of Washington OF WASHINGTON, VOLUME 35, NUMBER 2, JULY 1968 227 six incisures. Tail tapering to a rounded or pletely filling spermatheca. Lateral field with almost pointed terminus. Distal ends of spic- four incisures. Intestine indistinctly overlap- ules pointed. Caudal alae enclosing tail. ping rectum into tail. Tail tapering to almost Gubernaculum with small titillae. pointed, annulated terminus. JUVENILE : Similar to female except for three MALE (allotype): L = 0.50 mm; a = 29; or four incisures in lateral field and shorter b = 6.1; b' = 3.8; c = 12; c' = 3.6; II = 10 p.; hyaline area in tail terminus (2—7 p,). stylet = 13 yu,; gubernaculum = 8 p.; spicules TYPE HABITAT AND LOCALITY: Soil around = 15 p.. Lip region oblate, distinctly set off, roots of Codonocarpus cotinifolius in virgin without annulation. Stylet knobs with sloping area near Lake Mournpoul, Hattah, Victoria, anterior surfaces. Median bulb with valve. Australia. Caudal alae narrow, not enveloping tail. TYPE SPECIMENS: Female holotype slide JUVENILE: Similar to female except for a Radopholus 1, male allotype slide Radophohis shorter hyaline area in tail terminus (2-3 p.). 2, collection of Commonwealth Research Sta- HOLOTYPE: Female collected by S. D. Van tion, Merbein, Australia. Gundy, 20 February 1966, catalog number 1, DIAGNOSIS: R. inaequalis can be distin- U.C.R. Nematode Collection, Riverside, Cali- guished from R. similis and R. ivilliamsi by fornia. the longer stylet with pointed anterior pro- ALLOTYPE: Male, same data as holotype, jections on the stylet knobs and five or more catalog number 2. lateral incisures in the lateral field. PARATYPES: 37 9 5 , 30 $ $ , 9 juveniles same In a collection from the type locality of data as holotype distributed as follows: 399, R. inaequalis and R. neosimilis (supplied by 355, Department of Nematology, University M. R. Sauer) there appears to be two addi- of California, Davis; 25 9 $, 17 $ $, 8 juve- tional unclescribed species of Radopholus. niles, Department of Nematology, University These additional species in the same collection, of California, Riverside; 3 9 9 , ~4 $ $, USDA the sexual dimorphism of the species and the Nematode Collection, Nematology Investiga- scarcity of males (only two and three speci- tions, Beltsville, Maryland; 399, 3$ $, 1 mens of each species reported by Sauer in juvenile, Nematology Department, Rotham- 1958) makes the specific identification of sted Experimental Station, Harpenden, Eng- males for these species a more difficult prob- land; and 399, 3 $ $, Plantenziektenkundige lem than usually encountered. Although Dienst, Wageningen, The Netherlands. Sauer's identification of the males appears to TYPE HABITAT AND LOCALITY: Eucalyptus be correct, additional data would be desirable. sp., Acacia sp. and unknown grass soil in native habitat, bank of the Mt. Williams River Radophohis vangundyi n. sp. (near Dadswells Bridge), Victoria, Australia. (Fig. 3 A-F) DIAGNOSIS: R. vangundyi can be distin- MEASUREMENTS: 15 5 paratypes: L = 0.54 guished from the closely related R. neosimilis mm (0.47-0.65); a = 27 (22-30); b = 7.0 by the hemispherical female lip region shape, (6.3-8.1); b' = 3.7 (3.3-4.0); c = 14 (12- more tapering female tail, usually smaller fe- 16);c' = 2.7 (2.4-3.2); H = 6.5^ (5-9); V = male body size and the higher lip region with- i>4 (20-80) 59 (57-62) 21 (i8-24); stylet = 18 p. out annulation in the male. (17-19); O = 15 (11-24). In face view the female has six similar lips, 10 $ paratypes: L = 0.52 mm (0.41- the lateral lips slightly set off (similar to the 0.63); a = 31 (28-35); b = 7.3 (6.1-7.7); type species); amphid apertures are seen on b' = 4.2 (3.6-4.6); c = 13 (11-15); c' = 3.1 the lateral lips. In the male face view the (2.7-3.6); H = 9 p, (6-12); stylet = 13 p. lips are set off with the lateral lips distinctly (12-15); gubernaculum = 7 p. (6-10); spic- smaller (similar to the type species). The ules = 16 p. (14-18). female lip region has three to four annules. FEMALE (holotype): L = 0.52 mm; a = 26; The stylet knobs are round to slightly flattened b = 6.3; b' = 3.5; c = 15; c' = 2.7; H = 6 M; anteriorly. Incomplete aerolation of the lateral V = 22Q020; stylet = 18 /*; O = 16. Lip region field in the tail region is seen on some of the hemispherical, not set off, with three annules. females. The tail is tapering to a rounded Stylet knobs round. Rodlike sperms incom- (often almost pointed) terminus.

I.:*-: .-••^••••..O.'/ ^QvCKfrg-:.. lii6»Q5f>

^••:-D-

?|:.;9-"b:

0.02 mm

Figure 4. Radopholus magniglans n. sp.: A. Female, anterior region. B. Female, esophageal region. C. Female, longitudinal section at center of the body. D. Female, posterior region. E. Female, tail region.

Radopholus magniglans n. sp. with the lateral lips distinctly set off from the (Fig.4A-E) other lips. The lip region usually has two MEASUREMENTS: 2099: L = 0.53 mm annules, three annules are seen in a few (0.47-0.57); a = 27 (24-29); b = 5.5 (4.9- specimens. Stylet knobs are flattened ante- 6.1); b' = 2.6 (2,3-3.2); e = 16 (14-18); riorly or with slightly anterior projections. c' = 2.6 (2.4-2.9); H = 7 p (6-8); V = The intestine appears to extend into the tail is (is-20) 67 (64-71) I'Mii-i4>. stylet _ 22 /L region. Annulation around the tail terminus (21-23); O = 16 (11-23). is often irregular and indistinct. FEMALE (holotype): L = 0.56 mm; a = 27; Additional specimens of R. magniglans have b = 5.6; b' = 3.1; e = 15; c' = 2.8; H = 7 p, been identified from the following native habitats and localities in Australia: bank of the V = ^eS1-; stylet = 21 M; O = 19. Lip region hemispherical, slightly set off, with two dis- Onkaparinga River (above Mt. Bold Reser- tinct annules. Stylet knobs rounded, slightly voir), South Australia; Tyrenclarra, Victoria flattened surfaces. Two posterior esophageal and bank of the Murray River, about 10 miles glands coarser appearing than anterior esoph- above Renmark, South Australia. ageal gland; anterior gland with large nucleus, each posterior gland with one smaller nucleus. Radopholus trilineatus n. sp. Rodlike sperms filling spermatheca. Posterior (Fig. 5 A-F) ovary and spermatheca distinctly smaller than MEASUREMENTS: 20 9 paratypes: L = 0.59 anterior ovary. Lateral field with four inci- mm (0.53-0.66); a = 31 (28-34); b = 6.2 sures, inner lines closer together, coalescing (5.5-6.6); b' = 2.7 (2,5-3.1); c = 15 (13- to three incisures just anterior to level of anus. 18); c' = 2.7 (2.1-3.1); H = 6 ^ (5-7); Tail tapering, rounded terminus with an- V = 15 <14-18> 61 (59-64) " (13-ie). styiet = nulation. 20 ^ (18-21); O = 20 (16-24). MALE: Unknown. FEMALE (holotype): L = 0.60 mm; a = 30; JUVENILE : Unknown. b = 6.0; b' = 2.7; c = 15; c' = 2.7; H = 5 p; HOLOTYPE: Female, collected by S. D. Van v = ir.6ii«; stylet = 20 p.- O = 16. Lip region Gundy, 3 November 1965, catalog number 3, hemispherical, slightly set off, with two dis- U.C.R. Nematode Collection, Riverside, Cali- tinct annules. Stylet knobs round. Rocllike fornia. sperms filling spermatheca. Lateral field with PARATYPES: 16799, same collection as three incisures. Tail slightly tapering, rounded holotype distributed as follows: 799, De- terminus, annulated. partment of Nematology, University of Cali- MALE: Unknown. fornia, Davis; 1339 9, Department of Nema- JUVENILE : Unknown. tology, University of California, Riverside; 8 HOLOTYPE: Female, collected by S. D. Van 9 9, USDA Nematode Collection, Nematology Gundy, 19 May 1966, catalog number 4, Investigations, Beltsville, Maryland; 699, U.C.R. Nematode Collection, Riverside, Cali- Nematology Department, Rothamsted Experi- fornia. mental Station, Harpenden, England; 799, PARATYPES: 95 2 9, same data as holotype Plantenziektenkundige Dienst, Wageningen, distributed as follows: 799, Department of The Netherlands; and 699, Canadian Na- Nematology, University of California, Davis, tional Collection, Ottawa, Canada. 62 9 9, Department of Nematology, Univer- TYPE HABITAT AND LOCALITY: Unknown sity of California, Riverside; 6??, USDA grass and Eucalyptus sp. soil in native habitat, Nematode Collection, Nematology Investiga- bank of the Onkaparinga River (above Mylor), tions, Beltsville, Maryland; 799, Nematology South Australia. Department, Rothamsted Experimental Sta- DIAGNOSIS: R. magniglans can be distin- tion, Harpenden, England; 799 Plantenziek- guished from the preceding species by the tenkundige Dienst, Wageningen, The Nether- long conspicuous esophageal glands, usually lands; and 699, Canadian National Collec- two lip region annules and absence of male tion, Ottawa, Canada. specimens. TYPE HABITAT AND LOCALITY: EllCall/ptUS The face view is similar to the type species sp., unknown sedge, fern and grass soil in

A,G,M B,H,R A

Figure 5. Radopholus trilineatus n. sp.: A. Female. B. Female, face view. C. Female, anterior end. D. Female, lateral view near center of body. E-F. Female, tail regions. Radopholus rotundisemenus n. sp.: G. Female. H. Female, face view. I. Female, anterior end. J. Female, lateral view near center of body. K. Female, tail region. L. Juvenile, tail terminus. Radopholus vertexplanus n. sp.: M. Female. N. Juvenile, tail terminus. O. Female, tail region. P. Female, lateral view near center of body. O. Female, anterior end. R. Female, face view.

Copyright © 2011, The Helminthological Society of Washington OF WASHINGTON, VOLUME 35, NUMBER 2, JULY 1968 native habitat; bank of Scrubby Creek, near PARATYPES: 1822, 4 juveniles same data Narrarba, New South Wales, Australia. as holotype distributed as follows: 222, DIAGNOSIS: R. trilineatus can be distin- USDA Nematode Collection, Nematology Col- guished from the closely related R. magnigUms lection, Beltsville, Maryland; 322, Planten- by the three incisures of the lateral field in ziektenkundige Dienst, Wageningen, The the female, more anterior position of the vulva Netherlands; and 122 2, 4 juveniles, Depart- (59-63) and the usually shorter stylet length. ment of Nematology, University of California, It can be distinguished from R. loilliamsi by Riverside. the longer stylet, fewer annules on the lip TYPE HABITAT AND LOCALITY: Eucalyptus region and longer esophageal glands. sp., Acacia sp. and unknown native grass and In face view R. trilineatus is similar to R. fern soil in native habitat; Tyrendarro East, magniglans. The lip region usually has two Victoria, Australia. annules, some specimens without annulation. DIAGNOSIS: R. rotundisemenus can be dis- Stylet knobs are usually round, sometimes tinguished from the closely related R. inae- slightly flattened anteriorly. The intestine ap- qualis by the round sperms in the spermatheca, pears to extend into the tail region. The fewer incisures in the lateral field and usually annulation on the tail termination is irregular shorter stylet. and usually incomplete. The face view is similar to the type species. Additional specimens of R. trilineatus have Stylet knobs are flattened or usually with an- been identified from Eucalyptus sp. and un- terior projections (often pointed). Four to six, known grass soil in a native habitat west of usually irregular incisures are in the lateral Murray Bridge (Monarto South), South field near the center of the body. The intes- Australia. tine overlaps the rectum into the tail but is Radopholus rotundisemenus n. sp. indistinct on most specimens. The annulation (Fig. 5 G-L) around the tail terminus is distinct but sometimes irregular. MEASUREMENTS: 15 2 paratypes: L = 0.58 mm (0.54-0.61); a = 27 (24-33); b = Radopholus vertexplanus n. sp. 6.4 (5.6-7.2); b' = 3.9 (3.6-4.5); c = 14 (Fig. 5 M-R) (13-16); c' = 2.8 (2.4-3.6); H = 9 ^ (7- MEASUREMENTS: 15 2 paratypes: L = 11); V = - (1(i--T) 58 (56-61) -" d8-2e); stylet 0.51 mm (0.42-0.56); a = 30 (27-32); b = = 20^ (19-22); O = 16 (12-20). 5.8 (4.9-6.8); b' = 3.6 (3.2-4.0); c = 12 FEMALE (holotype): L = 0.58 mm; a = (11-15); c' = 3.8 (2.8-4.5); H = 9 p. (7- 32; b = 6.8; b' = 4.0; c = 15; c' = 3.3; H = 11); V = -4 (2a-->7> 58 (56-60) is d«-22). styiet 8 ^ V = ]!>611!>; stylet = 21 p.; O = 17. Lip = 16 fj, (15-17); O = 20 (16-25). region hemispherical, slightly set off, with FEMALE (holotype): L = 0.47 mm; a = 31; three indistinct annules. Stylet knobs with b = 5.4; b' = 3.4; c = 11; c' = 3.9; H = 10 ^ flattened anterior surfaces. Spermatheca with V = 255819; stylet = 16 M; O = 21. Lip region round sperms. Lateral field with four incisures flattened anteriorly with rounded edges, not in anterior and posterior part of body, five and set off from body, annulation indistinct. Stylet six incisures near center of body to three knobs with slightly flattened anterior surfaces. incisures on tail; incompletely aerolated on Spermatheca without sperms. Lateral field tail. Tail tapering slightly to rounded, annu- with four incisures narrowing to three incisures lated terminus. just anterior to level of anus. Tail tapering to MALE : Unknown. irregularly rounded annulated terminus. JUVENILE: Similar to female except for MALE: Unknown. visually fewer incisures in center of body and JUVENILE: Similar to female except for slightly shorter hyaline area in tail terminus shorter hyaline area in tail terminus (4-6 ^). (4-8,*). HOLOTYPE: Female collected by S. D. Van HOLOTYPE: Female collected by S. D. Van Gundy, 21 May 1966, catalog number 6, Gundy, 21 May 1966, catalog number 5, U.C.R. Nematode Collection, Riverside, Cali- U.C.R. Nematode Collection, Riverside, Cali- fornia. fornia. PARATYPES: 24 2 2, 11 juveniles, same data

6.m

0.02mm A,C-F,H,J-M O.O4 mm 0.01 mm , N Figure 6. Radopholus nativus n. sp.: A. Female, anterior region. B. Female, face view. C. Female, lateral view near center of body. D. Female, terminus. E. Juvenile, terminus. F. Female, tail region. G. Female, longitudinal section at center of body. Radopholus nigeriemis n. sp.: H. Female, anterior region. I. Female, face view. J. Female, lateral view near center of body. K-L. Female, tail termini. M. Female, tail region. N. Female, longitudinal section at center of body.

as holotype distributed as follows: 35? Clark, 29 March 1958, catalog number 7, USDA Nematode Collection, Nematology Sec- U.C.R. Nematode Collection, Riverside, Cali- tion, Beltsville, Maryland; 329 Plantenziek- fornia. tenkundige Dienst, Wageningen, The Nether- PARATYPES: 3099, 5 juveniles, same col- lands; 1892, 11 juveniles, Department of lection as holotype distributed as follows: Nematology, University of California, River- 399, 1 juvenile, USDA Nematode Collection, side. Nematology Investigations, Beltsville, Mary- TYPE HABITAT AND LOCALITY: EllC(lllJptilS land; 399, Plantenziektenkundige Dienst, sp., Acacia sp. and unknown native grass and Wageningen, The Netherlands; and 24 9 9, fern soil in native habitat; Tyrendarra East, 4 juveniles, Department of Nematology, Uni- Victoria, Australia. Same collection as R. versity of California, Riverside. rotundisemenus. TYPE HABITAT AND LOCALITY: Soil around DIAGNOSIS: R. vertexplanus can be dis- the native plants Carmichaelia monroi, Cel- tinguished from the most closely related spe- misia sp., Danthonia sp., Dracophylhim sp., cies R. williamsi by the longer tail with annu- and Senecio bellidioides in a native habitat, lation around the terminus, higher position of Porters Pass (3,100'), about 10 miles west of the vulva, the absence of sperms in the sper- Springfield, South Island, New Zealand. matheca and males and the flatter lip region. DIAGNOSIS: R. nativus can be distinguished Six similar lips without longitudinal stria- from the closely related species R. neosimilis tions are seen in face view (Fig. 5 R). The and R. vangundyi by the longer stylet and lip region is slightly or not set off with two absence of sperms in the spermatheca and or three faint often indistinct annules. Stylet males. It can be further distinguished from knobs are rounded to usually flattened an- R. vangundyi by the four lip region annules. teriorly. The intestine indistinctly overlaps the The face view is similar to the type species rectum. The spermatheca is often inconspic- (Fig. 6B). The lip region is slightly or not uous and without sperms. The tail tapers to set off, hemispherical or sometimes slightly a rounded or irregularly rounded, sometimes flattened anteriorly. Stylet knobs are flattened almost pointed terminus. anteriorly or with anterior projecting processes. Areolation of the lateral field varies from al- Radopholus nativus n. sp. most complete areolation to incomplete areola- (Fig.6A-F) tion only in the tail region. MEASUREMENTS: 15 9 paratypes: L = Additional specimens of R. nativus have 0.56 mm (0.50-0.69); a = 26 (23-30); b = been, identified from soil around native plants 5.8 (5.0-7,3); b' = 4.2 (3.3-5.0); c = 15 in a native habitat at Burkes Pass (1,650') (13-17); c' = 2.8 (2.1-3.3); H = 6 p. (4-8); about 10 miles east of Omarama, South Island, v = 23 (20-27) 60 (57-63) 22 do-*)), stylet = New Zealand. 20 yx (19-22); O = 16 (12-22). FEMALE (holotype): L = 0.55 mm; a = Radopholus nigerwnsis n. sp. 28; b = 5.2; b' = 3.5; c = 12; c' = 3.4; H = (Fig. 6 H-N) 7 p.; V = 2«582i; stylet = 21 ^ O = 17. Lip MEASUREMENTS: 20 2 paratypes: L = region hemispherical, slightly set off, with four 0.59 mm (0.49-0.71); a = 29 (25-34); b = annules. Stylet knobs with anterior projections. 6.6 (5.0-7,5); b' = 3.9 (3.4-4.4); c = 11 Spermatheca without sperms. Lateral field (10-14); c' = 3.7 (2.9-4.3); H = 2.5 ^ (2- with four incisures, inner incisures coalescing 4); V = 2

Lateral field with four and five indistinct At least nine additional undescribed species incisures. Intestine not overlapping rectum. of Radopholus have been identified from Phasmids in posterior portion of tail. Tail native habitats in Australia and New Zealand. tapering to slight ventral nipplelike terminus, Because of insufficient and/or poorly pre- without annulation. served specimens they are not described. MALE: Unknown. JUVENILE: Similar to female. Discussion HOLOTYPE: Female, collected by F. E. The genus Radopholus is especially inter- Caveness, 30 May 1960, catalog number 8, esting because of its distribution, sexual and U.C.R. Nematode Collection, University of usually juvenile dimorphism, rod-shaped sperms California, Riverside. in most species, and the apparent absence of PARATYPES: 50? 9, 94 juveniles, same data males in some species which have spermatized as holotype distributed as follows: 222,8 females. In addition, at least one species is juveniles, Department of Nematology, Uni- known to be widespread and of great economic versity of Califomia, Davis; 42 2 2, 62 juve- importance and contains physiological races. niles, Department of Nematology, University Nine of the eleven nominal species now in of California, Riverside; 222, 7 juveniles, the genus Radopholus are known from Aus- USDA Nematode Collection, Nematology In- tralia and New Zealand, and seven of these vestigations, Beltsville, Maryland; 222, 6 are from native habitats in Australia. Because juveniles, Nematology Department, Rothamsted of this, and the fact that six additional un- Experimental Station, Harpenden, England; described species are from native habitats in and 222, 11 juveniles, Plantenziektenkundige Australia, the genus Radopholus is considered Dienst, Wageningen, The Netherlands. to be indigenous to this area. The small num- TYPE HABITAT AND LOCALITY: Sandy loam ber of samples taken (75) in a limited area soil in native habitat around southern gamba of Australia (Victoria, Queensland and New grass (Andropogon tectorum) and spear grass South Wales), comprising 13 native species, is (Imperata cylindrical), 3 mile marker south of unique and quite remarkable for the Tylen- Igboora, Ibadan Province, western Nigeria. choidea. Four species (one described, three DIAGNOSIS: R. nigeriensis can be distin- undescribed) were identified in over 450 guished from the previous species by the short samples collected from native habitats in New hyaline area in tail terminus, phasmids in the Zealand. In contrast, over 1,000 soil samples posterior part of the body, usually shorter sty- from uncultivated areas in western and eastern let and the prominent vulval lips. Slightly Nigeria contained only one new species (R. protruding vulval lips have been seen in a few nigeriensis), Radopholus species indigenous specimens of R. similis from the roots. to other areas in the world are not known. The In face view six similar lips without longi- native habitats of R. williamsi and R. similis tudinal striations are seen (Fig. 6 I). The are not known but their distribution suggests lip region has two to three annules. Stylet they could have originated in Australia. knobs are flattened anteriorly to slightly pro- R. nigeriensis, from a native habitat in jecting anteriorly. Incisures of the lateral field Africa, shows the least affinity to the other vary from usually four or five to sometimes species. The absence of males in this species six near the center of the body. The phasmids is regrettable since their presence would fur- are often inconspicuous. The tail varies from ther indicate the relationship of this species almost pointed to irregularly rounded, some- to the rest of the genus. Such information times with a slight nipplelike terminus. would be most interesting for a species that Additional specimens have been seen from shows morphological differences from the other the following habitats and localities in north- species and is native to an area so far removed em Nigeria: soil around unknown native from the habitats of the other species. plants, ground nut (Arachis hypogaea) and Although males are known for only five of guinea corn (Sorghum margaritiferum), 2.5 the eleven nominal species of Radopholus, miles north of Katsina, Katsina Province; and they are all similar in having a reduced stylet, corn soil (Zee mays), 2 miles north of Katsina, esophagus, lateral lips and a higher, distinctly Katsina Province. set off lip region. Morphological differences

Copyright © 2011, The Helminthological Society of Washington OF WASHINGTON, VOLUME 35, NUMBER 2, JULY 1968 235 are seen between the juvenile specimens and species, R. trilineatus, might fit the specula- the adults in all species where juveniles are tions of Steiner and Buhrer, since the only known except in R. nigeriensis. These differ- stage found thus far in the soil is the sperma- ences, in addition to the normal smaller size tized female. of the juveniles, include the shorter hyaline More than 200 spermatized females but no area in the tail terminus and fewer lateral field males or juveniles of R. magniglans have been incisures in four of the species. found in four localities (during November and The rod-shaped sperms observed in most December 1965). Over 100 spermatized fe- Radopholus species are known only in the males but no males or juveniles of R. trilineatus closely related genus Radopholoides de Guiran, were found in two localities (during May 1967. The genus Schistonchus (Cobb, 1927), 1966). In addition, the ovaries in these speci- which is in the Aphelenchoidea, contains sim- mens appeared uniform in size and develop- ilar but longer flagellate sperms (Thorne, ment. This suggests that these species may 1961). These rod-shaped sperms were first have a more formal or restrictive life cycle, illustrated for R. similis in Cobb's 1915 paper, and that the other stages might be found in which gives a detailed description of this the soil during different times of the year species and contains excellent illustrations. and/or the other stages might be in the roots The apparent absence of males in R. mag- of plants. The rather short ovaries and ab- niglans, R. trilineatus, and R. rotundisemenus, sence of well-developed oocytes or eggs might and presence of spermatized females, cannot be further evidence that these species undergo be satisfactorily explained. In closely related further development in the roots of the plants. genera, Pratylenchus Filipjev, 1934 and R. rotundisemenus also has spermatized fe- Hirschmanniella Luc and Goodey, 1963, as males and no males but only 18 females are reported by Loof (1960) and Sher (1968) known from one locality. In addition, juveniles and in a closely related subfamily (Sher, of R. rotundisemenus, two additional species 1965, Sher, 1966), there is a good correla- of Radopholus and a few male Radopholus, tion between the presence of males and that could not be identified as to species, were sperms in the spermatheca, and the absence found in this collection. The ovaries in R. of males and no sperms in the spermatheca. rotundisemenus are variable, well-developed, Genera, in the more distantly related family and the spermatheca contains round sperms. Criconematidae, have been recorded with Round sperms are also seen in one of the spermatized spermathecas and absence of species of Radopholus that is not described males, as well as no sperms in the spermatheca because of insufficient specimens. and presence of males (Raski and Golden, The best known, most widely distributed 1965). and only known pathogen in this genus, Ra- Brooks and Perry (1962) report the appar- dopholus similis, has been recorded as para- ent reproduction of R. similis for three genera- sitizing 244 different plants (Poucher, et al., tions without males. It was not noted if 1967). Bally and Reydon (1931) first re- sperms were present in the spermatheca in ported the possibility of biological strains in these specimens. this species when they were unable to infect Steiner and Buhrer (1933) in a discussion Gigantochloa apus with specimens from Coffca of the sexual dimorphism of T. similis Cobb, robusta. Other workers have reported the 1893, speculated that, "Unless disproved, then, inability of Radopholus similis from Musa spp. the sexual dimorphism may be considered an to infect Citrus spp. (DuGharme and Birch- adaptation to the life-long endoparasitism of field, 1956; Blake, 1961; van Weerdt, 1957). T. similis, to its possible habit of copulation A study of specimens (races) from banana and within the host tissue, to the possible cessa- citrus has been made and no consistent mor- tion of nutrition by the adult male and its phological differences can be seen between death shortly after copulation." This is appar- specimens from banana or citrus roots; this ently not true for Radopholus similis, as nu- confirms the observations of DuCharme and merous juveniles, females, and males are found Birchfield (1956) and van Weerdt (1958). in the soil around roots parasitized by this Specimens from Java have been unavailable species. R. magniglans, and the closely related for study.

