University of the Pacific Scholarly Commons
University of the Pacific Theses and Dissertations Graduate School
1977
Some helminths from salamanders of California
José Joaquín Castro University of the Pacific
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Recommended Citation Castro, José Joaquín. (1977). Some helminths from salamanders of California. University of the Pacific, Thesis. https://scholarlycommons.pacific.edu/uop_etds/1937
This Thesis is brought to you for free and open access by the Graduate School at Scholarly Commons. It has been accepted for inclusion in University of the Pacific Theses and Dissertations by an authorized administrator of Scholarly Commons. For more information, please contact [email protected]. 6 ~ ~ C------SdME HELMINTHS FROM
SALA}~ERS of CALIFORNIA
!---~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~]"~-~'--=--c_c
A Thesis
Presented to
The Graduate Faculty
of the ------University of the Pacific
-~-
In Partial Fulfillment
of the Requirements for the Degree
Master of Science
==
I Jos~ Joaquin Castro
May 1977 ~--- This thesis, written and submitted by ~ ·--- Jose/ J. Castro
on Graduate Studies, University of the Pacific.
Department Chairman or Dean:
fR-.~·
Thesis Committee:
Chairman
·---
Dated~ __M__ ay~·--3~,_1_9_7_7 ______
- --- ·~ ACKNOWLEDGEMENTS
The author wishes to express his deepest gratitude to Dr. Fuad Nahhas for his helpul suggestions and criticism, his encouragement, and inexhe.ustible patience during this investige.tion; to Dr. Lee Christianson for his assistance in the identification of the salamanders, and very dearly to Dr. Alice Hunter whose influence made possible his acceptance to the graduate school and for her ever readiness to supply him with microtechnical materials needed in this study. The author is also very grateful to William Carey, Helen Ferguson, and Elisa Smith for their assistance in --- the collection of salamanders, and finally, to Janet Ahn whose moral support, patience, constant help and unders tanding made this study possible. ~------
TABLE OF CONTEN'rS .;::::;----
~ Page --- HISTORICAL REVIEW. • • • • • • •. ·• • • • • • • • • • •. • 1 M.A. TERIALS AND }lETHODS • • • • • • • • • • • • • • • • • 4
.-~r{r~· ------~---~'---'--
Distoichometra ensatinae sp. nov. • • • • • • • • • • 8 Baerietta diana (Helfer, 1948) Douglas, 1958. • • • .10 NEHATODA • • • • • • • • ... • • • • • • • • • • • • • 13 Cosmocercoides dukae (Roll, 1928) Travassos, 1931 •• 13
Dibulbiffer caballeroi sp. nov. • • • • • • • • • • 15 Pseudaplectana cablei sp. nov. • • • • • • • • • • 17 ------
Pseud~ple~~~~ ~ltoni sp. nov ••••••••••• 18
Rhabdias sp •• • • • • • • • • • • • • • • • • • • • 20 Thelandros magnavulvaris (Rankin, 1937) Schad, 1963 .21