Morphological variation in single female Thome. Soil and Crop Sci. Soc. Fla. Proc. 22: progeny of Radopholus similis was reported by 160-162. van Weerdt in 1958. Similar polymorphism Christie, J. R. 1957. The yellows disease of was seen within the many populations of R. pepper (Piper) and spreading decline. PL similis examined during the present study. The Dis. Rept. 41: 267-268. Cobb, N. A. 1893. Nematodes, mostly Austra- most common variations are in the female lip lian and Fijian. Macleary Mem. Vol. Linn. region shape and stylet knob shape (Fig. 1 Soc. N. South Wales, p. 252-308. A—B), as well as areolation of the posterior . 1909. Nematodes (In Fungus maladies portion of the lateral field of the female (Fig. of the sugar cane). Bull. ( 6) Hawaiian Sugar 1 K-M). Van Weerdt (1958) considered the Planters' Assn. Exper. Station, Div. Path, and areolation of the lateral field "an artifact as Physiology 51-73. a result of the fixation in formalin . . ." 1915. Tylenchus similis, the cause of a root disease of sugar cane and banana. J. Summary Agric. Research, U. S. Dept. Agric. 4(6): 561-568. The genus Radopholus Thorne, 1949 is diag- DuCharme, E. P., and W. Birchtield. 1956. nosed. The four nominal species [R. similis Physiological races of the burrowing nema- (Cobb, 1893), R. inaequalis Sauer, 1958, R. tode. Phytopath. 46: 615-616. neosimilis Sauer, 1958 and R. williamsi Siddiqi, Filipjev, I. N. 1934. Harmful and useful nema- 1964] are redescribed from type specimens. todes in rural economy (Russian Text). 440 Seven new species are proposed; five from pp., figs. 1-333, Moskva, Leningrad. Australia, one from New Zealand and one from . 1936a. Ueber freilebende und pflan- Nigeria; all from native habitats in these areas. zenparasitische Gattungen der Tylenchinen Trudy Zool. Inst. Akad. Nauk SSSR 3: 537- A key to the species and illustrations of the 550. species are presented. The distribution and . 1936b. On the classification of the some morphological characters of Radopholus Tylenchinae. Proc. Helm. Soc. Wash. 3(2): are discussed. 80-82. , and J. H. Schuurmans Stekhoven, Jr. Acknowledgments 1941. A manual of agricultural helminthol- Helpful assistance in the preparation of ogy. 878 pp. Leiden. nematode slides, measurements, and illustra- Goodey, T. 1932. The genus Anguittulina Gerv. & v. Ben., 1859, vel Tylenchus Bastian, 1865. tions was received from: A. H. Bell, K. W. J. Helminth. 10(2-3): 75-180. Brown, and L. Wang. de Guiran, G. 1967. Description de Radophol- oides litoralis n. g., n. sp. (Nematoda: Praty- Literature Cited lenchinae). Nematologica 13: 231-234. Allen, M. W. 1955. A review of the nematode Loot, P. A. A. 1960. Taxonomic studies on genus Tylenchorhynchus. Univ. Calif. Publ. the genus Pratylenchus ( Nematoda). Tijdschr. Zool. 61, 129-165. Plantenziekten. 66: 29-90. , and S. A. Sher. 1967. Taxonomic prob- Luc, M., and J. B. Goodey. 1962. Hirsch- lems concerning the phytoparasitic nematodes. mannia n. g. differentiated from Radopholus Ann. Rev. Phytopathology 5: 247-264. Thorne, 1949 (Nematoda: Tylenchoidea). Baker, A. D. 1953. Rapid method for mounting Nematologica 7: 197-202. nematodes in glycerine. Canad. Entom. 85: Menzel, R. 1929. De Plagen van de Thee in 77-78. Neclerlandsh—Indie (Java en Sumatra) en Bally, W., and G. A. Reydon. 1931. De tegen- Hare Bestrijding. Archief voor de Theecultur woordige Stand van het Vraagstuk van de wor- 1: 11-66. telaaltjes in die Koffiecultuur. Arch. Koffie- Micoletzky, H. 1922, Die freilebenden Erd- cult. Nederl. Indie 5: 23-216. Nematoden. Arch. Naturgesch. 87(A) (1921), Blake, C. D. 1961. Root rot of bananas caused 1-650. by Radopholus similis (Cobb) and its control Poucher, C., H. W. Ford, R. F. Suit, and E. P. in New South Wales. Nematologica 6: 295- DuCharme. 1967. Burrowing nematode in 310. citrus. Bull. 7. Florida Dept. Agric., Gaines- Brooks, T. L., and V. G. Perry. 1962. Appar- ville, Florida. ent parthenogenic reproductions of the bur- Raski, D. J., and A. M. Golden. 1965. Studies rowing nematode Radopholus similis (Cobb) on the genus Criconemoides Taylor, 1936 with

descriptions of eleven new species and Ba- Thorne, G. 1949. On the classification of the kernema variable n. sp. (Criconematidae: Tylenchida, new order (Nematoda, Phas- Nematoda). Nematologica 11: 501-565. midia). Proc. Helm. Soc. Wash. 16(2): 37- Sauer, M. R. 1958. Ploplolaimus gracilidens, 73. Radopholus inaequalis, and Radopholus neo- . 1961. Principles of Nematology. New similis, three new tylenchs native to Victoria, York, McGraw-Hill, 553 pp. Australia. Nematologica 3(2): 97-107. van der Vecht, J. 1950. Op planten parasi- Siddiqi, M. R. 1964. Radopholus williamsi n. terende aaltjes (Tylenchidae). pp. 16-42. In sp. (Nematoda: Pratylenchiclae), a parasite Plagen van de cultuurgewassen in Indonesie. of sugar cane roots at L'Etoile, Mauritius. In- W. van Hoeve, The Hague. dian J. Ent. 26(2): 207-208. van Weerdt, L. G. 1957. Studies on the biol- Slier, S. A. 1965. Revision of the Hoplolaiminae ogy of Radopholus similis (Cobb, 1893) (Nematoda) V. Rotylenchus Filipjev, 1936. Thorne, 1949. I. PL Dis. Rept. 41(10): 832- Nematologica 11: 173-198. 835. . 1966. Revision of the Hoplolaiminae . 1958. Studies on the biology of Ra- (Nematoda) VI. Helicotylenchus Steiner, 1945. dopholus similis (Cobb, 1893) Thorne, 1949. Nematologica 12: 1-56. II. Morphological variation within and be- . 1968. Revision of the genus Hirschman- tween progenies of single females. Nemato- niella Luc & Goodey, 1963 (Nematoda: Ty- logica 3: 184-196. lenchoidea). Nematologica (in press). Williams, J. R. 1960. Studies on the nematode Steiner, G., and E. M. Buhrer. 1933. The fauna of sugar cane fields in Mauritius. 4. nematode Tylenchus similis Cobb, as a para- Tylenchidae (partim). Occ. Pap. Maurit. Sug. site of the tea plant (Thea sinensis, Linn.), its Ind. Res. Inst. 4: 1-30. sexual dimorphism, and its nemic associates in Zimmermaiin, A. W. P. 1898. De nematoden the same host. Zeitsch. Parasitenk. 5(2): der koffiewortels. Deel I. Mededeel. 's Lands 412-420. Plantentuin (Buitenzorg) 27: 1-64.

Digenetic Trematodes of Marine Fishes from Ghana: Families Acanthocolpidae, Bucephalidae, Didymozoidae1

JACOB H. FISCHTHAL AND J. D. THOMAS2

This paper is the second in a series by the DATE: 22 December 1964. authors on the digenetic trematodes of marine SPECIMEN: USNM Helm. Coll. No. 63330. fishes from Ghana. Specimens have been DISCUSSION: Our collection consists of two deposited in the U. S. National Museum Hel- specimens from one host. Seven spines in the minthological Collection as indicated. All oral row of circumoral spines measure 72—93 measurements are in microns, unless otherwise by 17-22, while nine in the aboral row are noted. 68-82 by 19-26. The body spines end just short of the posterior extremity. Fifteen eggs Family Acanthocolpidae measure 66-74 by 42-47; they contain 8-16 Stephanostomum casum (Linton, 1910) cell embryos measuring 38-52 by 29-33. Our McFarlane, 1934 specimens compare favorably with specimens HOST: Lutjanits modestus Blecker, red snap- of S. casum in the U. S. National Museum per (Lutjanidae). Helminthological Collection from Florida (no. HABITAT: Rectum. 8743), Massachusetts (no. 8211), North Caro- LOCALITY: Tema, Ghana. lina (no. 37101), and Puerto Rico (no. 39341).

1 Contribution from the Department of Biology, State Stephanostomum ghanensis n. sp. University of New York at Binghamton, Binghamton, New York 13901 (J. H. Fischthal). (Figs. 1, 2) - Address of J. D. Thomas: School of Biological Sci- ences, The University of Sussex, Falmer, Brighton, Sussex, England. HOST: Trachinotiis goreensis Cuvier and This study was supported in part by a State University Valenciennes, pampano (Carangidae). of New York Faculty Research Fellowship (FRF67-40-004) awarded to the senior author. HABITAT: Stomach.

LOCALITY: Iture, Ghana. cular, somewhat winding anteriorly. Pars pro- DATE: 21 February 1966. statica short, just posterodorsal to acetabulum, SPECIMEN: USNM Helm. Coll. No. 63331 surrounded by few prostate cells. Cirrus long, (holotype). winding, proximal part spined. Cirrus sac DIAGNOSIS (based on one adult and two uniting with metraterm at anterior margin of immature specimens; adult measured): Body acetabulum, forming short genital atrium. 4,498 by 630 at testicular level. Tegument Genital pore median, just preacetabular. annulated preacetabularly, spined to level of Ovary 218 by 232, smooth, lying 700 post- posterior testis. Eyespot pigment present. acetabular. Ootype complex dorsal to ovary. Forebody 1,365 long, hindbody 2,858 long. Uterus extensively coiled between acetabulum Oral sucker 172 by 230, truncate posteriorly. and ovary, slightly overlapping latter dorsally, Circumoral spines 34-36 in number, in two sperm in proximal portion. Metraterm thick- alternating, uninterrupted rows; ventral oral walled, shorter than cirrus sac, spines not spines 45-57 by 22-23, dorsal 61-63 by 19- observed. Vitellaria extensive, commencing 24; ventral aboral spines 33-41 by 16-19, 340 postacetabular; follicles ventral, lateral and dorsal 65-75 by 21-23. Acetabulum 275 by dorsal to ceca, invading intercecal space dorsal 255, center at level of anterior one-third of to gonads but not confluent, filling posttestic- body length. Sucker length ratio 1:1.60. Pre- ular space except medianly, confluent dorsal pharynx 655 long, thick-walled, muscular; to proximal half of uterus between anteriormost pharynx 300 by 245, pyriform, glands antero- margin of vitellaria to just preovarian. Eggs laterally and posteriorly; esophagus 152 long, yellow, thin-shelled, operculate, eight measur- thick-walled, muscular, glands along length; ing 62-66 by 37—45, zygote undivided. cecal bifurcation just preacetabular; ceca con- Excretory bladder long, narrow, extending spicuously cell-lined, extending to posterior to ovarian level, ducts reaching posterolateral extremity, opening into excretory bladder. margins of oral sucker before turning back on Gonads tandem, contiguous, filling inter- themselves and running posteriorly at least to cecal space, may overlap ceca ventrally. Testes vitellaria (probably beyond), pore terminal. two, smooth; anterior testis 440 by 330, dorsally DISCUSSION: Our species could not be keyed overlapping ovary 51; posterior testis 605 by to any species listed in the keys given by 295; posttesticular space 955 long, also long Manter and Van Cleave (1951) and Caballero in immature specimens. Cirrus sac winding, (1952). The closest species appear to be S. 700 (longitudinal extent) by 152 at seminal sentum (Linton, 1910) Manter, 1947, and S. vesicle, commencing 425 postacetabular (three- anisotremi Manter, 1940, but it differs from fifths of distance from latter to ovary), just them in having a much longer posttesticular contacting vitellaria medianly. Internal seminal space, in the cirrus sac extending more than vesicle 375 (longitudinal extent) by 145, sac- halfway to the ovary, and in the vitelline

Figures 1-11. 1. Stephanostomum ghanensis, holotype, ventral view. 2. Same. Terminal genitalia, holotype, ventral view. 3. Stephanostomum trachinoti, holotype, ventral view. 4. Bucephaloides ghanensis, holotype, anterior part of body in dextrolateral view, from pharynx posteriorly in dorsal view. 5a. Prosorhynchus caudovatus, relative position of gonads, ventral view; b. dorsal view of second specimen; c. dextrolateral view of third specimen. 6. Rhipidocotyle ghanensis, holotype, ventral view. 7. Immature Didymozoid D, ventral view. 8. Immature Didymozoid E, from Scyris alexandrinus, ventral view. 9. Immature Didymozoid F, dextrolateral view. 10. Immature Didymozoid G, ventral view. Ha. Allonematobothrium ghanensis, paratype, anterior extremity, dorsal view; b. terminal genitalia, holotype, dextrolateral view; c. region of anterior part of testes, holotype; d. region of genital junction (intestinal ceca omitted), holotype; e. posterior extremity, paratype. Abbreviations: A, acetabulum; C, cirrus; CS, cirrus sac; E, esophagus; EA, excretory arm; EB, excretory bladder; GA, genital atrium; GC, gland cells; GJ, genital junction chamber; GP, genital pore; I, intestinal cecum; M, metraterm; MG, Mehlis' gland; O, ovary; OS, oral sucker; P, pharynx; PC, prostate cells; PP, pars prostatica; S, spine; SV, seminal vesicle; T, testis; U, uterus; UA, ascending uterus; UD, descending uterus; V, vitellarium; VD, vas deferens; VE, vas efferens; VR, vitelline reservoir; VW, ventral body wall.

Copyright © 2011, The Helminthological Society of Washington 240 PROCEEDINGS OF THE HELMINTHOLOGICAL SOCIETY follicles being confluent dorsal to the proximal in two alternating, uninterrupted rows, 5-7 by half of the uterus; it differs further from S. 4-5, oral spines smaller than aboral. Acetab- anisotremi in having fewer circumoral spines. ulum 94 by 102, center at level of anterior two-fifths of body length. Sucker length ratio Stephanostomum megacephahim 1:0.75. Prepharynx 118 long, thick-walled, Manter, 1940 muscular; pharynx 60 by 54, oval, four-lobed anteriorly; esophagus 184 long, lined with cells HOSTS: Caranx hippos (L.), jack or horse continuous with those of ceca, longer than mackerel (Carangidae); Myxus curvidens prepharynx or pharynx; cecal bifurcation 12 (Valenciennes), mullet (Mugilidae). preacetabular; ceca long, cell-lined, terminat- HABITAT: Small intestine. ing 46 from posterior extremity, opening into LOCALITIES: Cape Coast (Caranx), Tema excretory bladder. (Myxus); Ghana. Gonads tandem, 4-5 apart, filling intercecal DATE: 27 April 1966 (Caranx). space, may overlap ceca ventrally. Testes two, SPECIMENS: USNM Helm. Coll. No. 63332 smooth; anterior testis 118 by 93, posterior tes- (from Caranx); No. 63333 (Myxus). tis 115 by 103; posttesticular space 203 long. DISCUSSION: Our specimens readily keyed Cirrus sac 198 by 46, median, commencing 82 to S. megacephahim in the keys given by postacetabular, 48 previtellarian, 168 preovar- Manter and Van Cleave (1951) and Caballero ian; containing small seminal vesicle, short (1952). This species has been reported from pars prostatica surrounded by prostate cells, Caranx hippos from the Pacific coast of Mexico, and long, spined cirrus. Cirrus sac uniting Panama and Ecuador, and from the Gulf of with metraterm preacetabularly, forming short Mexico (Florida); it has also been found in genital atrium. Genital pore median, just pre- C. latus Agassiz from Tortugas (Florida) and acetabular. Jamaica (West Indies). Our collection con- Ovary 45 by 38, smooth, lying 250 post- sists of four adults and one immature specimen, acetabular. Ootype complex anterodextral to with 28—30 circumoral spines, from one C. ovary. Uterus with little coiling, ascending hippos, and four adults, with 31-32 spines, ventral to proximal part of cirrus sac. Metra- from one M. curvidens. The circumoral spines term 140 by 20, thick-walled, muscular, ap- in specimens from the latter host average pearing spined, shorter than and dextral to smaller (46-75 by 7-11) than, those originally cirrus sac, commencing 31 postacetabular. described; we believe this to be host influ- Vitellaria commencing 130 postacetabular, 120 enced. preovarian; follicles very small, ventral, lateral and dorsal to ceca, invading interceal space Stephanostomum trachinoti n. sp. slightly, filling posttesticular space. (Fig. 3) Excretory bladder long, narrow, extending dorsally to midlength of anterior testis, pore HOST: Trachinotus glanciis (L.), palometa terminal. (Carangidae). DISCUSSION: This form could not be keyed HABITAT: Small intestine. to any species given in the keys by Manter LOCALITY: Cape Coast, Ghana. and Van Cleave (1951) and Caballero (1952). DATE: 20 April 1966. The unique combination of characteristics SPECIMEN: USNM Helm. Coll. No. 63334 prompted us to describe it as a new species (holotype). even, though immature. It differs from all DIAGNOSIS (based on single immature specimen): Body 1,344 by 210 (preacetabular), species, except S. multispinosum Manter, 1940, extremities round. Tegument spinecl to ante- in having 80-84 circumoral spines. The latter rior testis level, extending anteriorly up to species differs in the oral sucker being cupuli- circumoral spines. Eyespot pigment abundant, form and smaller than the acetabulum, in scattered from oral sucker to micllength of having a pyriform pharynx and a short esoph- esophagus. Forebody 505 long, hinclbody 745 agus, in the vitellaria extending to the ace- long. Oral sucker 125 by 94, elongate, funnel- tabulum, and in the cirrus sac extending a shaped. Circumoral spines numbering 80—84, considerable distance postacetabular.