Thelandros minutus Re~d and Amreim, 1952. • • • • • 22 Thelandros salamandrae Schad, 1960 • • • • • • • • 23 SUM~IARY • • • • • • • • • • • • • • • • • • • • • • • 26 HOST-PARA.SITE LIST FOR THE WEST COAST • • • • • • • • 27 LITERATURE CITED • • • • • • • • • • • • • • • • • • • 29 TAB"LE I Heasurements on Distoichometra ensatinae sp. nov •• • • • • • • • • • • • • • • • • • • • • • 32 T.ABLE II Heasurements on Dibulbige.!. cabal.leroi sp. nov. • • • • • • • • • • • • • • • • • • • • • • 33 TABLE III Neasurements on Pseudaplectana cablei sp. nov. • • • • • • • • • • • • • • • • • • • • •• 35 TABLE IV }leasurements on Pseudaplectana. >val toni sp. nov. • • • • • • • • • • • • • • • • • • • • • • 37 PLATE I Distoichometra ~;;..;;;.,.;..;..;;ensatinae __ sp. nov~ • • . .. • • 39 PLATE II Baerietta diana (Helf'ei·, 1948) Douglas, 1985. 41
PLATE III Cosmocercoides dukae (Holl, 1928) Travassos, 1931;Dibulbiger caballeroi sp. nov •• • • 43
PLATE IV Pseudaplecta!l!l cablei sp. nov.; Pseudaplectana waltoni sp. nov. • • • • • • • • • • 45 PLATE V Thelandros magnavulvaris (Rankin, 1937) Schad, 1963; Hhabdias sp. • •••••••••••• 47
------~ --- .. =------~ PLATE VI Tholandros minutus Read and Amreim, 1952; Thelandros salamandrae Schad, 1960 • • • • • • • • • 49
::;:_:__::__ __-_=-:.=_-:..==--
----
- - " HISTORICAL REVIEW
Studies on the hel~inth parasites of salamanders from ~-- 'l'j------~ California have been very limited. Helfer (1948) described §------.-'--- ~ Proteocephalus diana from Batrachoceps ~· attenuatus and
Ens a tina e. eschschol tzii taken in Contra Costa and }Iendocino
counties, and Proteocephalus interaneides.from Aneides ~-
!?.!'=!-.!!. collected in Alameda and :Hendocino counties. Read and
t !'!a 0.:::- --- !--~----AA:-~u-r-e--i-m-(-1-9-5-2-)-cl-e-s-e-P-i-S.-e-El-rp_h-e--l-a-n-El-Po-s-m-i-n-u-tu-s-£-X!-&-m J3-a c hoc e P-~ ~------H_-:-_:--- ~· attenuatus collected in Los Ahgeles. Lehmann (1954)
reported Oxyuris dubia Leidy, 18,56, from the rectum of~-
~ ~· eschscholtzii taken in Sonoma County, and Batrachoceps
~· attenuatus collected in San Francisco County. Lehmann
(1954) also found Q• magnavulvar:i.s Rankin, 1937, in the rec-
tum of f!.Eeide~. flavipun~ctatus :collected :J.n Narin County, Rhab ---- ~ sp. in the lungs and Cosmocercoides dukae (Roll, 1928) Travassos, 1931 in the intestine and rectum of aquatic Taricha torosa collected in Cnntra Costa County. Douglas (1958) ----- transferred f• diana to the genus Baerietta. Lehmann (1960) reported Hegalodiscus americanus Chandler, 1923 and Q.• dubia
Leidy, 1856 from the intestine o!~ Dicamptodon ensatus, and
Rhabdias sp. from the lungs of the same host species. Leh;..
mann (1960) also found rarus Thomas, 1937 in the intestine o:f
Taricha. torosa; all from }larin and Sonoma counties. Yamaguti
(1959) transferred Proteocephalus interaneides to the genus ;_--- -- ·:- ---- Baerietta., Johnston (1962) reported .Q.• dtibia, Leidy, 1856,
1 2
f'rom Ane~ lugubris, Ensatina ~· eschschol tzj.i and
Taricha torosa sierrae, Capillaria sp. and :r.Jegalodiscus sp.