Family Bucephalidae genital atrium. Eggs few, thick-shelled, yellow- Bucephaloides ghanensis n. sp. brown, operculate, 10 measuring 16-21 by (Fig. 4) 10-15; many abnormal eggs present. HOST: Scomheromorus tritor (Cuvier and Excretory bladder very long, narrow, com- Valenciennes), Spanish mackerel or kingfish mencing 232 from anterior sucker, 320 from (Scombridae). anterior extremity; pore terminal. HABITAT: Small intestine. DISCUSSION: Our species most closely re- LOCALITY: Iture, Ghana. sembles Bucephaloides philippinorum Velas- DATE: 21 February 1966. quez, 1959, from a sphyraenid fish from Luzon SPECIMEN: USNM Helm. Coll. No. 63335 Island, Philippines, and B. tennis (Yamaguti, (holotype). 1952) Hopkins, 1954, from a platycephalicl DIAGNOSIS (based on one complete speci- fish from Celebes. Both these species differ men, and one with previtellarian body miss- from ours in having a much longer posttestic- ing) : Body 2,277 by 175 at ovarian level, ular space and cirrus sac, and in the uterus elongate, narrow, extremities round. Tegu- extending considerably previtellarian and pre- ment entirely spined. Anterior sucker 85 long, pharyngeal. B. tennis differs further in having 77 deep, subterminal ventral. Mouth ventral a much shorter intestine, a larger ovary which to pharynx, at midbody length; pharynx 92 is about the same size as the testes, and the by 86, lying 1,110 from anterior extremity; anterior tip of the excretory bladder being esophagus muscular, very short, directed dor- much nearer the pharynx. sally; intestine 560 by 46, very long, cell-lined, directed posteriorly to level of ovary or anterior Prosorhynchus caudovatus Manter, 1940 testis. (Figs. 5a-c) Gonads smooth, tandem, may be contiguous SYNONYM: Prosorhynchus crucibulus of Eck- or slightly separated from one another, extend- mann, 1932, nee Rudolphi, 1819. ing posteriorly from posterior part of middle HOSTS: Epinephelus goreensis (Cuvier and third of body. Testes two, median; anterior Valenciennes), sea perch or grouper (Ser- testis 123-145 by 102-114, posterior testis 116 ranidae); Lutjanus maltzani (Steindachner), by 94-97; posttesticular space 219-320 long. snapper (Lutjaniclae). Cirrus sac 257-340 by 52-58, thick-walled, HABITATS: Small intestine, stomach. muscular, straight to slightly sinuous, terminat- LOCALITY: Tema, Ghana. ing 30—38 from posterior extremity. Seminal DATE: 18 December 1964. vesicle 88-102 by 40-48, elongate oval. Pars SPECIMENS: USNM Helm. Coll. No. 63336 prostatica 155-195 by 23-31, cell-lined, sur- (from Epinephelus); No. 63337 (Lutjanus), rounded by dense mass of prostate cells. MEASUREMENTS AND SOME PERTINENT DATA Genital lobes unequal, projecting into genital (based on two specimens from E. goreensis, atrium. Latter 65-80 by 47-71; short duct and four from L. maltzani; five measured): leading to subterminal ventral genital pore. Body 1,715-2,245 by 465-675, entirely spined; Ovary 72-73 by 68-72, round, submedian rhynchus 285-330 by 205-245, wedge- to dextral, lying 440 postpharyngeal and 1,622 funnel-shaped; pharynx 77-100 by 95-120, from anterior extremity (holotype). Oviduct wider than long; narrow esophagus may be muscular, from posterodextral margin of ovary, present; intestine 222-435 by 125-170; testes extending posteriorly dorsal to anterior testis. 167-232 by 150-196, their position in relation Vitelline follicles in narrow lateral fields, ex- to each other and to ovary showing considera- tending from postpharyngeally (53 in holo- ble variation as illustrated in Figs. 5a-c; cirrus type) to level of ovary or anterior testis, sac 655-760 by 143-170, thick-walled, mus- follicles (right-left) numbering 16-16, 16-17; cular, may overlap posterior testis or may be vitelline duct from each field extending poste- entirely posttesticular; genital atrium 186-300 riorly dorsal to ovary and anterior testis, unit- by 167-200; ovary 124-198 by 111-155, over- ing at posterior part of latter to form short lapping testis dorsally; ootype complex dorsal common duct. Uterus with few coils, extend- to testis; Laurer's canal extending posteriorly ing from ovarian level to posterior margin of beyond testis to dorsal surface of body; sperm

Copyright © 2011, The Helminthological Society of Washington 242 PROCEEDINGS OF THE HELMINTHOLOGICAL SOCIETY in proximal part of uterus, uterine coils may 58-77, round to longitudinally elongate, at extend previtellarian; vitelline follicles in five level of anterior testis; esophagus long, 97-152 specimens numbering (right-left) 8-12, 11-13, by 42-58, muscular, passing anteriorly from 11-14, 12-14, 14-14, usually in separate pharynx to ovarian level; intestine 196—272 by lateral fields, forming inverted U in two; 126-150, oval, conspicuously cell-lined, dor- vitelline reservoir small; eggs thick-shelled, somedian, extending very slightly anterodorsal operculate, yellow to yellow-brown, 20 measur- before looping posteriorly, terminating anterior ing 32-43 by 21—25, usually with anopercular to or at pharyngeal level, may overlap median filament which may be up to 30 long, some parts of ovary and anterior testis. with anopercular knob only; some abnormally Gonads smooth, tandem, usually overlapping shaped eggs present in all specimens. adjacent one, dextral, posterior to midbody DISCUSSION: This species was originally de- length. Anterior testis 220-278 by 157-198, scribed from Epinephelus sp. from the Suez. posterior testis 222-240 by 150-193; post- The extreme variation in the relative positions testicular space 405—585 long. Cirrus sac 518- of the gonads raises some question regarding 720 by 78-95, thick-walled, muscular, C- the assignment of some species to the genus shaped to straight, extending from level of Neidhartia Nagaty, 1937, as they are based posterior testis to 100-240 from posterior on only one or a very few specimens. Had we extremity. Seminal vesicle 114-170 by 60-78, encountered our specimen illustrated in Figure elongate oval. Pars prostatica 276-392 by 5b by itself, we probably would have described 45-53, cell-lined, surrounded by dense mass it as a new species of Neidhartia rather than of prostate cells. Genital lobes muscular, un- recognizing it as a variant of Prosorhynchm equal, projecting into genital atrium. Latter caiidovatus. An anopercular process has been 114-140 by 100-125; narrow, very thick- described for at least some of the eggs in walled, muscular duct leading to subterminal several species of bucephalids: Paiirorhynchus ventral genital pore. hiodontis Dickerman, 1954, possesses a "knob"; Ovary 138-182 by 116-129, pretesticular, Bucephalus kathetofitomae (Manter, 1934) lying 715-1,380 from anterior extremity. Ovi- Manter, 1940, an "irregularly shaped cap"; duct muscular, from posterior or dextrolateral Bucephalus fragilis Velasquez, 1959, a "pro- margin of ovary, extending posteriorly dorsal tuberance." to anterior testis. Vitelline follicles in narrow lateral fields, extending from 505-840 from Rhipidocotyle ghanensis n. sp. anterior extremity to ovarian level; follicles (Fig. 6) numbering (right-left) 8-14, 13-16, 13-17, HOST: Psettodes belcheri Bennett (Psetto- 15—18 in four specimens, respectively; vitelline didae). duct from each field extending long distance HABITAT: Small intestine. posteriorly, dorsal to ovary and anterior testis, LOCALITY: Tema, Ghana. uniting to form short common duct at overlap SPECIMENS: USNM Helm. Coll. No. 63338 of anterior and posterior testes. Uterus exten- (holotype and two paratypes on same slide). sive, extending from 495-640 from anterior DIAGNOSIS (based on five specimens; four extremity (previtellarian) to level of posterior measured): Body 1,770-2,510 by 290-450 at margin of genital atrium or more posteriorly. postvitellarian or testicular level, dorsoventrally Eggs numerous, yellow-brown, thick-shelled, flattened anterior to midlength of vitellaria, operculate, 20 older ones measuring 23—27 by much rounded posteriorly, anterior extremity 16—18; younger eggs nearest ovary thin-shelled, truncate, posterior round. Tegument entirely larger and rounder. spined, anterior spines more scalelike. Anterior Excretory bladder very long, narrow poste- sucker 190-205 by 143-175, subterminal riorly, considerably expanded anteriorly and ventral, opening round to elongate oval, with filling most of body width, extending almost seven-lobed polygonal hood measuring 82-107 to anterior sucker to slightly overlapping latter by 220-242, lobes may show muscular papilla dorsally; pore terminal. when extended. Mouth 1,060-1,670 from DISCUSSION: Our species appears closest to anterior extremity, well posterior to midlength Rhipidocotyle longleyi Manter, 1934, from ser- of body, posterior to pharynx; latter 61-76 by ranid and lutjanid fishes from Florida and

Japan, and to R. laruei Velasquez, 1959, from Didymozoidae (Monilicaecum) a psettodid fish from Luzon Island, Philippines. larvae I Nikolaeva, 1965 Both these species differ from ours in having HOSTS: Vomer setapinnis (Mitchill), moon- the mouth near midbody length, lacking an fish (Carangidae); Brachydeiiterus auritus esophagus or having only a very short one, and (Cuvier and Valenciennes), burrito (Pomad- the excretory bladder extending anteriorly only asyidae); Scomberomorus tritor (Cuvier and to the level of the digestive tract. R. longleyi Valenciennes), Spanish mackerel or kingfish differs further in the intestine extending both (Scombridae). anterior and posterior to the mouth, and in HABITAT: Small intestine. possessing a papilla on the posterior margin of LOCALITY: Cape Coast, Ghana. the genital pore. R. laruei differs further in DATES: 29 November, 6 December 1965; the intestine extending anteriorly from the 4 March 1966. pharynx, having the gonads at midbody length, SPECIMENS: USNM Helm. Coll. No. 63341 the cirrus sac comprising almost one-half of (from Vomer); No. 63342 (Brachydeiiterus); the body length, the seminal vesicle being No. 63343 (Scomberomorus). coiled, and the uterus extending anteriorly DESCRIPTION (based on three specimens; all only to the level of the anterior vitelline measured): Body 1,748-2,030 by 213-280, follicles. elongate, anterior part narrow, gradually wid- Family Didymozoidae ening to rounded posterior extremity; body 7.2-8.8 times longer than wide. Forebocly Fischthal and Kuntz (1964) and Nikolaeva 395-452 long, hindbody 1,280-1,490 long. (1964, 1965) have reviewed previously de- Body filled with vesicular parenchyma. Paren- scribed immature didymozoids in addition to chymal glands conspicuous, filling entire body describing some new ones. Subsequently, in zone beneath tegument. Oral sucker 156— Fischthal and Kuntz (1965) reported Torti- 167 by 61-66, 59-73 of it protruding from caecum nipponicum Yamaguti, 1942, from a anterior body opening, elongate; composed of colubrid snake from North Borneo, and Paruk- outer thin layer of longitudinal and inner hin (1966) illustrated, without describing, thicker layer of circular muscles, and much "Didymozoidae gen. sp. larvae" from carangid greater inner area of vesicular parenchymatous fishes from the South China Sea; the latter cells, combined thickness of muscle layers .5—8. form is Torticaecum-\ike. Acetabulum 73-99 by 80-93, muscular, center Immature Didymozoid B Fischthal at level of anterior one-fourth of body length. and Kuntz, 1964 Sucker length ratio 1:0.44-0.63. Distance between suckers 266-342, its ratio to body length HOSTS: Euthynnus alleteratus (Rafinesque), being 1:5.9-6.6. Pharynx 22-27 by 23-27, false albacore or little tunny (Scombridae); round, inner part non-muscular and composed Brachydeiiterus auritus (Cuvier and Valen- of vesicular cells, contiguous with center of ciennes ), burrito (Pomadasyidae). posterior part of oral sucker; esophagus 186- HABITAT: Small intestine. 276 long, sinuous, thick-wallecl, muscular, LOCALITIES: Tema (E. alleteratus), Cape lumen much enlarged at distal end just before Coast (B. auritus); Ghana. cecal bifurcation; latter 38-72 preacetabular; DATES: 23 October, 17 December 1964; one cecum ascending very short distance before 6 December 1965. descent, ceca lined internally to acetabular SPECIMENS: USNM Helm. Coll. No. 63339 level with very thick layer of amber-colored, (from Euthynnus); No. 63340 (Brachydeii- large glands with free ends projecting into terus) . small lumen, length of glandular part of right DISCUSSION: Two albacore harbored four cecum 120-127, left cecum 114-140, no nuclei and 12 specimens, respectively; one worm was visible in glands, walls 6-23 thick; indistinct, recovered from the burrito. This immature somewhat diffuse, large, very granular, multi- didymozoid was originally described from nucleate glands external to thick part of ceca; Euthynnus yaito Kishinouye from Palawan 7-8 narrow, elongate chambers along post- Island, Philippines. acetabular length of each cecum, most cham-

Copyright © 2011, The Helminthological Society of Washington 244 PROCEEDINGS OF THE HELMINTHOLOGICAL SOCIETY bers with amber-colored amorphous contents; HABITAT: Small intestine. ceca terminating subequally, one 114-191 from LOCALITY: Cape Coast, Ghana. posterior extremity, other 275-360. No re- DATE: 4 March 1966. productive fundaments. Excretory bladder SPECIMEN: USNM Helm. Coll. No. 63344. elongate, narrow, followed anteriorly as far as DESCRIPTION (based on single specimen): acetabulum, but probably extends preacetab- Characteristics as for Didymozoidae (Moni- ular; pore terminal. licaecum) larvae I, except as noted below. DISCUSSION: This form was originally de- Body 2,085 by 240, abruptly narrowing post- scribed from the small intestine of Sardinops cecally to papillalike termination; body 8.7 ocellata (Clupeidae) and Zeus faber L. (Zei- times longer than wide. Forebocly 750 long, dae) from the Atlantic coast of Southwest hindbody 1,245 long. Oral sucker 119 by 50, Africa. We are referring our specimens to it within body, combined thickness of muscle even though some differences occur. Nikolaeva layers 4-5. Acetabulum 90 by 87, strongly (1965) studied much smaller specimens than muscular, center at level of anterior 38 per ours, but inasmuch as they are immature size cent of body length. Sucker length ratio 1: alone may not be significant for differentiation. 0.76. Distance between suckers 615, its ratio For the same reason and because of differences to body length being 1:3.4. Pharynx 20 by 37, in the state of extension or contraction of the contiguous with entire width of truncated body following fixation, the ratios of width and posterior end of oral sucker; esophagus 440 of distance between suckers to body length are long; cecal bifurcation 148 preacetabular; ceca also unreliable as criteria for differentiation. not ascending at beginning, winding, with very Nikolaeva described a strongly muscular oral thick, amber-colored glands internally to ace- sucker for her specimens, but her illustration tabular level, length of glandular part of right appears to show that the inner part is non- cecum 205, left cecum 198, walls 10-22 thick; muscular. Should our interpretation of the 15 distinct, isolated, large, multinucleate, illustration be incorrect, then our specimens agranular glands distributed dorsally and ven- represent a different species. The only other trally external to thick parts of ceca, some immature didymozoid with a glandular stom- glands bipartite, varying in size from 24 by 26 achlike structure at the cecal bifurcation is to 29 by 55; ceca continuing to within 90 of Monilicaecum ventricosum Yamaguti, 1942, posterior extremity, with 10 enlarged chambers from the flesh and body cavity of Cololabis along postacetabular length of each cecum, saira (Brevoort) (Scomberesocidae) from the chambers for short distance small and con- Japanese Pacific. The latter species differs spicuously cell-lined. Excretory bladder com- from Didymozoidae (Monilicaecum} larvae I mencing prebifurcal, 130 posterior to oral in having the stomachlike structure confined to sucker, arms postpharyngeal. the cecal bifurcation rather than extending DISCUSSION: This new form differs signif- posteriorly 011 the ceca to acetabular level, and icantly from Monilicaecum ventricosum Yama- in lacking glands external to the stomach. The guti, 1942, and Didymozoidae (Monilicaecum) presence of glands surrounding the anterior larvae I Nikolaeva, 1965, in having consider- part of the digestive tract has not been noted ably more of the ceca internally lined with for any other immature didymozoid; they have glands, a differently shaped pharynx, and an been described for adults of Nematobothrium oral sucker distinctly longer than the acetab- sardae MacCallum and MacCallum, 1916, and ulum. It differs further from the latter form species in the genera Paragonapodasmius in that the ceca do not ascend the body before Yamaguti, 1938; Allonematobolhrium Yama- extending posteriorly, and in having a different guti, 1965; Metanematobothrioides Yamaguti, type of gland external to the thick part of 1965; Nematobothrioides Yamaguti, 1965; and the ceca. Neonematobothrium Yamaguti, 1965. Immature Didymozoid E Immature Didymozoid D (Fig. 8) (Fig. 7) HOSTS: Scyris alexandrinus (Geoffroy St. HOST: Lagocephalus laevigatus (L.), smooth Hilaire), thread-fin horse mackerel; Caranx puffer or globe-fish (Tetraoclontidae). africanus Steindachner, African horse mackerel

(Carangidae); Cynoscion macrognathus (Blee- or nearly so). Excretory bladder commencing ker), large-mouth weakfish (Sciaenidae); Scom- pre- to slightly postbifurcal. beromorus tritor (Cuvier and Valenciennes), DISCUSSION: The distribution of the glands Spanish mackerel or kingfish (Scombridae); external to the anterior part of the digestive Pomadasys fubeUni (Cuvier and Valencien- tract is distinctive for this new form, dif- nes), burro; Brachydeuterus auritus (Cuvier ferentiating it from all other known immature and Valenciennes), burrito (Pomaclasyidae); forms of the family. The distribution is some- Psettodes belcheri Bennett (Psettodidae); what similar to that reported for the adult of Cynoglossus goreensis Steindachner, C. sene- Nematobothrioides kalikali Yamaguti, 1965, galensis (Kaup), tongue soles (Cynoglos- from Hawaii; however, the latter lacks an sidae); Rhinobatus albomaculatus Norman, acetabulum. The glands in specimens from white-spotted guitarfish (Rhinobatidae); Ptero- the guitarfish and ray were sparser and more platea micrura (Schneider), butterfly ray diffuse than those from the other hosts possibly (Trygonidae). because they, as selachians, are not suitable HABITATS: Stomach, small intestine. paratenic hosts. LOCALITIES: Tema, Cape Coast, Iture; Ghana. Immature Didymozoid F DATES: 19 December 1964; 31 March, 2 (Fig. 9) April 1985; 21 February, 10 March 1966. HOST: Cypsilums heterunis (Rafinesque), SPECIMENS: USNM Helm. Coll. No. 63345 flying-fish (Exocoetidae). (from Scyris); No. 63346 (Caranx); No. HABITAT: Small intestine. 63347 (Cynoscion); No. 63348 (Scombero- LOCALITY: Tema, Ghana. morus); No. 63349 (Pomadasijs}: No. 63350 DATE: 2 June 1965. (Brachydeutenis}; No. 63351 (Psettodes); SPECIMEN: USNM Helm. Coll. No. 63356. No. 63352 (Cijnoglossus goreensis); No. 63353 DESCRIPTION (based on one complete speci- (C. senegalensis); No. 63354 (Rhinobatiis); men in dextrolateral view, hence measurements No. 63355 (Pteroplatea). are length by depth; also, two fragments of DESCRIPTION (based on 19 specimens; 10 postacetabular part of body of two other speci- measured): Characteristics as for Diclymozoi- mens ): Characteristics as for Didymozoidae dae (Monilicaecum) larvae I, except as noted (Monilicaecum) larvae I, except as noted below. Body 1,500-3,501 by 165-510, pos- below. Body 3,597 by 116, very long. Fore- terior extremity round; body 5.8—12.5 times body 515 long, hinclbody 3,060 long. Oral longer than wide. Forebody 385-663 long, sucker 45 by 39, entirely but weakly muscular, hindbody 1,015-2,830 long. Oral sucker Si- within body. Acetabulum 22 by 26, center at ll 1 by 37-80, usually longitudinally elongate level of anterior 15 per cent of body length. but may be round, entirely but weakly mus- Sucker length ratio 1:0.49. Distance between cular. Acetabulum 100-222 by 98-218, center suckers 465, its ratio to body length being at level of anterior 17-29 per cent of body 1:7.7. Pharynx 27 by 21, entirely muscular; length. Sucker length ratio 1:1.71-2.52. Dis- esophagus 252 long, lumen not enlarged distance between suckers 270-542, its ratio to tally; stomach at cecal bifurcation, 45 by 33, body length being 1:4.5-8.6. Pharynx 18-48 lying 150 preacetabular, lined internally with by 14-36, round to longitudinally elongate, very thick, amber-colored glands; ceca 9-10 wide at exit from stomach, straight to short entirely muscular, contiguous with center of distance postacetabular, then sinuous with posterior part of oral sucker; esophagus 191- some small, elongate chambers, extending to 404 long, straight or sinuous, lumen not much within 32 of posterior extremity. Glands in enlarged at distal end; cecal bifurcation 46-123 compact mass external to stomach and ceca preacetabular, no glandular thickening of walls; for distance of 1,045, terminating 620 post- ceca winding to within 27-190 of posterior acetabular. Excretory bladder commencing extremity, each cecum with 12—16 chambers. prebifurcal, arms to 63 postpharyngeal. Glands in compact mass externally surrounding DISCUSSION: The present new form appears posterior 78-189 of esophagus, cecal bifurca- closest to Monilicaecum ventricosum Yamaguti, tion and 53-140 of ceca (to acetabular level 1942, but the latter differs in possessing non-

Copyright © 2011, The Helminthological Society of Washington 246 PROCEEDINGS OF THE HELMINTHOLOG1CAL SOCIETY muscular elements in the oral sucker, in the HABITAT: Encysted in subepidermal tissue retracted oral sucker being smaller than or the of buccal cavity. same size as the acetabulum, and in lacking LOCALITY: Tema, Ghana. glands external to the anterior part of the SPECIMENS: USNM Helm. Coll. No. 63358 digestive tract. The distribution of the latter (holotype, acetabulum prebifurcal); No. 63359 glands in our form is somewhat similar to that (paratypes). occurring in adults of Paragonapodasmius DIAGNOSIS (based on fragments of seven managatuwo Yamaguti, 1938, and Metanemato- specimens): Body very long, extremities bothrioides opakapaka Yamaguti, 1965. round; anterior part of body (prebifurcal) wide, gradually becoming narrower postbifur- Immature Didymozoid G cally, then gradually widening again to slightly (Fig. 10) greater width than prebifurcally; in one specimen 1,000 wide at pharyngeal level and 300 HOST: Larimus peli Bleeker (Sciaenidae). at distance 7,570 postbifurcal; in another 1,250 HABITAT: Small intestine. wide at pharyngeal level and 400 at distance LOCALITY: Cape Coast, Ghana. 5,355 postbifurcal; may be up to 1,300 wide DATE: 19 January 1966. more posteriorly. Oral sucker 540-677 by SPECIMEN: USNM Helm. Coll. No. 63357. 510-630, terminal, within body, well devel- DESCRIPTION (based on single specimen): oped, muscular, 12-30 from anterior extrem- Characteristics as for Didymozoidae (Moni- ity. Acetabulum 80-105 long, 104-108 wide licaecum) larvae I, except as noted below. (in four), 65—70 deep (in three), transversely Body 1,350 by 360, 3.8 times longer than oval; in four lying 180 prebifurcal (.1,0.1.0 wide. Forebody 340 long, hindbody 815 long. posterior to oral sucker), 218 (625), 465 Oral sucker 83 by 64, entirely but weakly (915), and 795 (1,065), respectively; in three muscular, within body. Acetabulum 195 by lying 180, 395, and 805 postbifurcal, respec- 200, center at level of anterior 32 per cent tively. Sucker length ratio 1:0.14-0.19. of body length. Sucker length ratio 1:2.35. Pharynx 127-208 by 109-140, muscular, in- Distance between suckers 260, its ratio to verted bell-shaped, contiguous with postero- body length being 1:5.2. Pharynx 27 by 27, median part of oral sucker, overlapping latter entirely muscular; esophagus 255 long, 19-100; esophagus 900-2,350 long, straight to straight, lumen not enlarged distally; cecal slightly sinuous, thick-walled, muscular; ceca bifurcation overlapping anteriormost part of very long, narrow, extending to posterior ex- acetabulum; ceca thick-walled, muscular at tremity. Glands surrounding pharynx, esoph- acetabular level for distance of 103, with outer agus, and ceca for distance of 265—475. thin longitudinal and inner thick circular mus- Testes two, symmetrical, tubular, uncle layers which are continuous with those of branched, sinuous, commencing posterior to esophagus, walls 3-5 thick, lumen narrow; genital junction, usually slightly subequal ante- glands in compact mass external to muscular riorly, anteriormost extent 11.6-14.7 mm from part of ceca; beyond latter ceca conspicuously anterior extremity of body. Vas efferens from cell-lined, with 12 chambers along length of anterior tip of each testis, slightly swollen. Vas each cecum, amber-colored material absent deferens very long, ascending next to me- from chambers, terminating 125 from posterior traterm, distalmost end thick-walled, muscular extremity. Excretory bladder followed ante- for very short distance. Genital atrium short, riorly as far as acetabulum, but probably ex- relatively narrow, thick-walled, muscular, pro- tending preacetabular; pore terminal. truding from genital pore in one specimen. DISCUSSION: This new form differs from all Genital pore median, ventral to approximate known immature didymozoids in having the micllength of oral sucker. ceca muscular at their beginning. Ovary median, tubular, unbranched, winding considerably, 18.5-28.7 mm in longitudinal Allonematobothrium ghanensis n. sp. extent (in three), commencing 1.2-7.4 mm (Figs, lla-e) posterior to anteriormost tip of testes, lying HOST: Epinephelus aeneus (Geoffroy St. 12.8-21.6 mm from anterior extremity of body. Hilaire), sea perch or grouper (Serranidae). Genital junction 40.1-43.7 mm from anterior

Copyright © 2011, The Helminthological Society of Washington OF WASHINGTON, VOLUME 35, NUMBER 2, JULY 1968 247 extremity (in three); forming enlarged cham- sucker length ratio 1:0.47. The pharynx is ber 97-150 by 126-250 (in four), ovary 77 by 87, slightly overlapping the oral sucker emerging from anteromedian surface, single dorsally; the anterior end is shaped like an duct emerging from posteromedian surface; inverted bell with the clapper protruding latter duct receiving vitellarium 15-30 (in slightly, while the posterior end is broad and four) from its origin, winding to posterior truncate. A vitelline reservoir is present; this extremity as uterus after receiving vitellarium. is not indicated in a paratype illustrated (Fig. Latter tubular, winding considerably, com- 10D) by Yamaguti. The Mehlis' gland is 1,060 mencing near posterior extremity, enlarging long, whereas it is illustrated in the same into vitelline reservoir 165-220 by 77-175 paratype as being considerably shorter. Fin- (in four) just before joining duct from genital ally, the distalmost ends of the vas deferens junction. Seminal receptacle absent. Mehlis' and metraterm as well as the genital atrium gland extensively developed, surrounding are thick-walled and muscular as described for genital junction and continuing posteriorly our species. around uterus for distances of 1,805 and 2,275 in two, respectively, filling much of body Literature Cited width. Metraterm very long; especially thick- Caballero y C., E. 1952. Revision de los gen- walled and muscular before entering genital eros y especies quc integran la familia Acan- atrium. Eggs numerous, yellow-brown, oval, thocolpidae Liihe, 1909 (Trematoda: Di- 43 measuring 15-20 by 10-13. genea). Rev. Med. Vet. Parasit, Caracas 11: Excretory bladder elongate, narrow, sinuous, 1-231. extending prebifurcal, arms extending ante- Tischthal, J. H., and R. E. Kuntz. 1964. Di- riorly as far as level of posterior margin of genetic trematodes of fishes from Palawan Island, Philippines. IV. Some immature Didy- oral sucker; pore terminal. mozoiclae, a bucephalid; a new hemiuroid DISCUSSION: Allonematobothrium epinepheli genus and subfamily. J. Parasit. 50: 253-260. Yamaguti, 1965, the type and only species in , and . 1965. Digenetic trema- the genus, was found encysted in pairs in the todes of amphibians and reptiles from North fins and underside of the operculum of Epi- Borneo (Malaysia). Proc. Helm. Soc. Wash. nephelus quernus Scale from Hawaii. Our form 32: 124-136. differs from the type species in possessing oval Manter, H. W., and H. J. Van Cleave. 1951. eggs, and a considerably larger oral sucker and Some cligenetic trematodes, including eight slightly smaller acetabulum, and in the genital new species, from marine fishes of Lajolla, Calif. Proc. U.S. Natl. Mus. 101: 315-319. junction forming an enlarged chamber. In A. Nikolaeva, V. M. 1964. [Metacercariae in the ghanensis four specimens had the acetabulum trematode family Didymozoidae (Monticelli, lying prebifurcal, and three postbifurcal. No 1888) Poche, 1907 from fishes of the Mediter- other difference was noted between them. ranean Sea basin.] Trudy Ukrain. Respublik. Therefore, in spite of the extensive variation Nauch. Obshchest. Parasit., Probl. Parasit. 3: in the position of the acetabulum (795 pre- 53-68. (Russian text.) bifurcal to 805 postbifurcal), we are listing . 1965. [On the development cycle of them as a single species until such time as life trematodes belonging to the family Didymo- zoidae (Monticelli, 1888) Poche, 1907.] Zool. history studies may determine otherwise. Zh. 44: 1317-1327. (Russian text.) Yamaguti (1965) noted the acetabulum as Parukhin, A. M. 1966. [Toward a study of the being 250—800 postbifurcal in the seven speci- helminthofauna of the fish family Carangidae mens of A. epinepheli studied. from the South China Sea.] In Gel'mintofauna Examination of the holotype specimen of Zhivotnykh luzhnykh Morei. Respublik. Me- A. epinepheli (USNM Helm. Coll. No. 63526) zhvedomst. Shorn., Ser. "Biol. Moria," Akad. Nauk UkSSR, p. 80-96. (Russian text.) has made it possible to slightly amplify Yama- Yamaguti, S. 1965. New digenetic trematodes guti's description. The oral sucker measures from Hawaiian fishes, I. Pacific Sci. .19: 245 by 257 and the acetabulum 115 by 196; 458-481.

Five New Species of Belondiroidea (Nematoda) from Sibsagar, India, with a Revised Classification of the Superfamily MOHAMMAD RAFIO SIDDIQI*

Soil samples collected on 25 July 1966 by D. paraclavatus n. sp.—Rose (Rosa sp.). Mr. Ajai Kumar of the Oil and Natural Gas D. parvulus Thorne, 1939—Rose, banana. Commission of India around roots of various D. salmis n. sp.—Banana. plants at Sibsagar, Assam State, contained 15 different species of belondiroid nematodes, Axonchium asacculum n. sp. five of which were previously undescribecl. (Fig. 1,A-F) These species are listed below, together with Measurements the names of the associated plants; the first FEMALES (12) (Paratypes): L = 0.97- name against a new species is that of the type 1.17 mm; a = 38-49; b = 1.9-2.4; c = 51-65; plant with which it is associated; botanical V = 53-57. names are mentioned only once, FEMALE (Holotype): L = 1.05 mm; a = Axonchium amplicolle Cobb, 1920—Coco- 40; b = 2.2; c = 57; V = 53-12. nut (Cocos nucifera L.), Elephant-foot DESCRIPTION: Female: Lip region amalga- yam (Amorphophallus campanulatus mated, rounded, continuous with body con- (Roxb.) Blume ex Dene.). tour. Spear well developed, 7-8 p, long. A. asacculum n. sp.—Sweet orange (Cit- Hemizonid 92 p. from anterior end in holotype, rus sinensis (L.) Osbeck), banana near anterior edge of nerve ring. Esophagus (Musa paradisiaca L.), bottle gourd 400-530 p,; anterior part 170-210 p.; esophago- (Lagenaria siceraria (Mol.) Stand!.), intestinal valve large, oval. Esophagus-vulva chillie (Capsicum frutescens L.), cot- distance usually 100-190 p, (in one young ton (Gosst/pium arboreum L.), jack female this distance is only 30 p,). Vagina tree (Artocarpus heterophyllus Lamk.). elongate, longitudinal to body axis. Anterior A. elegans Jairajpuri, 1964—Elephant ear uterine branch absent. Ovary with multiple (Colocasia esculenta (L.) Schott). rows of oocytes. Prerectum 103-140 p, long; A. nitidum Jairajpuri, 1964—Lemon (Cit- rectum equal to or longer than anal body rus limon (L.) Burm.), jack tree, litchi width. Tail obtusely rounded, equal to or (Litchi chinensis Sonn.). a little less than anal body width. Male not A. saccatum Jairajpuri, 1964—Chillie. found. Belondira caudata Thorne, 1939—Ber- TYPE MATERIAL: Holotype and two para- muda grass (Cynodon dactylon (L.) types at Nematology Department, Rothamsted Pers.), lemon; date palm (Phoenix Experimental Station, Harpenden, England; dactylifera L.), areca nut (Areca six paratypes at Commonwealth Bureau of catechu L.), cotton. Helminthology, St. Albans, England; two para- B. clavicaudata (Williams, 1958) An- drassy, 1963—Lemon, jack tree. types in Zoology Museum, Aligarh Muslim B. murtazai n. sp.—Litchi, date palm, University, Aligarh, India; two paratypes at coconut, areca nut. Crops Research Branch, USDA, Beltsville, B. neortha Siddiqi, 1964—Jack tree, Maryland, USA. lemon. RELATIONSHIP: A. asacculum n. sp. comes B. nepalensis Siddiqi, 1964—Elephant close to A. amalgans Thorne, 1939 and A. ear, litchi. indicum Siddiqi, 1964 from both of which it Donjlaimellus belondirelloides n. sp.— differs in the absence of the anterior uterine Bottle gourd. branch. From A. amplicolle Cobb, 1920; A. D. discocephalus Siddiqi, 1964—Bermuda bulbosum Williams, 1958 and A. nitidum Jairajpuri, 1964 it can easily be differentiated by its smaller body size, continuous lip region, * Plant Nematologist, Commonwealth Bureau of Helm- inthology, St. Albans, Hertfordshire, England. and the absence of an anterior uterine branch.

15 -M ADE

20^ BCF

Figure 1, A-F. Axonchiutn asaccnlum n. sp. A. Head end of female. B. A portion of female reproductive organs. C. Tail end of female. D. Junction of two parts of esophagus. E. En face view of female. F. Vulva near esophageal base in immature female.

Belondira murtazai n. sp.* Belondira nepalensis Siddiqi, 1964 (Fig. 2, A-D) (Fig. 2, E-H) Measurements Male and female specimens of Belondira FEMALES (12) (Paratypes): L = 0.85- nepalensis Siddiqi, 1964 were found in these 1.06 mm; a = 40-50; b = 4.3-4.9; c = 50-55; samples. Measurements and diagrams of these V = 36-38. are provided here and compared with those of MALES (12) (Paratypes): L = 0.75-0.99 the holotype. Males of this species are being mm; a = 43-50; b = 4.3-5.1; e = 47-55; reported for the first time. T = 33-48; spicules = 19-22 p.. FEMALE (Holotype): L = 0.98 mm; a = Measurements 43; b = 4.9; c = 54; V = 6-37-14. FEMALES (4): L = 0.83-0.95 mm; a = 36- DESCRIPTION: Female: Body slightly arcu- 43; b = 4.7-5.1; c = 44-54; V = 32-35; ate ventrally, tail end clavate. Inner layer of spear = 3-4 p,; anterior uterine sac = lVi-1% cuticle distinctly striated; striae fine. Spear body width. 3-4 p. long, aperture one-third its length; MALES (3): L = 0.84-0.92 mm; a = 43- extension in two parts, 12-16 p, long. Esoph- 55; b = 4.2-4.7; c = 49-52; T = 40-46; agus enlarging just behind its middle. Vagina spicules = 30-32 p. straight, leading two-fifths to halfway into FEMALE (Holotype, after Siddiqi, 1964): body. Anterior uterine sac 2% to 3 times body L = 0.89 mm; a = 42; b = 5.1; c = 56; V = width. Prerectum 2% to 3 times body width. 33.6; spear = 3 p. (spear length was mis- Tail rounded, about one anal body width long; printed as 7 p. in the original description). its inner mass convex-conoid, about 10 p, long. MALE: Body slightly arcuate ventrally in MALE: Spicules robust, ventrally bent near posterior half. Cuticle and subcuticle marked middle; copulatory muscles strongly developed. by transverse striae which are more prominent First supplementary, ventro-median papilla towards extremities. Testes dorylaimoid, with 42-47 p, from anus and the second 51-60 p. few spermatocytes. Spicules slender, ventrally from the first. Body with a sharp depression arcuate but not sharply bent, poorly ceph- behind anus. alated. Supplementary papillae consist of a preanal pair and two ventromedians: the first TYPE MATERIAL: Holotype and two pairs of located 42-60 p, from anus and the second paratypes (2 5,2 ?) at Nematology Depart- 50-60 p, from the first. Tail bluntly rounded, ment, Rothamsted Experimental Station, Har- equal to or a little longer than anal body penden, England; two pairs of paratypes (2,8, width; its inner layer of cuticle is more thick- 2 9 ) distributed to each of the following ened than the outer. centers: Commonwealth Bureau of Helmin- thology, St. Albans, England; Plantenziekten- Dorylaimellus belondirelloides n. sp. kundige Dienst, Wageningen, The Nether- (Fig. 2,1-K) lands; Department of Zoology, Aligarh Muslim University, Aligarh, India; and USDA, Belts- FEMALE (Holotype): L = 0.76 mm; a = ville, Maryland, USA. 40; b = 4; c = 23- V = ('-47.5~fi. RELATIONSHIP: B. murtazai n. sp. is very FEMALE (Paratype): L = 0.73 mm; a = closely related to B. nepalensis Siddiqi, 1964 37; b = 4,5; c = 21; V = 48. JUVENILE (I): L = 0.44 mm; a = 27; b = from which it differs in having a clavate tail 3.4; c = 18; (II): L = 0.53 mm; a = 33; b end, straight and shorter vagina (vagina open = 3.1; c = 18. S-shaped, more than half body width long in DESCRIPTION: Female: Body slightly arcu- B. nepalensis), longer anterior uterine sac and ate ventrally. Cuticle with faint transverse a little more posterior vulva. Also its males striae; hypodermis coarsely striated; lateral have shorter and differently shaped spicules. hypodermal chords one-fourth body width. Lip region rounded, slightly offset, with raised :;: Named after Mr. Miirtaza Ali Ouadri, Lecturer, S.N.I. papillae; inner papillae forming six-lobed, non- College, Pilibhit, U.P., India who arranged and supplied this collection. refractive disc; pharyngeal sclerotization around

15 A,D,E, I-K 25 -B,C,F-H

K Wif-^j/;;V^/S;-:^xl>>:ii.;•./:•;,;, "y!m K; >-.....m.

iiiii'•I'D •••'^l^'S&f'-^i-^^'Si

Figure 2, A-D. Belondira murtazai n. sp. A. Head end of female. B. Vulval region. C. Tail end of female. D. Tail end of male. E-H. B. nepalensis Siddiqi, 1964. E. Head end of female. F. Tail end of male. G. Tail end of female. H. Vulval region. I-K. Dorylaimellus belondirelloides n. sp. I. Head end of female. J. Tail end of female. K. Vulval region.

Figure 3, A-C. Dorylaimellus paraclavatus n. sp. A. Head end of female. B. Vulval region. C. Tail end of female. D-H. D. salvus n. sp. D. Head end of female. E. Tail end of female. F. Vulval region. G. Spicular region. H. Spicule.

Copyright © 2011, The Helminthological Society of Washington OF WASHINGTON, VOLUME 35, NUMBER 2, JULY 1968 253 mouth easily seen. Spear 5-6 ^ long, aperture rectum shorter than anal body width. Tail a quarter of its length; extension poorly flanged. dorsally convex-conoid at first, then elongate- Amphidial pouches large. Esophageal enlarge- clavate, 4.3 times anal body width long (tail ment muscular, 46 per cent of neck length; end in paratype female less clavate than that esophago-intestinal valve large, heart-shaped. in holotype as depicted in Fig. 3, C). Nerve ring 67 ^ from anterior end; hemizonid JUVENILE: Cuticle and subcuticle distinctly near its anterior edge. Ovaries paired, short. striated. Spear 6 ^ long; developing spear Prerectum 115 ^ long; rectum a little longer 7 ju, long. Esophagus enlarged at 53 per cent than anal body width. Tail cylindroid-sub- of its length. Tail as in female but less clavate, thrice anal body width long; cuticle clavate. at terminus thickened. Male not found. TYPE MATERIAL: Holotype at Nematology TYPE MATERIAL: Holotype at the Nema- Department, Rothamsted Experimental Sta- tology Department, Rothamsted Experimental tion, Harpenden, England; rest at Common- Station, Harpenden, England; rest at Common- wealth Bureau of Helminthology, St. Albans, wealth Bureau of Helminthology, St. Albans, England. England. RELATIONSHIP: In tail shape, D. paracla- RELATIONSHIP: D. belondirelloides n. sp. vatus n. sp. resembles D. davatus Thorne, differs from D. caffrae Kruger, 1965 and D. 1964, D. nygellurus Loof, 1964; D. spicatus projectus Heyns, 1962 in having a more an- Loof, 1964 and D. virginianus Cobb, 1913. teriorly located vulva and longer prerectum D. davatus and D. spicatus are smaller and and tail. have a smaller value for "b" ratio (3.2 or less), a postequatorial vulva and vulva- Dorylaimelhis paraclavatus n. sp. esophagus distance being 75 per cent or (Fig. 3, A-C) less of neck length. D. nygellurus is smaller, has a smaller spear extension, relatively longer Measurements tail and smooth cuticle. D. virginianus as re- FEMALE (Holotype): L = 1.1 mm; a — 40; described by Thorne (1939), is larger and b = 4.5; c = 15; V = (i-48-°. has a more posterior vulva and shorter tail. FEMALES (2) (Paratypes): L = 1.02-1.10 mm; a = 37-38; b = 4.3; c = 15; V = 47-49. Dorylaimellus salvus n. sp. JUVENILE (I): L = 0.8 mm; a = 36; b = (Fig. 3, D-H) 3.6; c = 14. DESCRIPTION: Female: Body slightly arcu- Measurements ate ventrally. Cuticle and subcuticle coarsely FEMALES (3) (Paratypes): L = 1.17-1.23 striated; striae more prominent over extrem- mm; a = 47-53; b = 6.5-7.5; c = 8-10; V = ities. On one side, about 10 lateral body pores 47-49. in neck region, 9 in esophagus-vulva and 17 MALES (3) (Paratypes): L = 1.10-1.17 in vulva-anus region, and 2 on tail. A ventral mm; a = 46-51; b = 6.5-7.0; c = 8.5-9.5; series of prominent body pores including 9 in T = 37-47. region of esophagus, 10 in esophagus-vulva, FEMALE (Holotype): L = 1.23 mm; a — and 18 in vulva-anus region. Lip region 53; b = 7.2; c = 9.2; V = 1Q-47.4-8. rounded, offset; papillae prominent. Amphid DESCRIPTION: Female: Body slightly arcu- pouches almost as wide as body at that region. ate ventrally. Cuticle and subcuticle marked Spear well developed, with wide lumen and by distinct transverse striae. Lateral chords aperture about one-third its length, 7 ^ long; with prominent glandular bodies. Lip region extension well developed, twice its length. rounded; inner margins forming a pseudolabial Hemizonid opposite nerve ring, which lies disc. Amphids large. Spear with narrow lu- 70 /A from anterior end. Esophago-intestinal men, 5 /ji long; extension 10—12 ^ long, poorly valve rounded. Posterior 53-56 per cent of flanged. Esophagus enlarging in its posterior esophagus enlarged. Esophagus-vulva distance third. Esophago-intestinal valve rounded. Va- 115 per cent of neck length. Vagina extending gina straight, extending halfway into body. one-third into body. Ovaries short, symmet- Ovaries paired, symmetrical. Prerectum 5-6 rical. Prerectum about four body widths long; times body width long; rectum a little longer

Copyright © 2011, The Helminthological Society of Washington OF WASHINGTON, VOLUME 35, NUMBER 2, JULY 1968 255 than anal body width. Tail elongate-conoid to Belondiridae and Nygellidae as synonymous a finely rounded terminus, 9-10 times anal with Dorylaimidae and Nygolaimidae respec- body width long. tively. Male: Essentially similar to female. Spicules In certain species of Discolaimus and Dis- stout, 24-26 p. long. Besides a pair of sub- colaimium of the family Dorylaimidae the median papillae near anus there are five esophageal enlargement is covered with a ventro-median papillae spaced from anus at a thick sheath of connective tissue which, in distance of 29-44 ^ 46-50 ^ 73-80 p., 92- Discolaimus bicorticus, is double-layered and 99 p, and 102-110 p.. appears thickened at three points in cross- TYPE MATERIAL: Holotype and a pair of section (Furstenberg and Heyns, 1966). How- paratypes (1 $, 1 2 ) at Nematology Depart- ever, such a sheath is not muscular and should ment, Rothamsted Experimental Station, Har- not be confused with the muscular spiral penden, England; a pair of paratypes (IS, sheath of the Belondiroidea. Yeates' (1967) 12) at Commonwealth Bureau of Helminthol- conclusions are based mainly on his examina- ogy, St. Albans, England; a pair of paratypes tion of Dorylaimettus tahatikus Yeates, 1967a, (1$, 12) at USDA, Beltsville, Maryland, in which this sheath is not recognizable in USA. living worms but a slight indication of it ap- RELATIONSHIP: In tail shape, D. salmis re- pears after fixation. I have examined the types sembles D. donjlaimoidunis Siddiqi, 1966; D. of D. tahatikus and consider that this species salimi Siddiqi, 1966; D. filicaudatus Thome, belongs to Tylencholaimus of the family Dory- 1964; and D. filiformis Jairajpuri, 1964. The laimidae because of the shape and structure first two are smaller, have an attenuated spear, of its cuticle, head, spear, esophagus, and fewer supplementary papillae in male and gonads. Accordingly I propose that it be spicules 17 //, long. The third has a longer redesignated as Tylencholaimus tahatikus esophagus and tail (b = 3, c = 5) and esoph- (Yeates, 1967) n. comb. agus enlarged in its posterior two-thirds, and Evidence for the muscular nature of the the fourth is smaller, has smooth cuticle and sheath around the esophageal enlargement of subcuticle, angular lip region devoid of labial Belondiroidea seems to be limited to that of disc and vulva at 43 per cent of body. Hopper (1961), who reported six conspicuous spiral muscles capable of changing the A revised classification of the length of the esophageal enlargement by their Belondiroidea contraction or relaxation in Swangeria bisex- The classification of the superfamily Belon- ualis Hopper, 1961. Likewise, in Falcihasta diroidea (Thorne, 1939) Thome, 1964 is paliistris Clark, 1964, I found six bands of mainly due to Thorne (1964). The major strong muscles around the esophageal enlarge- character of this superfamily is the presence ment which, in most of the specimens exam- of a conspicuous sheath of spiral muscles ined, were apparently responsible for the around the enlarged basal part of the esoph- dorsally bent condition of the esophageal en- agus. Clark (1961) was doubtful of the mus- largement (Fig. 4, c). I have also examined cular nature of this sheath as he mentioned a number of transverse sections of the esoph- a "sheath of connective tissues" in diagnosing ageal enlargement in Belondira, Donjlaimellus, the family Belondiridae. Yeates (1967) con- and Axonchium, in all of which about 12 siders that the muscular esophageal sheath of muscles were found surrounding the basal part belondirids is "a fixation artifact" and regards of the esophagus (Fig. 4, F-H, K). These

Figure 4. A-K. A. Belondira neortha Siddiqi: Anterior portion of esophagus (Belondirid type). B-D. Falcihasta palustris Clark: B. Head end of female, C, D. Basal region of esophagus. E. Axonchium asacculum n. sp.: Anterior portion of esophagus (Axonchiid type). F-K. T.S. through esophagus: F. Belondira murtazai n. sp.: Near middle of esophageal enlargement. G. Dorylaimellus parvulus Thorne: Through dorsal esophageal gland. H. Axonchium asacculum n. sp.: Near middle of esophageal enlargement. I-K. Axonchium nitidum Jairajpuri: I. Near base of anterior part of esophagus. J. Just behind nerve ring. K. Near middle of esophageal enlargement.