f'rom Taric~ torosa taken. in the Central Calif'ornia Sierra fi------n Nevada Hountains and Tuolumne County. Hizelle et ai. (1968)
.described a new monogenetic trematode, Gyrodactylus ensatus. ~-- ~---- f'rom Dicamptodon ensatus collected in Humbolt County, and
}lizella et al. (1969) described another new species Gyrodac
tylus ambystomae f'rom ;4-mbyst,oma macrodactylu,!1l taken in La:s-
~--~------:~~n-~oun-ty•• ------~------~------~------~------~------~~~=== H~- Although not f'rom Calif'ornia but pertinent to this
survey are the reports of' several investigators f'rom Oregon
and British Columbia: In Oregon, Lynch (1936) described
Phyllodistomum singulare; Senger and ::Hacy (1952) described
.T~uryhelniis pacif'icus and ( 1953) Cephalouterina dicamptodoni;
Hacy and Demott ( 1957) reported !:~alipegus £.S,£i~i.,! ---- Staf'ford, 1905, and McCauley and Pratt (1959) reported
J:.Iegalodiscus micropha~l~ Ingles, 1936; all f'rom Dicamptodon
ens a tus. Taricha eranulosa and Ens a tina e schschol tzii '\vere
-~-- f'ound by Lehmann (1954) to harbor Brachycoelium salamandrae
(Frtnich, 1789) Stiles and Hassell, 1898. Taricha torosa
was also found by Le.hmann ( 1954) to harbor }iegalodiscus
americanus Chandler, 1923. In the same host sp(;!cies, .I•
torosa, }facy and Demott (1957) and Macy et al. ( 1960)
recovered Halipegus occidualis Staf'f'ord, 1905. Lehmann (1956) " reported ~galodiscus americanus Chandler, 1923 in Taricha
sp. and Ambystoma gracile.
The only study made in British Columbia is that o:f Ef'f'ord and Tsumura ( 1969). These authors f'ound ~!e{l'alodiscus. rnicrophagns Ingles, 1936, £.
Senger and ?>1acy, 1953 and .9Ethocephalus truncatus PalJ as, n--- -- "10--;-:--;--_--=o.~ 1781 in Ambystoma gracile. They also found Negalodiscus E!~'--~~-- -'~-~- t_;_ ____ micronhagus Ingles, 1936 in Ambystoma macrodactylum and ~" ---~ p ~ --- Taricha ~ranulosa, and Cephalouterina dicamptodoni Senger and Macy, 1953 in Taricha granulosa.
------"'
!::;
-----
----- ;.; ~~TERIALS AND METHODS
;_j The 340 salamanders were.captured by hand from var- ~-- ~c~~ !§ '"'c--·-- ious localities in California by turning over logs and L_-__-__ =-_ rocks, and stripping the bark of fallen trees. The animals
~="-~ ~~ lvere brought to the laboratory and killed by beheading a ----
day or two after their capture, pinned down to a_dissecting
tray, the ventral side cut open, and the internal organs
-----~~~~~~-~~~~~~~~~~~~~~~-~~~~~~~----c--~1~--- removed and placed in a 0.7%NaCl solution. The intestine tL
and other internal organs were teased apart in a petri dish
with ~ dissecting needle under a dissecting microscope.
The parasites were removed with an eyedropper and placed in = fresh saline to. rid them of debris and host's tissue. Once
the.parasites were clean, they were processed as follows:
Cestoda ------' To facilitate the study of' their internal struc•
tures,~cestodes must be fl~ttened before they are killed.
The flattening process was carried out on a slide under -~-~~ slight cover slip preassure for about 15 minutes and at the
sam~ time, A.F.A. {alcohol-formalin-acetic acid) was
applied at the junction of cover slip and slide to .kill and
fix the worms. Upon completing the flattening process, the
. . . cestodes were removed from the slide, placed in stender
dishes and kep-t over night in A.F.A.
The A.F.A. solution was replaced by 50% ethanol, followed
by 70%, and stained overnight with acetocarmine. Destain
ning wa~ acomplished by washing the wb~ms with acid-alcohol
4 ------. 5
(0.5 HCl in 70% ethanol) • The -worms were dehydrated·. in a
graded series of' alcohol of' increasing strehgth, 80%, 90~~~
and 100% f'o~ 15 minutes ench, .cleared in a mixture of' 1:1 ~ a·~---~-~~ !!--·····--'-···· ethanol-methyl salicylate and later 1 007b methyl salicylate I ..
8-~-- washed "'tvith xylol and mounted in permount (Fisher Scientific g____ _
Co •, N • J.) • Nematoda
Because of' their tough cuticle, nematodes were
pouring boiling glycerin-..alcohol ( 1 07~ glycerin in 1 OO%
ethanol) over the worms to kill them in an extended posi-
tion. The neraatodes w·ere kept in stender dishes, covered
with lens paper. to prevent dirt and dust penetration and to
allow ~lcohol evaporation, leaving only the glycerin
which penetrates and dehydrates the worms. ·.The: nematodes -----
were mounted by placing a drop of' glycerin-jelly (Carolina
Biological Supply House) on a slide, immersing the wprms in
it, and applying a small amount of permount (Fisher Scien-
----~ tific-C6., N.J.) around the glycerin-jelly to geal the
cover slip and prevent dehydration.
All measurements are in mm except f'or eggs and paruterine
capsules which are given in microns~ Drawings were made by
micro-projection. New· host records are indicated by an
asterisk, The numbers in parentheses f'ollo1dng host name
represent-the number of individuals that harbored that
particular parasite (numerator) and total number of' sala-
manders examined (denominator). Measurements on each new 6 species are given in Tables I, II, III, and IV. These
represent the smallest and largest mature specimens and
structures f'ound, the mean and the standard deviation. ~------
~------~------~------~--~----~------~------!~-----
= THE NEHATOTAENIID CESTODES
There is a general agreement among cestode spec- ~ n------~ ;;~~~- ialists (Lawler, 19.30; lfardle and }1cLeod, 1952; Yamaguti, i...... c~.
W--- 1959) that the four genera Cylindrotaenia, Baerietta, Q! ---- stoichometra, and Nematotaenia belong to the family
Nematotaeniidae, of the order Cyclophyllidea. lvardle et al.
(1974), however, removed this family from the order Cyclo-
------phyl-riaea, and placed~i t in a new order Nema totaeniiaae; th=-e=--~--:~=~=-- authors justified this change on the following basis:
11 Nematotaeniids are readily distinguishable from cyclophyllideans by the almost cylindrical body lvi th segmentation restricted to the posterior body region; by the nonbothridiate, nonrostellate hold fast (although Law·ler 1 s fiffUre of Cylindrotaenia . auadrijugosa showed, no doubt by mistake, a hold fast with a rostellum and well-developed bothridi form suckers ( lvardle and 1-'lcLeod, 19 52, f'igure 186); by the testes numbering only one or two per segment; and by the almost complete disappearance of the q transverse tubular uterus from each gravid segment. The eggs pass one or more at a time into egg cap sules formed from the uterine wall and from thence eventu~lly into hollow, somewhat pear-shaped spaces within the parenchyma." --- \vardle et al. ( 1974) ch~racterize the genus Distoichometra
Dickey~ 1921, as follows:
11 '1'wo testes per segment; paruterine organs numerous, each containing three to six uterine capsules with fused bases, arranged in parallel rows. Genotype and only species, bufonis Dickey (1921) from Bufo species in Georgia."
The above authors were evidently unaware of' Douglas' (1958) work on the Nematotaeniid cestodes.
The same authors spearate Distoichometra from Nematotaenia
Ltlhe, 1899, by 11 egg capsules clustered together 'vith bases
7
tl 8 fused compared to egg capsules scattered throughout the parenchyma." Other closely related genera are Cylindrotaenia
Jewell~ 1919 and Baerietta HsU, 1935. Wardle et al. (1974) characterize the genus Cylindrotaenia Jewell, 1919 as follows: ,,~ ___ ''One testis per segment and two paruterine organs each with three uterine capsules and united basally."
More recently Ulmer and James (1976) emended the family
with the following characteristics:
11 Testes 3 to· 10 (usually 8) dorsal, medullary. Ovary median, posteroventral to testes. Paru terine organ single, containing approximately 20-40 eggs. Parasitic in int~stines of frogs."