Copyright © 2011, The Helminthological Society of Washington 256 PROCEEDINGS OF THE HELMINTHOLOG1CAL SOCIETY muscles are similar to somatic muscles in con- name, since Nygolaimellinae was already avail- sistency. With this evidence at hand and the able. Nygolaimellinae is here raised to family fact that the group is also recognizable by rank as Nygolaimellidae (Clark, 1961) n. rank. other subsidiary characters, I recognize Belon- Moreover, since the bibulbar appearance of cliroidea as proposed by Thome (1964) and the esophagus in Nygolaimellus abnormis Loos, differ from Yeates (1967) who considers 1949 is considered less important because an- Belondiridae and Nygellidae as junior syn- other species, N. capitivitatus Andrassy, 1962, onyms of Dorylaimidae and Nygolaimidae has almost a normal esophageal enlargement, respectively. Nygellidae Jairajpuri, 1964, as well as Nygel- Thorne (1964) recognized the following linae Jairajpuri, 1964, are here suppressed as families under Belondiroidea: Belondiridae synonyms of Nygolaimellidae. Thorne, 1939 (subfamilies Belondirinae Jairaj- puri, 1964; Axonchiinae* Thorne, 1964); Oxy- Family Belondiridae Thorne, 1939 diridae** Thorne, 1964; Roqueidae Thorne, Syn. Belondirinae Jairajpuri, 1964 1964; Dorylaimellidae (Jairajpuri, 1964) Belondirinae de Coninck, 1965 Thorne, 1964; Mydonomidae Thorne, 1964; Dorylaimellinae Jairajpuri, 1964 Nygellidae Jairajpuri, 1964 (subfamilies Ny- Dorylaimellidae (Jairajpuri, 1964) gellinae Jairajpuri, 1964; Nygolaimellinae Thorne, 1964 Clark, 1961). I feel that the proposal of Dory- Dorylaimellinae de Coninck, 1965 laimellidae is unjustified, as the structure of Opailaimidae Kirjanova, 1951 the esophagus, buccal armature, and the num- DIAGNOSIS: Belondiroidea: Spear usually ber of muscles around the enlarged part of the small with narrow lumen; spear extension esophagus are the same in Belondira and in appearing in two parts. Esophagus with a Dorylaimellus. In many species of Belondira, spinclle-shapecl swelling often with visible for example B. dcwicaudata, B. bulbosa, B. radial musculature following spear extension, nepalensis, the spear extension, like that in becoming extremely thin and nonmuscular as Dorylaimellus, is divisible into two parts, the it passes through the nerve ring, gradually basal part bearing flanged swellings. The enlarging into a cylindrical bulb enveloped by other feature, the presence of three or four narrow bands of spiral muscles, about twelve sclerotized platelets around the vestibule, is in number. Esophago-intestinal valve simple, sometimes hard to detect and in D. parvulus conoid rounded. Intestine two to four cells in these pieces are hardly discernible even when circumference. Spicules well developed; sup- viewed en face. plements few. The esophagus in Axonchium (Fig. 4, E) is TYPE GENUS: Belondira Thome, 1939. so different from that in Belondiridae (Fig. 4, OTHER GENERA: Belondirella Thorne, 1964 A) that the subfamily Axonchiinae is here Dorylaimellus Cobb, 1913 given a familial status, necessitating the sup- Yunqueiis Thome, 1964 pression of Belondirinae and the redefinition Bullaenema Sauer, 1968 of Belondiridae. A new family, Falcihastidae, Axonchoides Thorne, 1967. is proposed to contain Falcihasta Clark, 1964, Family Axonchiidae (Thorne, 1964) n. rank which has an asymmetrical head, mural spear, Syii. Axonchiinae Thorne, 1964 and six spiral esophageal muscles. Jairajpuri (1964) proposed the family Ny- DIAGNOSIS: Belondiroidea. Moderately long gellidae in which he included Nygellinae (1 to 5 mm). Lateral chords narrow. Spear Jairajpuri, 1964 and Nygolaimellinae Clark, asymmetrical, rather spindle-shaped, axial, 1961. According to Article 36 of the Interna- with wide lumen. Anterior part of esophagus tional Code o£ Zoological Nomenclature, Ny- narrower towards cephalic end, comparatively gellidae would be invalid as the family-group wider posteriorly with distinct radial musculature and lumen, offset from the posterior '•'•• Original spellings of Axonchinae and Swangerinae enlarged part by a constriction or a short amended as Axonchiinae and Swangeriinae respectively. isthmus; enlarged part usually greatly devel- :::::: Thome's (1964) citation of this family as Oxydiridae (Jairajpuri, 1964) n. rank is an error, as this family-group oped, enveloped by about 12 bands of spiral name was not proposed by Jairajpuri (1964) or any other worker before Thorne (1964). muscles. Dorsal esophageal gland large, with

Copyright © 2011, The Helminthological Society of Washington OF WASHINGTON, VOLUME 35, NUMBER 2, JULY 1968 257 prominent nucleus near beginning of esopha- behind. Tails filiform in both sexes. Large geal enlargement. Esophago-intestinal valve spicules, with lateral guiding pieces. No gu- usually elongate-conical. Spicules with inter- bemaculum. Supplements few. nal stiffening piece; spicular guiding pieces TYPE GENUS: Falcihasta Clark, 1964. present; supplements numerous. No other genus. TYPE GENUS: Axonchium Cobb, 1920. Falcihastidae has a subventrally located No other genus. mural spear as in Nygolaimellidae (Clark, This family is at once recognized by the 1961) n. rank but differs from this family in anterior part of the esophagus being very the form of the head, spear and pharynx and muscular and offset from the posterior part the absence of three esophago-intestinal (Fig. 4, E). The genus Axonchoides Thorne, glands. 1967, which is somewhat similar to Axonchium, has a belondirid type of esophagus and thus Family Nygolaimellidae (Clark, 1961) belongs to Beloncliridae. n. rank Syn. Nygolaimellinae Clark, 1961 Family Swangeriidae (Jairajpuri, 1964) Nygellidae Jairajpuri, 1964 n. rank Nygellinae Jairajpuri, 1964 Syn. Swangeriinae Jairajpuri, 1964 DIAGNOSIS: Belondiroidea. Pharynx wide, long and tapering posteriorly, evertible; spear DIAGNOSIS: Belondiroidea. Medium-sized located on its left subventral sector. Esophagus (1-2 mm), with slender body. Vestibule narrow, gradually enlarging to a well-devel- moderately wide and thick-walled in type oped cylindrical portion which may appear genus. Spear symmetrical, elongate, axial. bibulbar due to disintegration of tissue in Esophagus slender with poor musculature, midregion. Esophago-intestinal valve small, extending back to spindle-shaped, muscular, with three glandular organs. Tail bluntly basal enlargement which is enveloped by six rounded. spiral muscles. Esophago-intestinal valve large, TYPE GENUS: Nygolaimelhif; Loos, 1949. elongate-cylindrical, with intestine attached to its posterior end in fixed specimens. Ovaries OTHER GENUS: Nygellus Thorne, 1939. paired. Males with large, slender spicules An outline classification with lateral guiding pieces. No gubemaculum. of the Belondiroidea Supplements few, widely spaced. Tail elon- Belondira Thorne, 1939 gate, filiform in both sexes. Belondirella Thorne, 1964 Beloncliridae Donjlaimelhis Cobb, 1913 TYPE GENUS: Stcangeria Thorne, 1939. Yunqueus Thorne, 1964 Bullaenema Saner, 1968 OTHER GENUS: Qudsiella Jairajpuri, 1967. Axonchoides Thorne, 1967 This family is recognized by the structure Axonchiidae Axonchium Cobb, 1920 of the esophagus, esophago-intestinal valve and few supplements in the male. From Falci- Mydonomidae Mydonomiis Thome, 1964 hastidae n. fain, it differs, besides the structure Roqueidae Roquens Thorne, 1964 of the esophagus, in having a symmetrical lip Oxydiridae Oxijdirus Thorne, 1939 region and axial spear. Swangeriidae Swangeria Thorne, 1939 Qudsiella Jairajpuri, 1967 Family Falcihastidae n. fam. Falcihastidae Falcihasta Clark, 1964 Nygolaimellidae Nygolahnelhis Loos, 1949 DIAGNOSIS: Belondiroidea. Medium-sized Nygellus Thorne, 1939 (1-2 mm). Few body pores. Lip region asymmetrical, lower ventrally than clorsally. Summary Pharynx broad, not sclerotizecl, carrying an Fifteen species of Belondiroidea have been asymmetrical spear on its subventral sector. reported from soil around roots of various Esophagus in three almost equal parts as in plants in Sibsagar, India. New species de- Myclonomidae, basal part not marked off by scribed are: Axonchium asacculum, Belondira a constriction, enveloped by six bands of murtazai, Dorylaimellus belondirelloides, D. muscles. Esoghago-intestinal valve large, cylin- paraclcwatus, and D. salmis. Males of B. ne- drical. Sensillar sacs of amphids rather far palensis Siddiqi are also described.

Comments are made on the spiral muscular Furstenberg, J. P., and J. Heyns. 1966. Four sheath around the esophageal enlargement of new species of the genus Discolaimus Cobb, Belondiroiclea disapproving of Yeates' (1967) 1913 (Nematoda: Dorylaimoidea) from South action in regarding it a fixation artifact. Donj- Africa, with a description of the male of D. laimellus tahatikus Yeates, 1967, on which si-nulls Thorne, 1939. Nematologica 11 (1965): 467-479. Yeates based his discussion in suppressing Heyns, J. 1962. A report on South African Belondiridae is redesignated as Tylencholaimus nematodes of the families Longidoridae, Be- tahatikus (Yeates, 1967) n. comb. Falcihasta londiridae and Alaimidae (Nemata: Dory- palustris Clark has been shown to possess six laimoidea), with descriptions of three new spiral muscles around esophageal enlargement species. Nematologica 8: 15-20. which are apparently responsible for the dor- Hopper, B. E. 1961. Sicangeria bisexualis n. sally bent condition of the latter in many fixed sp. (Belondiridae: Nematoda) from Florida. specimens. Transverse sections of the esoph- Canacl. J. Zool. 39: 69-72. ageal enlargement in Belcndira, Donjlaimellus Jairajpuri, M. S. 1964. Studies on Nygellidae n. fam. and Belondiridae Thorne, 1939 and Axonchium show 12 muscles around it, (Nematoda: Dorylaimoidea) with description similar in consistency to the somatic muscles. of ten new species from India. Proc. Helm. Following families have been recognized Soc. Wash. 31: 173-187. under Belondiroidea: Belondiridae Thome, Kruger, S. P. 1965. New species of the genera 1939; Axonchiidae (Thorne, 1964) n. rank; Ti/lencholaimiis and Donjlaimellus from South Mydonomidae Thorne, 1964; Roqueidae Africa. Proc. Helm. Soc.'Wash. 32: 1-7. Thome, 1964; Oxydiridae Thorne, 1964; Loot', P. A. A. 1964. Free-living and plant- Swangeriidae (Jairajpuri, 1964) n. rank; Falci- parasitic nematodes from Venezuela. Nema- tologica 10: 201-300. hastidae n. fam. and Nygolaimellidae (Clark, Siddiqi, M. R. 1964. Four new species in the 1961) n. rank. Belondirinae Jairajpuri, 1964; family Belondiridae (Nematoda: Dorylaim- Belondirinae de Coninck, 1965; Dorylaimel- ida). L. J. Sci. Tech. (India) 2: 37-41. linae Jairajpuri, 1964; Dorylaimellinae de Con- . 1964a. Six new nematode species in inck, 1965, and Dorylaimellidae (Jairajpuri, the Superfamily Dorylaimoidea from India. 1964) Thorne, 1964, have been synonymisecl Ibidem 2: 136-144. with Belondiridae. Swangeriinae Jairajpuri, 1966. Studies on species of Belondiro- 1964, and Axonchiinae have been given family idea (Nematoda: Dorylaimida) from India. Proc. Helm. Soc. Wash. 33: 139-149. rank and described. Falcihastidae n. fam. has Thorne, G. 1939. A monograph of the nema- mural, asymmetrical spear; esophagus in three todes of the superfamily Dorylaimoidea. almost equal parts and large, cylindrical esoph- Capita Zool. 8: 1-261. ago-intestinal valve. Nygellidae Jairajpuri, . 1964. Nematodes of Puerto Rico: Be- 1964, is regarded as invalid owing to the londiroidea new superfamily, Leptonchidae, availability of coordinate name Nygolaimellinae Thorne, .1.935, and Belonenchidae new family (Nemata, Adenophorea, Dorylaimida). Univ. Clark, 1961, which is raised to the rank of a Puerto Rico Tech. Paper No. 39, 51 pp. family and the former as well as Nygellinae . 1967. Nematodes of Puerto Rico: Ac- Jairajpuri, 1964, have been put down as tinolaimoidea new superfamily with a revision synonyms of the latter. of its genera and species with addenda to Belondiroidea (Nemata, Adenophorea, Dory- Literature Cited laimida). Univ. Puerto Rico Tech. Paper 43: 48 pp. Clark, W. C. 1961. A revised classification of Williams, J. R. 1958. Studies on the nematode the Order Enoplida (Nematoda). N. Z. J. Sci. soil fauna of sugar cane fields in Mauritius. 4: 123-150. 2. Belondiridae (Dorylaimoidea, Enoplida). . 1964. Falcihasta palustris n. gen. et sp., Occ. Paper No. 2, Mauritius Sug. Iiid. Res. family Belondiridae (Nematoda). N. Z. J. Sci. Inst. 1-9. 7: 177-180. Yeates, G. W. 1967. Observations on phylog- Cobb, N. A. 1913. New nematode genera found eny and evolution in the Dorylaimina (Nema- inhabiting fresh water and non-brackish soils. toda). N. Z. J. Sci. 10: 683-700. J. Wash. Acad. Sci. 3: 432-444. . 1967a. Studies on nematodes from dune . 1920. One hundred new nemas. Contr. sands. 6. Dorylaimoidea. N. Z. J. Sci. 10: Sci. Nematol. 9: 217-343. 752-784.

Studies on the Freshwater Cercariae of Northern Colorado

ALEXANDER D. AcHOLONU1 Department of Zoology, Colorado Slate University, Fort Collins, Colorado

Introduction where each snail collection was made was Cercariae emerging from their molluscan strained through four layers of cheesecloth to hosts, serve as a clue to the digenetic trema- filter out larger particles of debris. Wide- tode fauna of a region. More often than not, mouth 50 cc bottles were filled to the three- a cercaria and its corresponding adult are de- quarter level with this water. About 5 to 10 scribed and named separately and indepen- snails of the same species were put into each dently. It is not until the link between them bottle and the mouth was covered with a is connected by life history studies that the double-folded cheesecloth held in position by identity of the former is recognized. a rubber band to prevent the snails from Very little previous work has been done on crawling out. The bottles were placed in the the cercariae of Colorado. The earliest sig- part of the laboratory best lighted and allowed nificant work is that of Hurst (1923), who to stand for a maximum of 14 days. Snails described Cercaria giinnisoni as a new species which died during this period were crushed of xiphidiocercaria from Lijnmaea stagnates and examined for infection. The water in the appressa Say and L. proximo Lea. He stated bottles was checked daily for cercariae by that the former host harbored three different viewing the contents against a light source species of cercariae simultaneously. Of the and changed every other day. The snails in three, he described only C. gunnisoni. Lym- a bottle showing cercariae were isolated into naea Lamarck, Physa Draparnaud, and Planor- individual bottles to determine which were bis Guettard were abundantly represented in infected and to provide a source of cercariae his area of study, and cercariae were found in for morphological studies and information on all of them; he stated, "I am confident that emergence rate. At the end of 14 days, snails many of these cercariae are undescribed spe- which had not shed any cercariae were cies." Another significant work is that of crushed, examined for infection, and the inci- Crawford (1939, 1940, 1943), who conducted dence of infection tabulated. Some shed cer- studies on the life cycle of Colorado trema- cariae were transferred to a stender dish to tocles at Rocky Mountain Biological Labora- study the swimming habits, tropisms, and tory, Gothic, Gunnison County, Colorado. The longevity (Holliman, 1961). Unless otherwise only other report on cercariae in Colorado is stated, descriptions are based on shed cercariae. that of Hunter and Birkenholz (1961), who For microscopic study, live cercariae were worked on the larval trematodes of Gunnison transferred in a drop of water to a glass slide County. They found a high incidence of cer- and a No. 1 coverslip placed over it. Differ- cariae distributed in six major groups, with ent kinds of media used by various investiga- the echinostome type being predominent. tors were tried, especially those said to facili- The present study was undertaken in order tate the visibility of flame cells, such as in- to acid to the meager knowledge on the larval travitam stains combined with normal saline trematodes of Colorado and gain an insight (Lundahl, 1941) and Amphibian Ringer's into the digenetic trematode fauna of this state. solution (Hall, 1959). With echinostome cercariae, collar spines were studied by the Materials and Methods method of Abdel-Malek (1952). The genital Mollusks were collected between the springs anlage was best studied from stained speci- of 1962-1964 from rivers, lakes, creeks, ponds, mens. Cercariae were studied alive, both un- and irrigation canals which abound in north- stained and stained with intravitam dyes of ern Colorado. neutral red and nile blue sulfate, which proved In the laboratory, water from the location very helpful in determining the number of penetration glands. Details of penetration 1 Present address: Department of Biology, Southern University, Baton Rouge, Louisiana. glands and the excretory system were obtained

Copyright © 2011, The Helminthological Society of Washington OF WASHINGTON, VOLUME 35, NUMBER 2, JULY 1968 261 by quickly compressing living cercariae with Results a coverslip (Bums, 1961). Cercarial reaction A total of 2,189 mollusks was examined to intravitam dyes was sometimes checked by over the 2-year period. Of these, 150 were the method used by Holliman (1961). pelecypods (Pisidium Pfeiffer), and the re- Coverslip pressure was controlled by the mainder gastropods. None of the pelecypods addition of water to its edge or by absorption were infected. A total of 778 snails, represent- of water from under it with blotting paper. ing 37.2% of the 2,039 examined, were in- The evaporation of mounting solution was pre- fected with larval trematodes; 116, or 14.9% vented (or controlled) by sealing the margins of the infected snails, shed cercariae which of the coverslip with vaseline. were identified. Different methods were used to slow down The mollusks were infected with four major the activity of living cercariae under a cover- groups of cercariae, viz., echinostome, furo- slip for study. The use of intravitam stains cercous, monostome, and xiphidiocercaria (see produced some effect. Coverslip pressure was Table I), including apparently 26 species. helpful as well as the application of a drop Nine species were studied in detail, with four of chloretone solution at the concentration of described as new. 1 to 10 parts of water. Cercariae were fixed An examination of the data on molluscan by adding 10% formalin of equal volume to infection with larval trematodes studied in this the water in a stender dish containing cer- survey brings out some interesting aspects of cariae, a method similar to that used by Talbot this work. Of all the cercarial species studied, (1936) and Holliman (1961). Sporocysts and that of Notocotylus urbanensis (Cort, 1914) rediae were fixed in the same manner. appeared to be distributed more widely than For toto mounts, preserved materials were the others in terms of locality. Table II shows stained with either Mayer's acid carmine or the percentage of cercarial infections for each Harris' alum haematoxylin. Fast green was host species from northern Colorado. sometimes used as a counter stain, and proved to be especially good for sporocysts and rediae. Description of new species and statements All the drawings of cercariae were made to concerning identified ones scale from preserved material with the aid A. Xiphidiocercaria of a projection prism. Measurements, in 1. Cercaria alganeshi n. sp. microns, of preserved material, unless other- (Plate I, Figs. 1-5) wise specified, represent the average for five to ten well-extended or fairly well-extended Diagnosis specimens. The minima and maxima are re- Belongs to Armatae group of Xiphidiocer- corded in the descriptions, with the average caria. Body 198-231 (209) long, 67-110 (87) being in parentheses. Cercariae obtained from wide at level immediately preacetabular. Tail crushed snails were identified only to the ma- 168-209 (185) long, 19-24 (22) wide at base. jor groups to which they belong, and were Oral sucker 43-48 (47) in diameter; mouth designated with letters for the ease of differen- subterminal; pharynx 14—17 (16) in diam- tiation. eter; esophagus longer than prepharynx, bi-

Abbreviations for all figures: bp, birth pore; eg, cystogenous gland; cp, caudal pocket; cs, collar spine; e sp, eyespot; es, esophagus; ev, excretory vesicle; fc, flame cell; g, gut; gd, gland duct; gp, genital primordium or genital anlage; ic, intestinal cecum; la, locomotor appendage; os, oral sucker; p ph, prepharynx; pg, penetration gland; ph, pharynx; s, spine; st, stylet; t, tail; vs, ventral sucker or acetabulum. Explanation of Plate I Figures 1-5. 1. Cercaria alganeshi n. sp. Ventral view showing body and tail structures. 2. Stylet of same, ventral view. 3. Side view of cercaria showing position of genital anlage and stylet. 4. Sporocyst. 5. Tail of cercaria shed under coverslip pressure showing crenate appearance and rounded mass at its base.

Table 1. List of cercariae studied.

Cercarial type Host Locality I. Echinostome 1. C. coloradensis n. sp. Physa gyrina Ft. Collins 2. C. denverensis n. sp. Physa integra Denver 3. Echinostome cercaria A Physa gyrinn Bellvue 4. „ „ B Psysa sp. 1 mi S of Bellvue 5. ,. i. C Psysa sp. Ft. Collins 6. „ u D Psysa sp. Ft. Collins 7. „ u E Psysa sp. Ft. Collins 8. u u F Psysa sp. Buckhorn Mt. 9. C. olseni n. sp. Lymnaca palustris Chambers Lake area, 42 mi W of Ft. Collins II. Furcocercous 1. C. of Cotylurus flabclliformis Lymnaea auricularia Dixon Lake, near Ft. Collins 2. Furcocercous cercaria A Physa sp. Ft. Collins longifurcous pharyngeate monostome Gyrauhis sp. Ft. Collins III. Monostome 1. C. of Notocotyltis stagnicolac Lymnaea auricularia Dixon Lake, near Ft. Collins 2. C. of N, urbanensis Phijsa gyrina Ft. Collins, Laporte 3. Monostome cercaria A Physa gyrina Ft. Collins IV. Xipliidiocercaria 1. C. alganeshi n. sp. Physa gyrina Ft. Collins 2. C. of Plagiorchis micracanthos Physa gyrina Laporte 3. C. of P. vespertilionis parochis Lymnaea auricularia Dixon Lake, near Ft. Collins 4. Xiphidiocercaria A Physa gyrina Bellvue 5. M B Helisoma trivolvis Dixon Lake, near Ft. Collins 6. M C Hclisoma trivolvis Dixon Dam, near Ft. Collins 7. M D Lymnaea palustris Ft. Collins 8. u E Physa sp. Ft. Collins 9. n F Stagnicola caperata Ft. Collins 10. n G Physa gyrina Ft. Collins 11. n H Stagnicola caperata Greeley 12. n I Lymnaea palustris Chambers Lake area, 42 mi W of Ft. Collins furcates about midway between pharynx and Tail inserted in caudal pocket; circular muscles acetabulum; ceca extending almost to pos- more prominent than in body, giving it striated terior end of body. Stylet (Fig. 2) elongate, appearance; filled with 30-35 small central sharply pointed and wider at base; 22 long cells; when shed (in specimens under cover- and 5 wide; with basal bulb. Acetabulum 41- slip pressure) a rounded mass seen at base 48 (46) in diameter, subequatorial in posi- (see Plate I, Fig. 5). Genital rudiment rep- tion. Tegument with minute spines concen- resented by large mass of small cells extend- trated anteriorly and sparse at posterior part ing from anterior to posterior level of acetab- of body, tail smooth. Penetration glands six ulum presenting G-like appearance. Develop- pairs; located anterolateral to acetabulum. Ex- ment in sporocysts bearing constrictions; sporo- cretory bladder consists of cornua located close cysts 627-814 (759) long, 176-220 (194) to posterior margin of ventral sucker and pos- wide, containing one or two immature em- terior circular vesicle composed of a single bryos, suggestive that cercariae emerge while layer of large cells. With alternate filling and yet immature and complete development in emptying of anterior arms (cornua) and pos- digestive gland of snail host. terior vesicle, excretory bladder sometimes HOST: Physa gyrina Say. assumes Y-shape. Excretory tubules and flame INCIDENCE OF INFECTION: One of 45 snails cells obscured by numerous cystogenous shed cercariae. glands. Body with scattered refractile con- LOCALITY: Pond at tree dump, Fort Collins, cretions. Spines present in caudal pocket. Larimer County, Colorado.

Table 2. Percentage of cercarial infection for each species of molluscan host.