'.rhe above authors did not accept the ordinal status of the nematotaeniids as proposed by Wardle et al. (1974} and -- retained.the family in the order Cyclophyllidea.
In the present study two nematotaeniid genera are represented: Distoichometra. and Baerietta.
Distoichometra ensatinae sp. nov.
PLATE I Figs. 1-5
.TABLE I
Host: Ensatina ~· eschscholtzii Gray (11/52) !• !.• sierra.e Storer (9/52)
Site: Small intestine
Locality: Oakland
Description based on J4 specimens, measurements on
14: Tota.l length 17.00-40.00; scolex without a rostellum, 9 0.200-0.450 in width; sucke~s four, circular, 0.090-0.120
in diameter, segmentation noticeable orily in mature and most
posterior proglottids; immature segments wider than long,
mature segments almost as long as wide, and posterior seg-
ments longer than wide; the last segment 0 •. 200-0 • .500 X o. 350-
1.210. Testes two, 0.055-0.100 X 0.090-0.200; the seminal
vesicle is long and somewhat coiled. The cirrus sac, 0.033-
0.050 X 0.042-0.120, is almost spherical in the immature and
------;.---.-----T.iu.ew-ly-mature-s-e-gnrent-s--,--ovo-r-d-~n-t_h_e rna ture segments, and_------~;=_=.. === flask-shaped in the most mature and gravid segments. The
cirrus sac and vagina are medullary in the immature and
newly mature proglottids but gradually become cortical as
the segments become more mature. The ovary is almost spher-
ical 0.-55-0.088 X 0.060-0.114; the vitelline gland, o.o44-
o.o66 X o.055-0.100, is usually posterior to the ovary.
The uterus :forms f'our to seven paruterine organs, each 0.0.50-
0.11.0 X 0.050-0.120; the encapsulated eggs or paruterine
capsuies-are 24-JJ X 44-66 microns. ----
Two species of' Distoichometra are.known to date: !!•
buf'onis Dickey, 1921, f'rom ~ lentiginosus in Georgia and
Q• kozlof'f'i Douglas, 1958 f'rom Hyla ~gilla in Oregon. Distoichometra ensatinae sp. nov. may be distinguished f'rom
Q. buf'onis by its smaller size {17-40 compared to 100),
fewer paruterine organs (4-7compared to 8-12), relatively
larger testes {0.073 X 0.140 compared to 0.067); and larger
cirrus sac 0.039 X 0.079 compared to 0.031 X o.o48). . 10 Q. ensatinae sp. nov. is very similar to and may be
identical w·i th Q. kozlof'f'i Douglas, 19 58. Q• kozlof'f'i
Douglas, was described f'rom the tree, f'rog Hyla regilla 1958 ': R-Ci .. in Oregon. The main dif'f'erences are in host specif'icity and
;l- geographic distr~bution. The measurements f'or both species ~----- show a great deal of' overlap except f'or the average number
of' eggs which is 2-9 (average 6) f'or D. ensatinae, and 2-5
(average J) f'or Q• kozlof'f'i.
Baerietta diana (Helf'er, 1948) Douglas, 1958
PLATE II Figs. 104
Synonym: Proteocephal.~ diana Helf'er, 19l.J.8
Host: Batrachoceps ~· attenuatus (Eschscholtz) (55/250)
·site: Small intestine
Locality: Oakland
------Description and measurements based on f'ive mature ~------
specimens: Total length 16-24; scolex without a rostel.lum;
suckers four, circular, each o.oso-0.120. Segmentation
noticeable only in the mature and most posterior proglottids;
immature segments wider"than l.ong, mature segments almost as
wide as·. long and posterior segments longer than '"'ide • The
last six segments measure 0.190-0.400 X o.4oo-o.soo. Testes
two, 0.033-0.073 X 0.055-0.120. Ovary ovoid, o.o44-o.o66 X
o.o66-0.088; vitelline gland ovoid, o.030-o.o66 X 0.055-0.120;
cirrus sac, 0.031-0.066 X 0.055-0.090. Paruterine organs
o-..-.~ two, o.o66-0.088 X 0.066-0.120, united basally; the encap- ~-
sulated eggs or paruterine capsules are 22-33 X 24-48.