Infected with any larval form Positive with eercaria Host Collected Number Percentage Number Percentage

Gyratilus sp. 26 6 23.1 4 15.4 Holisoma trivolvis 94 4 4.26 3 3.9 Lymnaca aiiricularia 668 595 89.1 28 4.18 Lijmnaea palnstris 60 8 13.3 5 8.33 Lymnaea sp. 46 0 0 0 0 Physa gyrina 546 99 18.1 48 8.80 Physa Integra 46 9 19.6 3 6.52 Physa sp. 453 34 7.51 12 2.65 Pisiditim sp. 150 0 0 0 0 Stagnicola caperata 100 23 23.0 13 12.0

Total 2,189 778 35.5% 116 14.9%

Cercariae emerge from their snail host a few (mucilagenous plug), caudal pocket with at a time, during the morning hours up to spines, and intestinal ceca extending to almost noon. Their longevity is about 10 hours. Neu- the posterior end of the body. Cercaria of tral red stain is mildly toxic, while nile blue T. robustus differs from C. alganeshi by its sulfate is toxic, killing the cercariae in short possession of nine penetration glands (four time. The cercariae swim with the body on one side and five on the other), less pro- flexed ventrally and the tail lashing in a nounced genital anlage, and shorter esophagus figure-eight motion. They swim intermittently, bifurcating farther preacetabular. C. alganeshi alternating this with creeping on the substrate. is similar to C. concavocorpa with respect to Swimming is mostly vertical. From the sub- the decaudation process of specimens under strate, they swim up until they reach the top coverslip pressure, leaving the base of the tail of the water in the container. Here they stop with an accumulation of a rounded mass of wriggling and start sinking to the bottom of tissue (Fig. 5) and the presence of a basal the container with the body half flexed and bulb on the stylet. The presence of eight pairs the tail bent forward over it. The rate of of penetration glands in C. concavocorpa, in- sinking may be slow or comparatively fast. testinal ceca extending almost to the level of The tail is longer than the body when swim- posterior part of acetabulum, and spines cover- ming but is contracted to about half the length ing the body and anterior half of the tail serve of the body when creeping. When subjected to separate it from C. alganeshi. to coverslip pressure, the body contracts, the anterior region becoming much wider with a 2. Cercaria o£ Plagiorchis micracanthos consequent reduction of the posterior region McMullen, 1937 and the tail. Five of 20 Physa gyrina Say, collected from C. alganeshi resembles C. goodmani Najar- a pond opposite Laporte Lions Park, Laporte, ian, 1952, the cercaria of Telorchis robustus were parasitized by xiphicliocercariae identi- Krull, 1936, and C. concavocorpa Sizemore, fied as those of Plagiorchis micracanthos. The 1936. It resembles C. goodmani by the nature general features of body and dimensions as and operation of the excretory bladder and reported by McMullen (1937) agree so closely possession of caudal pocket spines. C. good- with mine that I do not hesitate considering mani can be distinguished from C. alganeshi the two identical. This is a new host and new by its stylet which is devoid of a basal bulb, locality record. its short intestinal ceca, possession of eight pairs of penetration glands and tegumental 3. Cercaria of Plagiorchis vespertilionis spines which cover the entire body and tail. parorchis Macy, 1960 C. alganeshi appears to be closely related to This species was found infecting one of 131 the cercaria of T. robustus in the two having Lymnaea auriciilaria (L.) collected from Dixon stylets wider at the base and with a basal bulb Lake, near Fort Collins, The specimen agrees

Explanation of Plate II Figures 6-11. 6. Cercaria denvercnsis n. sp. Ventral view showing details of body and tail, 7. Cephalic region from ventral view showing collar spines. 8. Redia. 9. Metacercaria. 10. Lateral view showing occasional posture on substrate. 11. Lateral view showing a different posture on substrate.

Copyright © 2011, The Helminthological Society of Washington OF WASHINGTON, VOLUME 35, NUMBER 2, JULY 1968 265 with the description given by Macy (1960). Development in colorless rediae (Fig. 8), A new host and new locality record is estab- spotted with yellowish-brown pigment, latter lished for this cercaria. less numerous in anterior one-fifth. Redia B. Echinostome cercaria 682-1,650 (1,265) long, 132-264 (205) wide; 1. Cercaria denverensis n. sp. with only posterior locomotor appendages (Plate II, Figs. 6-11) which tend to be less pronounced in mature redia. Appendages located at about three- Diagnosis fifths distance from anterior end. Pharynx Body 187-253 (238) long, 83-110 (98) 41-62 (48) long, 41-53 (45) wide, leading wide at level of acetabulum; collar 62-84 (73) to short gut. Birth pore lateral and just pos- with about 34 spines in single row; tail 130- terior to collar. Tegument thin. Usually 14- 220 (155) long, 43-46 (44) wide at base. 16 cercariae in mature redia. Oral sucker 35-41 (37) in diameter, mouth Metacercaria (Fig. 9) found in the same subterminal, prepharynx 10—12 (11) long, snail as cercariae. Some cercariae encysted in pharynx 14-22 (18) long, esophagus 14-22 vials containing snails. Cysts spherical, 101— (16) long, bifurcating just preacetabular, ceca 113 (109) in diameter, with thick, irregular extending to near posterior margin of body. outer layer and thin inner membrane. Meta- Acetabulum 43-58 (46) in diameter, with cercaria has more concretions in primary ex- small superficial and spherical concretions. cretory tubules than cercaria, primary tubules Body devoid of spines; with row of about 17 arranged in a loop and crossing each other. large and conspicuous cystogenous glands ex- HOST: Physa integra Hald. tending along each lateral margin from phar- INCIDENCE OF INFECTION: One of 46 snails ynx to posterior end of body, those located shed cercariae. posteriorly being larger; smaller glands fill LOCALITY: Cherry Creek, Denver, Colorado. enclosed area in random arrangement. Tail This larva emerges from its snail host during devoid of spines; crenate along margin; with the daytime, between 9:00 AM and 4:00 PM. numerous nuclei of parenchyma, visible in Only a few are shed at a time. The life span stained material. Cercariae decaudate readily. of the cercaria is about 12-14 hours if encyst- Excretory bladder small, and oval; from its ment does not occur. No phototropism was anterior part two primary collecting tubules observed. Neutral red stain used for the study extend in wavy manner to posterolateral mar- of live cercariae was mildly toxic. The cer- gins of acetabulum enlarge and undergo three caria is not an active swimmer. Swimming is or four undulations, then taper as they extend accomplished by the vigorous wriggling of the to the level of prepharynx where each makes tail with the body flexed. However, it mostly an almost deltoid loop, passing under itself stays at the bottom of the container with the and descending to the level of excretory vesicle anterior part bent up or down, but most com- as the secondary tubule, loops again passing monly up (Figs. 10 and 11). In either of under itself as the tertiary duct and is seen these positions, movement is from side to side as far anterior as level of ventral sucker. Pri- accompanied by a slight wriggling of the tail. mary collecting tubules filled with concretions, An attempt to infect leghorn chicks with meta- extending from ventral sucker anteriorly to the cercariae was unsuccessful. midlevel of esophagus. Tuft of cilia in the Cercaria denverensis bears some resem- secondary tubule from level of anterior part blance to the cercaria of Echinopanjphium of acetabulum to excretory vesicle, beat rapidly flexum as described by Najarian (1954) with in living specimens flattened under coverslip respect to its digestive system, genital anlage, pressure. Excretory tubule in tail forks at about and excretory system. These two species, how- posterior part of first fourth of length of tail ever, differ in the following respects: C. into branches which open laterally to outside. denverensis is smaller in size; it has 34 collar About 16 flame cells observed on each side of spines, while cercariae of E. flexum have 45; body. Reproductive primordium consists of the flame cells are 32 (16 in each side), but two irregular anterior and posterior masses of 40 for the cercariae of E. flexum; the bifurca- cells connected by thin strand of cells. The tion of the caudal excretory tubule is about posterior mass shows vitelline duct rudiment. one-fourth of the length of the tail from its

Explanation of Plate III Figures 12-14. 12. Cercaria coloradcmis n. sp. 13. Redia. 14. Lateral view of cercaria showing occasional posture 011 substrate.

Copyright © 2011, The Helminthological Society of Washington OF WASHINGTON, VOLUME 35, NUMBER 2, JULY 1968 base, but one-eighth for the cercariae of E. INCIDENCE OF INFECTION: Ten of 47 snails flexum; it is devoid of tegumental spines which shed cercariae. are present in the cercariae of E. flexum. The LOCALITY: Pond on highway No. 14 in Fort arrangement of cystogenous glands in conspic- Collins, Larimer County, 1 mile NE of Colo- uous lateral row on each side is not mentioned rado State University. for cercariae of E. flexum. Cercariae are shed during the day. Their swimming habit is characteristic of echino- 2. Cercaria coloradensis n. sp. stome cercariae, being with body flexed and (Plate III, Figs. 12-14) tail expanded and lashing vigorously. The cercaria folds over itself intermittently with Diagnosis the ventral sucker protruding like a short Body 242-286 (257) long, 88-138 (108) tube (Fig. 14) after creeping for a while on wide at level of acetabulum. Tail 297-385 the substrate. Neutral red is mildly toxic. (341) long, 26-36 (32) wide; mouth sub- Cercaria coloradensis is similar to C. reb- terminal; prepharynx 7-17 (12) long; pharynx stocki McCoy, 1929, in some respects. While 14-19 (16) long, 11-14 (12) wide; esoph- the esophagus of C. coloradensis bifurcates just agus bifurcating just below anterior margin of posterior to the anterior margin of the acetab- acetabulum; ceca end near posterior margin ulum that of C. rebstocki bifurcates anterior of body. Acetabulum 43-60 (50) long, 46- to the acetabulum. It has a total of seven 65 (52) wide; located just posterior to mid- gland ducts, while C. rebstocki has eight. By body. Body and tail devoid of tegumental possessing two eyespots which C. rebstocki spines. Collar 74-84 (78) wide with spines lacks, C. coloradensis resembles a marine cer- of undetermined number obscured by numer- caria, C. fuscata Holliman, 1961. ous cystogenous glands. Seven gland ducts observed dorsal to oral sucker; two spherical 3. Cercaria olseni n. sp. ducts laterally located, five elongate ducts (Plate IV, Figs. 15-17) medially located (Fig. 12). Excretory vesicle This species is named after Dr. O. Wilford bulbous; from anterior part two primary col- Olsen, who helped in working out the flame lecting tubules extend in wavy form to lateral cell pattern of this cercaria. margins of acetabulum, dilate and undergo three or four undulations, then taper as they Diagnosis extend to level of prepharynx, here each forms Body 170-215 (189) long, 83-99 (90) wide an almost triangular anastomosis; rest of at level just above acetabulum. Tail 187-242 tubule obscured by cystogenous glands; pri- (223) long, 19-22 (20) wide at base. Oral mary collecting tubules filled with numer- sucker 29-31 (30) long, 26-31 (29) wide; ous concretions which extend from ventral mouth subterminal; prepharynx 5-8 (6) long; sucker to level of pharynx. No bifurcation of esophagus long, bifurcating just preacetabular; caudal excretory tubule observed. Two eye- ceca extending to level of excretory vesicle; spots lateral to prepharynx. Tail with numer- esophagus and ceca filled with single row of ous nuclei of parenchyma (seen in stained spherical bodies with posterior part of each material), especially crowded in posterior tip. overlapping anterior part of adjacent one. Genital primordium represented by two un- Acetabulum 35-43 (39) long, 36-43 (40) connected anterior and posterior masses of wide; located subequatorial. Body and tail small cells. Development in fairly large recliae, devoid of spines. Collar 53-60 (58) wide, measure 935-1,925 (1,247) long, 209-352 with about 32-36 spines in single row (Fig. (271) wide just above posterior locomotor ap- 16). Excretory vesicle comparatively small, pendage. Pharynx 38-86 (54) in diameter. oval. From anterior part of vesicle two pri- Birth pore lateral, just posterior to short collar; mary collecting tubules extend to level of tegument comparatively thick; locomotor ap- prepharynx in wavy manner; here each makes pendages not very prominent in either young circular loop and continues posteriorly as sec- or mature recliae; gut short. About 16-20 ondary tubules; latter obscured by cystogenous cercariae in various stages of development. glands filling body; from acetabular level each HOST: Physa gyrina Say. primary duct increases in diameter and under-

Explanation of Plate IV Figures 15-17. 15. Cercaria olseni n. sp. Ventral view showing details of body and tail structures (cystogenous glands omitted). 16. Cephalic region of same, ventral view showing collar spines. 17. Redia.

Copyright © 2011, The Helminthological Society of Washington OF WASHINGTON, VOLUME 35, NUMBER 2, JULY 1968 269 goes three or four undulations, then tapers to excretory duct at about one-fifth of the length level of prepharynx. Primary tubule with of the tail from the base. spherical concretions extending from the ventral sucker to level of pharynx. Caudal excre- Discussion tory duct bifurcates at about one-fourth length At the initial stage of this investigation snail of tail into branches opening laterally to out- collections were maintained in 50 cc wide- side. About 16 flame cells observed on each mouth bottles filled three-quarters full with side of body. Tail crenate at anterior two- tap water and closed with regular bottle fifths; stained specimen with numerous nuclei covers. The frequently observed outcome of in parenchyma. Genital primordium repre- this method was the death of all the snails sented by a mass of small cells lying posterior within 2 or 3 days. Later, the writer adopted to acetabulum and a smaller mass lying aiitero- a method which involved putting the snails in dorsal and close to the latter. Development filtered water, obtained from the respective in elongate redia with prominent posterior bodies of water where collections were made locomotor appendages seen in both young and covering the bottles with cheesecloth. and mature specimens. Redia (Fig. 17), This was found very advantageous. It in- 1,760-2,354 (2,110) long, 154-209 (183) creased to a considerable extent the longevity wide at level just above locomotor appendages. of the snails. Many uninfected and slightly Birth pore not observed; tegument thin. Four- infected snails lived up to the set limit of 14 teen to 18 embryos per mature redia in various days, after which they were crushed for exam- stages of development. ination. Some snails which shed cercariae lived HOST: Lymnaea palustris (Mull.). up to 5 or 6 days. This better result could INCIDENCE OF INFECTION: One of three be attributed to the following reasons: (1) snails shed cercariae. Water, filtered with cheesecloth in order to LOCALITY: Pond near Chambers Lake, 42 remove large particles of debris, still contains miles west of Fort Collins, Larimer County, some plankton and food particles on which Colorado. the snails may feed. (2) By maintaining snails Neutral red was mildly toxic to cercariae. in the water from which they are collected, The swimming habit is characteristic of echino- they are left in a condition close to their stome cercariae, being with body flexed and natural habitat. (3) This water, usually with tail extended to about two times body length dissolved salts (e.g. calcium carbonate) which and lashing vigorously in a figure-eight mo- the snails require for shell construction, is more tion. This tail movement helps to propel the propitious than tap water for their mainte- cercaria rapidly. Attempts to infect leghorn nance. (4) Covering the bottles with cheese- chicks with metacercariae were unsuccessful. cloth not only prevents the snails from crawl- Cercaria olseni is similar in some respects ing out, but allows more aeration than the to Cercaria equispinosa Brown, 1926, and regular bottle covers, thus increasing the dis- C. indicae XXIII Sewell, 1922, but differs solved oxygen content of the water which may from them by its smaller size, arrangement be a limiting factor for most snails; some snail of collar spines, number of flame cells, genital species require a high concentration of dis- rudiment and bifurcation of the caudal ex- solved oxygen. (5) Changing of the water cretory duct. The digestive and excretory sys- every other day prevents "souring" which is tems of C. olseni and C. equispinosa have often caused by the defecation of the snails. striking resemblances. The genital anlage of It is thus advisable for investigations of this C. olseni is dissimilar to that of C. equispinosa, nature to maintain the snails in the water being represented by only a mass of cells post- from which they are collected and change it acetabularly. While the number of flame cells regularly. in C. olseni is about 16 pairs, they are said to Of the nine species of shed cercariae studied be several (4) 011 each side of the posterior in detail, four are described as new ones, region of the body of C. equispinosa. C. augmenting the list of United States cercariae indicae XXIII differs from C. olseni by pos- awaiting life history study. The similarities session of a double ring of collar spines, 18 and differences of these species to their most pairs of flame cells, bifurcation of the tail closely related forms are given. Although

Copyright © 2011, The Helminthological Society of Washington 270 PROCEEDINGS OF THE HELMINTHOLOGICAL SOCIETY three of the new species are echinostomes and the parasite has undoubtedly led to the de- the other a xiphidiocercaria, the overall result scription of cercariae that would look much of the study indicates that xiphidiocercariae different if they had been allowed to emerge are more abundant than those of the other naturally." While the practice of describing groups represented here. This observation is cercariae obtained from crushed snails is dis- not in agreement with that of Hunter and couraged, it is advisable to crack and examine Birkenholz (1961), who reported a greater for infection, snails which did not shed cer- abundance of echinostome cercariae. The dif- cariae instead of discarding them, as is com- ference (3) in the number of species of monly done by some investigators. This has xiphidiocercaria and echinostome cercariae is, the advantage of giving an insight into the however, too small to be considered very trematode fauna of the locality or region being significant. surveyed. Moreover, there have been reported The percentage of infection (35.5%) indi- cases of precocious development of metacer- cated in Table II apparently shows that the cariae inside the sporocyst or redia without the trematocle fauna in northern Colorado is rather cercarial stage emerging from the snail (Stun- meager, as might be expected in a region of kard, 1934; Ingles, 1935; Cort and Brackett, limited precipitation (Gittings 1941). Table 1937; McMullen, 1937, 1938; and Acholonu, II shows the interesting fact that the snail 1965). Cases of this nature will escape the Lijmnaea auricularia (L.) had the highest notice of an investigator who discards snails percentage of infection (89.1%). It is a Euro- that did not shed cercariae. It is advisable, pean species previously introduced into the therefore, to crush and examine such snails. eastern part of the United States and now found in Colorado. Summary A considerable decrease in larval trematocle Cercariae falling into four main groups and infection of snails was noted during the late 26 apparent species, including four uncle- fall and winter seasons. Some of the snails scribed species, were collected; nine species collected during these periods were infected were shed by the snail hosts; the remainder with larval annelids (Oligochaeta). Such snails were obtained from crushed snails, and were had no larval trematode infections. Snails, identified only to major group. They include especially Lijmnaea auricularia (L), harbored Cercaria denverensis n. sp. from Physa Integra mostly the metacercarial stage during the Hald., C. coloradensis n. sp. from P. gyrina winter. However, from the latter part of Say, and C. olseni n. sp. from Lijmnaea palus- March to the summer months there was a tris (Mull.) of tlie echinostome group; cercaria significant increase in the other larval stages of Cotyhmis flabelliformis (Faust, 1917) from in the snails. This observation is in agreement L. auricularia (L.) and one unidentified spe- with that of Faust (1922) and Ulmer (1957) cies of the furcocercous group; cercaria of with respect to Tetracotyle flabelliformis Notocotylus stagnicolae Herber, 1942, from L. (Faust, 1917). auricularia (L.) and cercaria of N. urbanensi.s The inadvisability of crushing infected (Cort, 1914) from P. gyrina Say of the mono- snails to obtain larval trematodes for study stome group; cercaria of Plagiorchis micra- has been emphasized by various investigators. canthos McMullen, 1937, from L. auricularia Stunkard (1930) said: "When infected snails (L.), C. alganeshi n. sp. from P. gyrina Say and are crushed, the larval trematodes are freed, eight unidentified species of the xiphidio- but the cercariae are usually immature and cercaria group, the most prevalent. frequently differ, even in morphological fea- The 2,189 mollusks collected represent six tures, from those which emerge normally. recoznized species and several identified only Many published descriptions of cercariae are to genus; 35.5% were infected with larval trem- based on the study of such immature speci- atodes of one stage or another; 14.9% of the mens, and consequently the observations, al- infected snails contained cercariae. though accurate for that stage of development are misleading." Dunagan (I960) also pointed Acknowledgments out that "the very common practice of cracking A dissertation submitted to the Graduate snails to obtain and study different stages of School, Colorado State University, in partial

Copyright © 2011, The Helminthological Society of Washington OF WASHINGTON, VOLUME 35, NUMBER 2, JULY 1968 271 fulfillment of the requirements for the degree Mosesia chordeilesia McMullen, 1936. (Trem- of Doctor of Philosophy. atoda: Lecithodendriidae). J. Parasit. 45: The author wishes to express sincere grati- 327-336. tude to Professor O. Wilford Olsen under Holliman, R. B. 1961. Larval trematocles from whose supervision this research was conducted, the Apalacbee Bay area, Florida, with a check- list of known marine cercariae arranged in a to Mrs. Dorothy E. Beetle, malacologist, key to their superfamilies. Tulane Stud. Zool. Charlotte Children's Nature Museum, Char- 9: 3-74. lotte, North Carolina, who helped in identify- Hunter, G. W., Ill, and D. H. Birkenholz. ing some of the snail species, and Dr. Hugo 1961. Notes on larval trematodes of Gunni- Rodek, Director, University of Colorado Mu- son County, Colorado. Tr. Am. Micr. Soc. 80: seum, who allowed the author the use of some 358-364. museum specimens of the University's mala- Hurst, C. T. 1923. Cercaria gunnisoni, a new cological collections for identification purposes. xiphidiocercaria from Colorado. Bull. West- Due appreciation is also extended to Dr. Nell ern State Coll. Colo. 12: 1-15. B. Causey, Zoology and Physiology Depart- Ingles, L. G. 1935. Notes on the development of a heterophyid trematode. Tr. Am. Micr. ment, Louisiana State University, for reading Soc. 54: 19-21. the manuscript and making helpful suggestions. Lundahl, W. S. 1941. Life history of Caecin- This investigation was supported by a fel- cola parvulus Marshall and Gilbert (Crypto- lowship from the Federal Republic of Nigeria. gonimidae, Trematoda) and the development of its excretory system. Tr. Am. Micr. Soc. Literature Cited 60: 461-484. Abdel-Malek, B. T. 1952, Cercaria chandleri, McMullen, D. B. 1937. The life histories of a new echinostome species from the snail three trematodes parasitic in birds and mam- H&lisoma corpulentum in Lake Itasca, Minne- mals, belonging to the genus Plagiorchis. J. sota. Tr. Am. Micr. Soc. 71: 277-281. Parasit. 23: 235-243. Acholonu, A. D. 1965. Contributions to the . 1938. Observations on precocious meta- life history of Cotylurus flabelliformis (Faust, cercarial development in the trematode super- 1917) (Trematoda: Strigeidae). Proc. Helm. family Plagiorchioidea. J. Parasit. 24: 273— Soc. Wash. 32: 138-140. 279. Burns, W. C. 1961. Six virgulate xiphidiocer- Macy, R. W. 1960. The life cycle of Plagiorchis cariae from Oregon, including redescriptions vespertilionis parorchis n. sp. (Trematoda: of Allassogonoporus vespertilionis and Acan- Plagiorchiidae), and observations on the ef- thatrium oregonense. J. Parasit. 47: 919-925. fects of light on the emergence of the cercaria. Cort, W. W., and S. Brackett. 1937. Pre- J. Parasit. 46: 337-345. cocious development of the metacercaria stage Najarian, H. H. 1954. Developmental stages of Diplostomiim flexicaudum in the snail in- in the life cycle of Echinopan/phium flexum termediate host. J. Parasit. 23: 545-546. (Linton, 1892) Dietz, 1910. (Trematoda: Crawford, W. W. 1939. Studies on the life Echinostomatidae). J. Morph. 94: 165-197. cycle of Colorado trematocles. J. Parasit. 25 Stunkard, H. W. 1930. An analysis of the (6 Suppl.): 26. methods used in the study of larval trema- . 1940. The life history of a gorgoderid todes. Parasitology 22: 268-273. trematode, presumably of the genus Phytto- . 1934. The life history of Typhlocoeluin distomum. J. Parasit. 26 (6 Suppl.): 38. cymbium (Dies., 1850) Kossack, 1911 . 1943. Colorado trematode studies. I. (Trematoda: Cyclocoelidae). A contribution A further contribution to the life history of to the phylogeny of the monostomes. Bull. Crepidostomum farionis (Miiller). J. Parasit. Soc. Zool. France 59: 447-466. 29: 379-384. Talbot, S. B. 1936. Studies on schistosome der- Dunagan, T. T. I960. Cercariae belonging to matitis 2. Morphological and life history the Opisthorchioidea. Proc. Helm. Soc. Wash. studies on three dermatitis-producing schisto- 27: 44-52. some cercariae, C. elvae Miller, C. stagnicolae Faust, E. C. 1922. Notes on larval flukes from n. sp. and C. physettae n. sp. Am. J. Hyg. 23: China. Parasitology 14: 248-267. 372-384. Gittings, E. B. 1941. Climate of Colorado. In Ulmer, M. J. 1957. Notes on the development "Climate and Man." U. S. Dept. Agric. Yr. of Cotylurus flabelliformis tetracotyles in the Bk., pp. 789-808. second intermediate host (Trematoda: Strige- Hall, J. E. 1959. Studies on life history of idae). Tr. Am. Micr. Soc. 76: 321-327.

MINUTES Four Hundred Twenty-ninth Through Four Hundred Thirty-sixth Meetings

429th Meeting: Adult Education Center, 431st Meeting: Plant Industry Station, Belts- University of Maryland, College Park, Mary- ville, Maryland, 18 December 1967. Formal- land, 11 October 1967. Presentation of Hel- ities of installation of new officers completed. minthological Society Anniversary Award to Papers presented: "A nematode parasite of Dr. Gerard Dikmans (in absentia). An- the face fly of cattle and horses," by W. R. nouncement of Eighth International Congress Nickle; "Reproduction in Diploscapter coro- of Tropical Medicine to be held in September nata," by H. C. Hechler; "Plant growth regu- 1968. Slate of officers for 1968 presented: lators and resistance to root knot nematodes," D. R. Lincicome (President), A. C. Pipkin by V. Dropkin; and a voluntary paper from (Vice-President), E. J. L. Soulsby (Recording the floor: "Metabolic differences in isolates Secretary) and E. M. Buhrer (Corresponding of T. lewisi from different parts of the world," Secretary-Treasurer). Papers presented: "The by D. R. Lincicome. filaria, Macacanema formosana of the Taiwan 432nd Meeting: National Institutes of monkey, Macaca cyclopis," by J. F. Bergner; Health, Bethesda, Maryland, 19 January 1968. "Morphological variation in the rat nematode, Papers presented: "Genetic aspects of the sus- Nippostrongylus brasiliensis," by R. Lichten- ceptibility of Biomphalaria glabrata to infec- fels "Effects of concurrent infections of Nip- tion with Schistosoma mansoni," by D. S. postrongylus brasiliensis and Hymenolepis Richards; "Activity of Lincomycin analogs nana" by J. Poiley; "Blood protozoan para- against Plasmodium cynomolgi in Rhesus mon- sites of the redwing blackbird and the common keys," by K. G. Powers; "Studies on antigens grackle: A survey of two populations of Kent from axenically ciiltured Entamoeba histoly- Island, Maryland," by R. G. Bagley; "Studies tica," by M. N. Lunde and L. S. Diamond; on the biology and life history of Hepatozoon "Studies on parasitic gastroenteritis of cattle grisesciuri, a haemogregarine of the gray squir- in Germany," by J. Eckert and H. J. Burger, rel," by B. Reclington; "Nippostrongylus bra- "Infection of Meriones unguiculatus, M. hur- siliensis in castrates and normal male rats," by rianae, Mesocricetus auratus, and Cricetuhts R. Salerno. griseus with Dtpetalonema ivitei" by G. 430th Meeting: Beltsville Parasitological Pacheco. Laboratory, Beltsville, Maryland, 17 Novem- 433rd Meeting: Patuxent Wildlife Research ber 1967. Letter of thanks from Dr. Dikmans Center, Laurel, Maryland, 9 February 1968. read. Illness of senior member, Dr. J. R. Report on the progress of the Program Com- Christie, announced. W. R. Nickle, J. H. mittee of the Second International Congress of Fischthal, and J. T. Lucker elected to Editorial Parasitology given. Papers presented: "Mech- Board. Slate of officers presented at 429th anism of red cell destruction in Leucocytozoon meeting elected by unanimous vote. Papers simondi infections," by R. M. Kocan; "Breed- presented: "Continuous flow flotation proc- ing sites and collections of immature stages of ess for separating oocysts from fecal debris," black flies in Michigan (Diptera: Simuliidae)," by J. M. Vetterling; "Host-parasite relations: by I. B. Tarshis; "Epizootiology of Plasmodium Infection of cattle with Oesophagostomiim co- circumflexum in birds: Host and geographical lumbianum, a parasite of sheep," by H. Her- distribution," by C. M. Herman and E. Snyder; lich; "The development of a system of classifi- "Comparative mitotic activity in two species cation of the Nematoda," by M. Chitwood; of slime mold-like Ameboid Protozoa," by T. "The esophago-intestinal sphincter of Rictularia K. Sawyer. lucifigiis (Nematoda: Thelaziidae)," by F. 434th Meeting: Naval Medical Research Douvres; "Time-lapse photographs of equine Institute, Bethesda, Maryland, 22 March 1968. Babesias," by W. M. Frerichs. The Auditors' report was read and approved.

Report on the progress of the organization of cuniculi in the rabbit," by T. J. Hayes; "Ascaris the Second International Congress of the lumbricoides in the unnatural host," by S. R. World Federation of Parasitologists was given. Sylk; "A parasitological fairy tale," by G. L. Papers presented: "Oxygen uptake by Try- Graham; "Storage of Ascaris larvae at low tem- panosoma lewisi grown in young and old rats," peratures," by D. Post; "The ultrastructure of by J. K. Adaramola; "Oxygen uptake by cells reacting with antibody-sensitized Ascaris Trypanosoma lewisi grown in albino and black larvae," by D. J. Morseth; "Lymphocyte trans- rats," by M. S. El Helu; "Electron-microscopic formation in helminth infection," by E. J. L. study of kinetoplastic DNA from Trypanosoma Soulsby. Cocktails were served in Allam cruzi," by G. Riou; "A look at tropical med- House, courtesy of the School of Veterinary icine in Central America and Mexico," by B. Medicine, University of Pennsylvania, followed Fried. by dinner enjoyed by members and guests. 435th Meeting: Naval Medical Research The following were elected to membership Institute, Bethesda, Maryland, 17 April 1968. at the meetings indicated: 429th: D. Gershon, Summary of business executed by Executive C. L. Heagle, P. Krupa, F. Lamberti, D. H. Committee given to members. Papers pre- MacDonald, R. D. Romanowski, K. S. Todd, sented: "Drug studies on the exoerythrocytic F. G. Whittacker, G. W. Yeates. 430th: J. C. stages of malaria," by R. L. Beaudoin; "Ex- Burke, B. L. Duncan, R. J. Gee, E. Panitz, perimental infections of malaria in a natural D. Quigley. 433rd: J. K. Adaramola, M. El population of sparrows," by J. E. Applegate; Helu, J. Parris, D. K. Sen, W. B. Willers. "The sporogonous cycle of P. vivax as a system 434th: E. G. Bergquist, L. R. Crawley, S. for evaluation of anti-malarial drugs," by L. A. Draggan, S. J. Edwards, B. Fried, J. S. Keithly, Terzian; "Culture of Trypanosoma rangeli in H. C. King, R. M. Kocan, C. M. Lee, R. D. an insect cell culture system," by A. C. Pipkin; McAnnally, W. E. Martin, E. O. Morrison, "Histology of cercariae of Schistosoma man- B. J. Muirhead, C. A. Porter. 435th: K. Daib, soni: A demonstration," by M. A. Stirewalt. G. Germani, H. H. Kreis, G. Merny, C. 436th Meeting: University of Pennsylania's Netscher, G. A. Wells, D. Williams. New Bolton Center, Kennett Square, Penn- sylvania, 25 May 1968. Papers presented: E. J. L. SOULSBY "The early histotropic stages of Obeliscoides Recording Secretary

INDEX TO VOLUME 35

ACHOLONU, A. D. Studies on the freshwater cercariae of northern Colorado 259 Actinolaimiis armatus, new nematode from Indian soil 96 ALLEN, M. W., and E. M. NOFFSINGER. Revision of the genus Anaplectus (Nema- toda: Plectidae) 77 Allonematobothrium ghanensis, new digenetic trematode from Ghana 237 ALLRED, K. R. (see Tseng) 57 Anaplectus (Nematoda: Plectidae), revision of 77 Anaplectus arenicola as synonym of A. granulosus 77 Anaplectus granulosus, designation of male as neotype 77 Anaplectus rnagnus, redescription of 77 Anaplectus porosus, new nematode from Indian soil 77 Anaplectus similis, new nematode from Wisconsin soil 77 Anaplectus submersus, emended description of 77 Anaplectus varicaudatus, new nematode from Florida soil 77 Anniversary award, G. Dikmans 115 Anthelmintic action of thiabendazole on Ascaridia columbae 117 Ascaridia columbae, effect of thiabendazole on 117 Ascocotyle chandleri, ultrastructure of metacercarial cyst 212 Axonchium asacculum, new nematode from soil in India 248 BARKER, K. R. (see Webber) 34 BECKLUND, W. W. (see Walker) 49 Belondira murtazai, new nematode from soil in India 248 Belondiroiclea, a revised classification of the superfamily 248 BIER, J. W. (see Holloway) 30 Bivesiculoides posterotestis, new digenetic trematode from marine fish in New Caledonia __ 143 Bobcat, helminths of in southeastern United States 118 BOYD, E. M. Two new species of Harpyrhynchus from herons in North America (Acarina, Trombidiformcs, Harpyrhynchidae) 18 Bucephaloides ghanensis, new digenetic trematode from Ghana 237 BUSSING, W. A. (see Price) 54 Capillaria americana, emended description of 206 Capillaria bonnevillei as a synonym of C. americana 206 Carcharolaimus nuisoodi, new nematode from Indian soil 96 Car'charvidimus mujtabai, new nematode from Indian soil 96 Cercaria- alganeshi, C. denverensis, C. coloradensis, and C. olseni, new cercaria from Colorado 259 Cercaria. allomacarapensis, C. guaraunensis, C. heteroglandula, C. marisa, and C. pomacea, new cercaria from Venezuela 67 CLARK, G. M. In Memoriam 37 Clethrionomys gapperi, new host for Capillaria americana 206 Clinostomum hylaranae (metacercaria), new trematode from Ghana 1 COLGLAZIER, M. L. (see Wehr) 117 Cooperia punctata and C. spatidata morphology of 49 CROLL, N. A., and A. R. MAGGENTI. A peripheral nervous system in Nematoda with a discussion of its functional and phylogenetic significance 108 Cultivation of Meloidogyne incognita on Begonia and tomato 106 DASGUPTA, D. R., D. J. RASKI, and S. A. SHER. A revision of the genus Rotylcn- chulus Linforcl and Oliveira, 1940 (Nematoda: Tylenchiclae) 169 Diabrotica u. iindecimpunctata as host for Filipjevimermis leipsandra 161 Didymozoids, description of four new immature forms from Ghana 237

DIKMANS, GERARD, anniversary award 115 Diphtherostomum tropicum, new digenetic trematode from marine fish in New Caledonia 143 Diphyllobothrium sebago, occurrence of plerocercoids in Maine fish 92 Diplodiscus magnus (Trematoda), new description of 1 Diploscapter coronata (Nematoda: Rhabditidae), development and reproduction in 24 Ditylenchus dipsaci, population of in alfalfa field soil 57 Dorylaimellus belondirelloides, D. salvus, and D. paraclavatus, new nematodes from soil in India 248 DURIO, W. O., and H. W. MANTER. Some digenetic trematodes of marine fishes of New Caledonia. Part I. Bucephalidae, Monorchiidae, and some smaller families 143 Echinostoma Undoense (Trematoda), life history 74 Environmental factors, effect of on populations of Ditylenchus dipsaci in soil 57 Esophagus, morphology of in Hexatylus and Neotylenchus 154 Eumasenia ghanensis, new digenetic trematode from Ghana 126 Euparyphium anhingae, new echinostome from Florida birds 197 Fergusobia curriei (Nematoda: Sphaerulariiclae), classification and life history of 40 FISCHTHAL, J. H., and J. D. THOMAS. Digenetic trematodes of amphibians and reptiles from Ghana 1 FISCHTHAL, J. H., and J. D. THOMAS. Digenetic trematodes of some freshwater and marine fishes from Ghana 126 FISCHTHAL, J. H., and J. D. THOMAS. Digenetic trematodes of marine fishes from Ghana: Families Acanthocolpidae, Bucephalidae, Didymozoiclae 237 Filipjevimcrmis leipsandra, parasitic development of in Diabrotica 161 FISHER, J. M., and W. R. NICKLE. On the classification and life history of Fergusobia curriei (Sphaerulariidae: Nematoda) 40 Foxes, helminths of in southeastern United States 118 Ganeo africana (Trematoda), redescription of 1 Gerbils, experimental infection of with Trichostrongylus axei and T. colubriformis 102 GRIFFIN, G. D. (see Tseng) 57 HALEY, A. J. (see Lichtenfels) 206 Halipegus ghanensis, new trematode from Ghana 1 HARKEMA, R. (see Miller) 118 Plarpyrhynchus butorides and H. herodius, new mites from herons in North America 18 Hatching response of Plypsoperine ottersoni to root emanations 34 HEARD, R. W., III. Parasites of the clapper rail, Ralhis longirostris Boddaert. I. The current status of the genus Levinseniella with the description of Levinseniella byrdi n. sp. (Trematoda: Microphallidae) 62 HEARD, R. W., III. Levinseniella huntcri sp. nov., a new species of microphallid trematode from the Wilson's plover, Charadrius wilsonia Ord. 140 HECHLER, H. C. Postembryonic development and reproduction in Diploscapter coronata (Nematoda: Rhabditidae) 24 Helicometm antarcticae, new trematode from Antarctic fish 30 Heterorchis ghanensis, new trematode from Ghana 1 HILL, L. G. (see Whittaker) 15 HOLLOWAY, H. L., JR., and J. W. BIER. Helicometm antarcticae sp. nov. from Antarctic coastal fishes , 30 HOPKINS, S. H. (see Little) 46 Plypsoperine ottersoni (Nematoda), hatching of 34 Hysterorchis vitellosus, new digenetic trematode from marine fish in New Caledonia 143 In Memoriam Gordon M. Clark 37 George R. LaRue 37 F. Ben Struble .... 37

Arthur C. Walton 37 Isoglaridacris hexacotyle (Glaridacris hexacotyle), reclescription of 193 JAIRAJPURI, M. S. Three new species of Actinolaimidae (Nematoda: Dorylaimoidea) from India 96 JORDAN, W. C. (see Shepperson) 106 KATES, K. C., and D. E. THOMPSON. Susceptibility of gerbils and young white rats to simultaneous infection with Trichostrongylus axei and Trichostrongylus colubriformis 102 Key to the species of Anaplectus 77 Key to the species of Carcharolaimus 96 Key to the species of Neoechinorhynchus 46 Key to the species of Radolphus _ 219 Key to the species of Rotylenchulus 169 LARUE, G. R. In Memoriam 37 Lasiotocus longitestis, new digenetic trematode from marine fish in New Caledonia 143 Levinseniella bijrdi, new trematode from the clapper rail in Georgia 62 Levinseniella hunteri, new digenetic trematode from Louisiana plover 140 LICHTENFELS, J. R., and A. J. HALEY. New host records of intestinal nematodes of Maryland rodents and suppression of Capillaria bonnevillei Grundmann and Frandsen, 1960 as a synonym of C. americana Read, 1949 206 LIE, K. J. Further studies on the life history of Echinostoma, lindoense Sandground and Bonne, 1940 (Trematoda: Echinostomatidae) with a report of its occurrence in Brazil 74 Life history of Echinostoma lindoense 74 Life history of Fergusobia curriei (Nematoda: Sphaerulariidae) 40 LITTLE, J. W., and S. H. HOPKINS. Neoechinorhynchus constrictus sp. n., an acanthocephalan from Texas turtles 46 LUMSDEN, R. D. Ultrastructure of the metacercarial cyst of Ascocotyle chandleri Lumsden, 1963 (Trematoda: Heterophyidae) _„_ 212 MACKIEWICZ, J. S. Isoglaridacris hexacotyle comb. n. (Cestoidea: Caryophyllidea) from catostomid fishes in southwestern North America 193 MAGGENTI, A. R. (see Croll) .._ _. 108 MAMIYA, Y. Rotylenchus pint n. sp. (Nematoda: Hoplolaimidae) from forest nurseries in T

Neotoma magister, new host for Capillaria americana and Trichuiis sp. 206 Neotijlenchus as a synonym of Hexatylus 154 Nervous system, description of in Thoracostoma californicum (Nematoda) 108 New combination Isoglaridacris hexacotyle (Linton, 1897) Mackiewicz, 1968 193 Laiogonimiis tananarivense (Deblock and Capron, 1962) Fischthal and Thomas, 1968 1 Nothanguina arcuatus (Thome, 1941) Nickle, 1968 __ 154 Paurodontus consobrinus (de Man, 1907) Nickle, 1968 154 Paurodontus linfordi (Hechler, 1962) Nickle, 1968 154 Reynoldstrema berghei (Vercammen-Grandjean, 1960) Fischthal and Thomas, 1968 1 Reynoldstrema luurenti (Vercammen-Grandjean, 1960) Fischthal and Thomas, 1968 1 New family Falcihastidae Siddiqi, 1968 (Nematoda: Belondiroidea) 248 New grade Dlplodiscus magnus (Srivastava, 1934) Fischthal and Thomas, 1968 1 New species (new genus indicated by * ) Actinolaimus armatus Jairajpuri, 1968 96 Allonematobothrium ghanensis Fischthal and Thomas, 1968 237 Anaplectus porosiis Allen and Noffsinger, 1968 77 Anaplectus similis Allen and Noffsinger, 1968 77 Anaplectus varicaudatus Allen and Noffsinger, 1968 77 Axonchium asacculum Siddiqi, 1968 248 Belondira murtazai Siddiqi, 1968 248 Bivesiculoides posterotestis Durio and Manter, 1968 143 Bucephaloides ghanensis Fischthal and Thomas, 1968 237 Carcharolaimus masoodi Jairajpuri, 1968 96 Carcharolaimus mujtabai Jairajpuri, 1968 96 Cercaria alganeshi Acholonu, 1968 259 Cercaria allomacarapanensis Nasir and Diaz, 1968 67 Cercaria coloradensis Acholonu, 1968 259 Cercaria denverensis Acholonu, 1968 259 Cercaria guaraunensis Nasir and Daiz, 1968 67 Cercaria marisa Nasir and Diaz, 1968 67 Cercaria olseni Acholonu, 1968 259 Cercaria pomacea Nasir and Diaz, 1968 67 Clinostomum hylaranae Fischthal and Thomas, 1968 1 Diphtherostomiim tropicum Durio and Manter, 1968 143 Dorylaimellus belondirelloides Siddiqi, 1968 248 Dorylaimellus paraclavatus Siddiqi, 1968 248 Dorykiimellus salvus Siddiqi, 1968 248 Eumasenia ghanensis Fischthal and Thomas, 1968 126 Euparyphium anhingae Premvati, 1968 ____ 197 Halipegus ghanensis Fischthal and Thomas, 1968 1 Harpyrhijnchus butorides Boyd, 1968 18 Harpyrhijnchus herodins Boyd, 1968 18 Helicometra antarcticae Holloway and Bier, 1968 30 Pleterorchis ghanensis Fischthal and Thomas, 1968 1 *PIysterorchis vitellosus Durio and Manter, 1968 143 Immature Didymozoid D Fischthal and Thomas, 1968 237 Immature Didymozoid E Fischthal and Thomas, 1968 _____ 237 Immature Didymozoid F Fischthal and Thomas, 1968 _ 237 Immature Didymozoid G Fischthal and Thomas, 1968 237 Lasiotocus longitestis Durio and Manter, 1968 143

Levinseniella bijrdi Heard, 1968 62 Levinseniella hunteri Heard, 1968 140 Mesolecitha ghanensis Fisclithal and Thomas, 1968 126 Metaplagiorchis biloborchis Fisclithal and Thomas, 1968 1 '•'•'•Myorhynchus pritchardac Durio and Manter, 1968 143 Neidhartia coronata Durio and Manter, 1968 143 •••Onchobdella aframae Paperna, 1968 200 *Onchobdella krachii Paperna, 1968 200 '•'•'•Onchobdella pterigyalis Paperna, 1968 200 '•''Onchobdella spirocirra Paperna, 1968 200 '•'•'•Onchobdella voltensis Paperna, 1968 200 '•'Palombitrema heteroancistrium Price and Bussing, 1968 54 Paracryptogonimus ghanensis Fischthal and Thomas, 1968 126 Pelorohelmins ghanensis Fischthal and Thomas, 1968 126 Petasiger floridus Premvati, 1968 197 Pleorchis ghanensis Fischthal and Thomas, 1968 126 Prosorhynchus longisaccatus Durio and Manter, 1968 143 Prosorhynchns serrani Durio and Manter, 1968 143 Prosotocus exovitellosus Fischthal and Thomas, 1968 1 Proteocephalus chologasteri Whittaker and Hill, 1968 15 Pycnadenoides ghanensis Fischthal and Thomas, 1968 126 Radolphus magniglans Slier, 1968 219 Radolphus nativus Slier, 1968 219 Radolphus nigeriensis Slier, 1968 219 Radolphus rotundisemenus Sher, 1968 219 Radolphus trUineatus Sher, 1968 219 Radolphus vangundyi Sher, 1968 219 Radolphus vertexplanus Sher, 1968 219 Rhipidocotyle ghanensis Fischthal and Thomas, 1968 237 Rotylenchulus anamictus Dasgupta, Raski, and Sher, 1968 169 Rotylenchulus clavicaudatus Dasgupta, Raski, and Sher, 1968 169 Rotylenchulus leptus Dasgupta, Raski, and Sher, 1968 169 Rotylenchulus macrodoratus Dasgupta, Raski, and Sher, 1968 169 Rotylenchulus macrosomus Dasgupta, Raski, and Sher, 1968 169 Rotylenchulus variabilis Dasgupta, Raski, and Sher, 1968 169 Rotylenchus pini Mamiya, 1968 38 Stephanostomum ghanensis Fischthal and Thomas, 1968 237 Stephanostomum trachinoti Fischthal and Thomas, 1968 237 New rank Axoiichiidae (Thome, 1964) Siddiqi, 1968 248 Nygolaimellidae (Clark, 1961) Siddiqi, 1968 248 Swangeriiclae (Jairajpuri, 1964) Siddiqi, 1968 248 NICKLE, W. R. Observations on Plexatylus viviparns and Neotylenchus abulbosus (Neotylenchidae: Nematoda) 154 NICKLE, W. R. (see Fisher) 40 NOFFSINGER, E. M. (see Allen) 77 Onchobdella voltensis, O. spirocirra, O. pterigyalis, O. krachii, and O. aframae, new nionogenetic trematodes from Ghana 200 Ostioloides rappiae (Trematoda), redescription of 1 Otter, helminths of in southeastern United States 118 Palombitrema heteroancistrum, new trematode from Costa Rica 54 PAPERNA, I. Onchobdella n. gen. new genus of monogenetic trematodes (Dactylogy- ridae, Bychowski 1933) from cichlid fish from West Africa _. 200

Paracryptogonimus glumensis, new digenetic trematode from Ghana 126 Pelorohelmins glumensis, new digenetic trematode from Ghana 126 Petasiger floridus, new echinostome from Florida birds 197 Pigeons, removal of Ascaridia columbae from, by thiabendazole 117 Pleorchis glumensis, new digenetic trematode from Ghana 126 POINAR, G. O., JR. Parasitic development of Filipjevimermis leipsandra Poinar and Welch (Mermithidae) in Diabrotica u. undecimpunctata (Chrysomelidae) 161 PREMVATI, G. Echinostome trematocles from Florida birds 197 PRICE, C. E., and W. A. BUSSING. Monogenean parasites of Costa Rican fishes. II. Proposal of Palombitrema heteroancistrium n. gen., n. sp. 54 Prosorhynchus longisaccatus and P. serrani, new digenetic trematodes from marine fish in New Caledonia 143 Prosotocus exovitellosus, new trematode from Ghana 1 Proteocephalus chologasteri, new cestode from Kentucky 15 Pycnadenoides glumensis, new digenetic trematode from Ghana 126 Radolphus (Nematoda), revision of the genvis 219 Radolphus vangundyi, R. magniglans, R. trilineatus, R. rotundisemenus, and R. vertexplanus, new nematodes from soil in Australia 219 Radolphus nativus, new nematode from soil in New Zealand 219 Radolphus nigeriensis, new nematode from soil in Nigeria 219 RASKI, D. J. (see Dasgupta) 169 Rats, experimental infection of with Trichostrongylus axei and T. colubriformis 102 Reproduction in Diploscapter coronata 24 Rhipidocotyle glumensis, new digenetic trematode from Ghana 237 Rotylenchulus, revision of the genus 169 Rotijlenchulus macrosomiis, R. clamcaudatus, R. leptus, R. variabilis, R. anamictus, and R. macro do rat us; new tylenchicls 169 Rotylenchns pini, new nematode from forest nurseries in Japan 38 SAMSON, K. S. The synonymy of the ruminant parasites Nematodirus oiratiamis Raevskaia, 1929 and Nematodirus lanceolatus Ault, 1944 52 SHEPPERSON, J. R., and W. C. JORDAN. A technique for isolating and maintaining cultures of Meloidogyne 106 SHER, S. A. Revision of the genus Radolphus Thome, 1949 (Nematoda: Tylenchoidea) 219 SHER, S. A. (see Dasgupta) 169 SIDDIQI, M. R. Five new species of Belondiroidea (Nematoda) from Sibsagar, India, with a revised classification of the superfamily 248 Stephanostomum ghanensis and S. trachinoti, new digenetic trematocles from Ghana 237 STRUBLE, F. B. In Memoriam 37 lamias striatus, new host for Rictularia coloradensis 206 Tamiasciurus hudsonicns, new host for Strongyloides robustus and Capillaria americana .... 206 THOMAS, J. D. (see Fischthal) 1, 126, 237 THOMPSON, D. E. (see Kates) 102 Thoracostoma californicum (Nematoda), peripheral nervous system of 108 Trematode, metacercarial cyst, infrastructure of 212 Trichostrongylus axei and T. colubriformis in gerbils and white rats 102 TSENG, SHU-TEN, K. R. ALLRED, and G. D. GRIFFIN. A soil population study of Ditylenchus dipsaci (Kiihn) Filipjev in an alfalfa field 57 Ultrastructure of the metacercarial cyst of Ascocotyle chandler i .___ 212 VIK, R. (see Meyer) 92 WALKER, M. L., and W. W. BECKLUND. A note on the morphology of Cooperia punctata (Linstow, 1907) and Cooperia spatulata Baylis, 1938 49 WALTON, A. C. In memoriam 37

WEBBER, A. J., JR., and K. R. BARKER. Hatching response of Hypsoperine ottersoni to canarygrass root emanations 34 WEHR, E. E., and M. L. COLGLAZIER. Thiabendazole as an anthelmintic against Ascaridia columbae in pigeons 117 WHITTAKER, F. II., and L. G. HILL. Proteocephalus chologasteri sp. n. (Cestoda: Proteocephalidae) from the spring cavefish Chologaster agassizi Putman, 1782 (Pisces: Amblyopsidae) of Kentucky 15 Zapus hudsonhis, new host for Longistriata dalrymplei __.. 206

MEMBERS OF THE HELMINTHOLOG1CAL SOCIETY;OF WASHINGTON ; . ^(Massachusetts through \fenezuela; first half m January issue) '->/

•' '-''/'• '••;.- iMbssdctiusetts) /Y ' / Stunkard,B. W. - . Harlow, D. R. ''; Germany ;A, ,/'{-••' '.' -:y. ; i v-- • .';.'- Boyd, Elizabeth/M. Tiner,VJ. D. Hopkins, S. H. - Greene, R. E. V . i/. , ' :: •.'"•',/ ' , Hurley.'F. J. / ' ,', f • White, ,1. V. •' ;"',>; ; Kvntz, R."E. Gt. firlt, & N. Ireland :'••:./• •-'•"•'! Riser/N.W. \. , ; North Carolina , ' i ' JLittle, J. W. ; v : , FrankUn, Mary T. ' -; :-',-\:.:'-, /\'- - ',Rohdev R.' A. j • v / , Barker, K. R. !'•;. "/<<. ,v Meade, T. G. - •• ' /- •'' Zuckerman, B. M.-Y Gort, W, W.>s * '< "'. ' Moore, D. V. Siddiqi, M. R. ' . ' ' ~' '• ', YY-/;"'--., -Michigan j £ Y ,/ ^.'V... Harkema, R. , /- '. V Myers, Betty' June !,' 'Williams, I.e. ' ', . / *• Y'/.'y / Ager.A. L. Y '','•'- Hunter; Wanda S./ ' ;:9 Read, C. P. Winslow, R. D.1 x '* r,' '--• ; Y.. ;1 . Clark, D. T. ; ^ . Vi • Khan, Sekender A. ; Santmyer. P. H..,- , {Greece ' \ H' '' :/• ••',_ ;r r • I y ' ', Cook, Alice R. '" >•" • McDaniel, Jl S. , /Smith, W. N. i Himonas, C. A. '..-, '• S''j: " ,' DeGiusti, D. »'. ; ''•', ,' Miller/G. C. ('*^>;f Thames, W. H., Jr. v Honduras ;' . ;''> -' , ' Y " ' ''•'Y '" K Dieter, C. E. - ."/ ' •'-• .' Reid, W. A., Jr. Utah ' .'-, .-• - Wehunt, E. J. ' '-N vV V-'-; •,-'/.' • DikrnansYG, . •', Y . Roman, J. - , / Griffjn, (G. D. ' lndi«i ; : -v -y:.--''v.-^,;' 'Peters. L:'E.,'Jr..''r ; Shepperson,.•;•;':, /:«55S»fe c. -R. - , ,• , Tilton, Beverly E.. / Havertz.'iD. S. ,- .' --..-Das, D. N. " '.' . -/i -• /.;••.-• ;! ••'- :..'• ; Minnesota '. Triantsp^hyllou, < " : '•' ' ' Virginia ' ... • .' ' Jairajpuri, M. S. ' •••Y- \ MaeDonald, D. H. ' ,HedwlgH.! ';-/- ' Freund, Frances E. .";/ -Kahn, Shahid H. , ,./ . 'V Palmer, T. T. ' (, 'North Dakota ': v . "' Hargis, W. J., Jr/ -, Khan, Abrar-M. , '/ - /:J: ''' '. ••'••• Vahde Vusse, F. J. Dyer.W.G. ?> r';.''\ Holloway, H. L.f Jr. li Porter, /D. A. V , W>f:7;. *V»i««Ppi ..;• - ,. / Larson, O. R. r ;; Hutton.R. F. v /Saxenai S.'K. :, '1 .'.-" . ,''"• ' -.'• ••>•'•'- \. , Owens, V. H. , Iieiby, i P. JD.^ • ' '' Miller, L. I. '.••'•; / Israel , ' -'/" \ /,-:.. ".•<'•:'. , 'YY' - Ward, J. W. , .'• •} Ohio. ,' \ •'•'. •"/- . " ;Reardon, Lycy V. 'V/-/;Mlnzi"p. i • ' :-;.-' -'••' '/Y/VU Missouri / ,!•'.'-,.. '. •-- , s Grites, J. 'it. •' •',•... Sonepshine, :D.' v'" ' ^ '• • "jEftperna • J. ' "-'\ ' ^ --''' '/' .•'•••"•,• -''•, ' ' Mullee, Mary T. /.•'/,.-'•' ;' Etges> F. J. ' .' ' /'\~-' • Washington i ', '\ -• ,i : 'Wertheirri Guta i ? • ;/.'"• 1-7 " .'.• Montana ' '• Harwood, P. t). '', ;; ;Chitvvood, B.'G. - ' Ivory .Coast / r •-.;.;>Ki-1 '• > •-,• S/ifc'j'A' '''/••; Canaris, A. G. ' '/ •••?V.K«rr, E.-D. " '. ':.. ' xS' ;Martin,G.W. , f S , iLuc "*M. " '.; ,''"'•'• '.;- ;''-' • , •! ' ; •-/•—' '• v'Camey W. P. v. ' Watson, D.E., / ; . : Saunders,'.T.:L. ' ,,. Jamaica',. :.-r'^ .''/'/• ''.«- '"'.. '•: -.'V-V/' Kinsella, J. M. ; '.'''. --' . Whartoh,G. W. N Senger. G>M. •/. 'BeverleyiBiufton; ;Mary Oklahoma West Virginia ;, ''->.'-• . ;- ''''.,.:• '/•-'i-!';,', :Yunker' .G;-iE. • '•' • .-•• "';Beschj-E..'D. '•,'<• ;. ''.;• Hall J. >E.' ' '-" ;. Japan ,• ! ' •',.' '. ,-. '", •• ',' . • ' •*• "•••• i Y'- 'Nebraska ' •('•''" -Hopper, F.,Ai, Jr. ; ' Hbffman, G/L. / "Ichinohe JM[. •' >: f'f;..'-i/|V..;". Y- Janovy» J-» jr. ./'''-' < ~ ~.*SelJ, J. ;T. '' ' ' ' v •'<' •'Putz/R/'E. , , -/ '/. , ' ', Hlnatomi, S. •,• /" .'.--./;"'• "Y •'• ! ;Yi's ••' 'Manter, H. 'W. •'•. ••' ' •'. - .Srnith, P. E> ! Wisconsin \ 'C : '. 'Maraiyat'y.. -•'.',•' •• ''. . ; .' '; ','i: ;7 ^Pritchard^Mary H. !, -Oregon '' ,. '•• ~- -;'. \, /J.3Ehorne Ri , G./ i 'i . ;Radke,.M.:rG,-;. - ./, .• / / () ";1; _.'•..-;. •Vii;Heva/da, , rl Y // - '• Todd, A.-C.,' !;,.•• l . Yokogawa, M. /Jenseni.'.H. J.' " ....-'•. "~-;!"." - Wyoming . ' »»J' VîMalaysfe , 'VV : ';/, [,,.•• Y'v V/'' "v : New Hampshire . ' • '•Knapp.iS.jE. ' Honess, R.'F:1' ; ' -/ '-. -flphde %K. '-' i \- !• 'i /V'Y ,;.' / ;;- ; v Bullock, W^L. : ^; .' '.'": Macy^R. W, --- • ,.; •'--";' Australia Mexico '' V. '' \:\'^ - ; . .• ' ^ ' '''';,.;; New Jersey ' , 1 ;Millemann, R. iE. - Sprent, J. F. A. : / ; ' ,-Briseno, C. -H. /;"-';'-":'/• >''('.'' ; '"' " -Balasubramiinian, 'M. i •' 'Nybergj.JP. A.\ •>" -I;V '/ Belgium ' „, ;-,-,;., ; . i ) l . / iGabftllero-y;C. '£• ••'(.'- • v'-' . -PanitZ.vE. !. ,'";-'''1' ;-';' '' ' ,'Goornans,'i A.1 -' \' /-,, - •,;• < ^•'•\\ Franco, E- j -^ 'v^ y^ Col|înt'E :;"'''" Nctherlonds '7.' 7''|rr'6;'/ ^ :'A/ ',' S^'vCuckler.Â.X:. • .-'Sen,. 'A/*.'.'"'" ','--. •''- ; • ' .. . .NDorsrhan,,vW. .. '-,, •,.-,*.. :. ,'. -' /' ;•-', ;Doscher,'Mary Ehlers . - ;Smithson;;Harrjei:jR. / .:•' Lordello, L. G. E. ' ,", -'New^Zealand '' ' ;- , < " ; 1. ,-•, •• *l^1',.v^'' -.-Gbble, F.,;C. i. >,- '^'"Pennsylvonia "\-';-.'//i -.'"'K' V^ Canada , . l ; , ' -.' Yeates, G. W. *•<• -v .;.;'.; ' \'." / --'v:/' '','-'." ' ffT Green, 'D. F. 1 ' •; •-.-- . .'Barron,:.C. -N.'.o '.; ' •' ; '.'.'{ , , Anderson,! K. G. / - Nigetio ... .:i-;'",'';rv.;": ''--•; :?''; j ; v ;4-Haege'lc, C,.L- -,•;. '.' . 1 Graham, G. L; ., -" '";, Anderson,?R. V. . '• '' ^ ;.,'•' V :• '-"'' ''"'Hung, ,Chia-ling / '' ':,H'eridtix, 'S. ,S. ,/.)'"; /';' r - Baker, A. D. J f? '"'!-' Pakistan .'-, ' V., ,. />• ', '/• .'•/ ,,--'•' ^y.: . >^:;/ IngaSs, 'J. W.,:jr.: ' : \Kingston. !N. -,; ' -' ' VBird,' G. W.,./. Vpr; ; ;'' - ,' ::«aleyi A. J. ,• V/;- - .- '.'. -': ?"- 'Jenkins, W. R. V , :Martin, H. M. / - 1 Bosher, J, E. Timm, R. W. ' ; ,:,/ ',••, j •' ..,.' ;• •-.. ;, (Kantor.'S. .!/H> ""v /'• > Meinkoth, N; A. " " V - Ching, 'Hilda; Lei . • "'Philippines ,•-• /' •' "•') ••'- ', •, ';; : •• - * -'Katz,..F. F. ';•'/ -;"'....'• '-', >Morseth, D. j. \ :\/Muncey, D'Dorney. W. , R/ST'- './A. ,' iDeJesus, ^. , t/i'.;^ ; Hopper, R. E. V' Velasquez/Carmen C. • i,< , i" " -, — , J'ankavich, J. A. . "-•./ ,.< "-Ogren,^. E.:;. '• • 'V-H':! Mountain, W. B. .1 Poland ~^':/ „ .v,;' V ;.;- '' ,, ' , ' Petriello; R. P. \ Soulsby, E. T. L. /'; ' ' ; ' 'Mulvey, R. H. /Bezuteik.B, J ^' ! ^, 'W\-tf '•!!'• " -; Rohrbacher, G.-H., Jr. • , Syljc, S. R. '.' ] ' ' ,:J Orchard, W./R. • '-Portugal-''. • >/'r •'' '' ' • ,:'-, '• . •'- Y ' ' '/- ' • .->'•' '- Rnwse, T." W, "/; • •; ,< " . ,;, pUricchio, W, A.1 Sanwal, K. C. ,• >•> .Bravo Lima, M. , '/ / T ' T ;--' ' ''Stbll,:N.,R.; .-,'-.' ]>'";;-^ ; , 'Puerto Rico ., .-;. ,\ ',"•'. Welch, ;H.E. « - South Africa ',' ; ;;",i -,' -;." - v-'l.' >V- • ••:•< /.- • New Mexico , t - ,-:',. L .'. Ayala, A. . \ 'Wu, Liang- Yu ' - - ' 'Heyns.J.. • : ' '•'''' -,'.;'•',; '.' . •. f -:v ? / / Allen, R: W. (. '-;,;;. V 'Oliyer^-Gorizales, J. -•, chile; ; ..",. :, /-'' • i<; ; x -Kruge'r, S.'Pi^ ,,,-,-, -\.V 'T'7^ ~. -'• ,• ;,'-' ' ,; >; ( Massey,/C.;Lv •,'•. . Rhode Island " V . ITagle, lisaias^V; : ' ; Switzerland ' .'./a ''/';.'. ,''-;,'»/ • .;!;'rv '-/;'...,.', /. "Riffle, J.'W. .'-;-/- 21nn, D. T. ' > . Colombia / •,- > - -.'• - iBaer J. )G. ''/* (v ' ","••.•':'. ." :!-".' k' -" i ^V SfltHSOIU'K* o» ':.. " .South Carolina - : • Gonzales-Mugaburu, L. , D.iibois, G. ;l^' . :7. >- '. •'j r '/ -'• •- '• lA'ilsoDj' G« I« •" ''"" r Adkins, T. R., Jr. Costa Rica '.. Taiwan ," . JV'' ";';''. v-:\'y- / ' Graham, T. W. - ^ .BrenesiM., R./R. o Armstrong, -D. E.:''' •'• \- >' ^•; ••':•'• \,:\'-: ,'N«w,York , ' - ;,.•";• -'',; - - ' rt; /"I T- ' . ,?:,' ""•',', ' "V •'•'• /: '; /'.Braun, A. J. -, ' • /', Rall, ij. J. i •' V-^ . Salas, F,, L. A. "; . Bergner, J.JF., Jr. ' , •-.- •?,-/;/'> -/''-'Tischthal/J.'JI. •.,.' '' South Dakota, ' ''',,. Cyprus - ' , '•-'•: .'. ','• ! '- ',' ' " ''Ghiu,. Jui-Kuang _ • ' ' '» t/^'i ,'.•;•'),:-:/;:',/, Granek,'!/.... j 'i -" :' Greichus, A. / ,\), '''/>*' Phijis, J. - ' ' '- (;' - .Gross,!, J,-'«,: - " ,v, . '., . ' ,:<; /:v ' Hasselwander, A, J. . Hucghins, E. J. ' ' , , Egypt . ,,'--" , Thailand ' • •••-.-;; ?.--•;• '.•/- \Wckson, G. J. : ? ,' ' /v McDaniel, B., Jr. v, ;> : Ibrahim, I. K.T^.- . ''.•s ' Ratanaworabhah, S.' ! ,J'/v,, '-i/v ;:i ij'Mackiewicz, J. :S. Tennessee' ;Oteifa,iB. A: . U.S.S.R. , •' "*'••'-;: . '-ii IV Bpgitsh, B. J."-; , Shafiee, M. F.! '"', ', ' '•'• • Ershov.'V, S. .' .. ,;x ~4 . >-/ ^' :! ^/^ '.,a ^ Mueller^ J.*F. ' • • - v , / . /Cosgrove, G. E. ;: .-,•••'•'• " •France- ' •'• ''. •-, ' ~v '.; . Venezuela ' < . ; : '"'-.- '.. ,- '.' •' \\ \ iNasSr,'?.' ,..'-'}'( !;,/••; :' 'i/-' ' ''.•/! Y .•.'; '."' }y'S.chroedevP. ' ;., !'V ' Texas , - ' ;; ;'' .• /' \. . Dollfus; R. xPh. ' /X'.-V-' .'.'•-, /.( Sirh'aren, J. O."',. .';, ', • ^Bello, T.R. ^ , | /- ;Yitiell6, P. ' , ' }v" ;>

'- VOLUME 35 JULY 1968

, vCQNIENTS ((Continued from Front^Cover)

FlSCHTHAL, JAC'QB H,, AND J. D. ,THOXiAS. Digenetic Trematodes iof Some Freshwater and Marine Fishes from Ghana Ii~-L. ^j- 126 FISCHTHAL, JACOB H.i AND J. D. THOMAS. Digenetic Trematodes of Marine Fishes , • from Ghana: Families A^an&ocolpidae, Bucephalidae, Didymozoidae '".„ --.:-_ •ssn. HEARD, RICHARD W'., III. Levinseriiella hunteri sp. nbv., a New Species,of Microphallid ; /; Trematode-from the Wilson's^ Plover,; Charadfius wilsonia Qrd ".;1_..T J..X :-—..„> 140

Index, to Volume's 35 ... „!'.—^-^.—._„ r T-,-.J- . ;_ „ .„.. _/. ^.„....^:. 272

\• • ^ " . ' , • y • J,- - ;' ' —- • • ^'"r • ' - '<>.—- ' / LiCHTENFELSj'J. RALPH; AND A. JAMES HALEY. .New-Host Records of Intestinal Nema- •'''•:"' ,. todes of Maryland Rodents and Suppression of CapiUnria bormebillei Grundmann s' -and 'Frai^dsen, 1960 vas a Synonym of C. 'americana Read, 206 LUMSDEN, RICHARD Dl , Ultrastructure of tlie Metacercarial Cyst of Ascocotyle chandleri- \2 \ •••••'• ILumsden, 1963 ^Trematoda: Heterop^yidae ) -i.:.-^..^...^.:..!-!....^.-....;:...,^..^..^

!MAGKIEWICZ, JOHN S. Isoglatidacris hexacotyle comb. n. /(Gestoidea:xCaryophylIidea) >•. from Gatostomid ^Fishes in Southtwestern /North America; •'.•'^•..tlXi.-^-— ."-.-—--«-..- _____ — - --^ 193 MILLER, GRQVER C., rAND REIN Aiu) HABKEMA. Helminths' f Monqgenetic T3?ematodfes ('Dacty- Ipgyridae, Rychowski 1933) frbm Qichlid 'Fish from West Africa _-,-.-...-.-_.^-._..._.. V200 POINAB, GEORGE O., JR. r Parasitic IDievelopment of Fitipjevtinermis -leipsandra IPoinar / and[Welch ( Mermithidae ) in £>iabfot ica 'u. undecimpunotata (Chiysomelidae)—-„ . 16,1

G. Echinostome .Trematodes from .Florida Birds \_l_j:^l^-_.-^;-:----..j.^>^—- 197 Report ofthe Brayton H.jRansom?Memorial Trust Fund— ^ill.-.-....:____ _ .....--^-.-.i-._._.:.-..- 160 .SHER, S. A. ' Revision of the ..-Genus Rddopholus Thome, 1949 K Nematodac Tylenchoidea ) 219 RAFIQ. Five .New Species of ;Belondifpidea ( Nematoda ) Sibsagar, iridia, .witli -a, Revised Classification, of the Siiperfamily ,„.,______.. .____ _ . 248 WEHR, EVERETT E., AND MERLE L. COLGLAZIEH. Thiabendazole as an Anthehniritifc ..:'.-"' •; A Against Ascatidia colunibae in Pigeons — — .-- <117

•>'4^}:^^^i'S: . - MAILING DATES FOR VOLUME 34 --- 'y.r •' , ,yi>. V.- <( ;"y:^Number ,1, 14"February;1967 ' ;-; ':"•'• Number 2, 7 July

••• \j- • -'-;-•.... •;•-'-•• . "• ,'• - ^^^'l^'.}^^';1^^^ .:-' g ' ' ~':<^:^-^.J,A\' 'J^'^Y-y • -•r-.:''. -;r' .':• ;»-.!.'".V,'• • .v .••'•-"";-"~ '" v ' -; -'•••.' .'">•;- . •-\ii^ MArLmG'DATEs FQR VOLUME '35 -, j \\: Number'l^e Jmiuary 1968 1 : , ! Number2,16 July 1968 ; ^

ALLEN PRESS, INC LA\WRENCE, KANSAS

•'^:A);(;> '.^•;•':'/''':'> -:>]YY..: j-^'-,;