Each paruterine organ houses two to eight paruterine 11 capsules. The hexacanth embryos are 11-15 X 15-17.
The genus Baerietta was erected by Hst\ (1935) to accomodate a new species Baerietta baeri from ~ asiaticus ~r:------
1]-~~ in Nanking, China, and !l• jM.gerskitHdi ( :Tanicki, 1928) from t W--- ~ in Africa. The genus Baerietta is characterized by the i _____-~ ~--- presence .of tw·o paruterine organs united basally, and two testes per segment. Yamaguti (19.59) transferred.Proteoce- phalu!!, interaneides to the genus Baerietta. I agree with
------Yamaguti that this species does not belong in the genus E.!.2,- " ~------teocephalus; ho1vever, its placement in the genus Baerietta is also questionable. Helfer's ( 1948) dra1vings of this para- site show six paruterine organs in the gravid proglottids, and the description reads as follows:
"Then the uterine sac grows for1-1ard, the eggs move out into the sac, and it separates off: into six
separate round egg capsules." ------
The author considers ,!le interaneides as species insertae sedis. In addition to ~· diana and ~· j~gerski8ldi, seven other species have been described: !l• janicki (Hilmy, 1936)
Douglas, 19.58 f'rom -Rana and Xenopus in Liberia; .:§.• japonica
Yamaguti, 1938 f'rom Hyla and Polinedates in Japan; !!• clavii'ormis Yamaguti, 19.54 f'rom ~ in Japan; !l• montana
Yamagp.ti, 1954 f:rom ~ in Japan; !l• desmognathi Douglas, 1957 in Desmognathus, locality unknown; !!• idahoensis 1vaitz and Herha, 1961 f'rom Plethodon in Idaho, and !l• gerrhonoti Telford, 1965 f'rom Gerrhonotus in California. _The name :!!.• desmognathi appeared in an abstract (Douglas, 1957)-under the title "Fertilization, Naturation and Early Clevage in 12 !i• desmorrna thi sp. nov. Nema totaeni.idae. 11 I have been
unable to f'ind any description of' this parasite and there-
f'ore, this name must be considered nomen nudum.
------!------~------:5--··_··--= !L--=:c -- -- ·--
..
a-----
---- NEl-fATODA
Cosmocercoides dukae (Holl, 1928) Travassos, 1931 ~- PLATE III Figs. 1, 2, and 3
~----- ~ Synonyms.: Cosmocerca dukae Holl, 1928 ~----- " Oxysomatium variabilis Hardwood, 1930
Host: Taricha torosa (Rathke) (1/6)
Site: Large intestine
1--'------c---•L'o"c""a"':ti ty: Sacramento valley (exact location unknol'ln)'------·~[=_-=._=•.-.-·=-··=--
Description and measurements based on seven speci-
mens; £our females and three males:
FEl-ULES: Brown in color, 5.00-6.00 in length; width
through the vulva 0.264-0.390. The cuticle is .thick;
lateral allae extend £rom the anterior part of the body~
0.150-0.250 from the mouth and gradually taper near the ------
posterior end; transverse striations and longitudinal ridges
are present. The mouth is bound by six small lips; the
pharynx is o.o66-0.099, the esophagus, 0.560-0.690, and ------the esophageal bulb 0.09.9-0.110 X 0.110-0.143; the intes-
tine is tubular, extends to near posterior end; anus-tail
distance o.286-0.J50; the posterior end gradually tapers
into·a sharp pointed tail. The excretory pore is anterior
to the level of the esophageal bulb, o.450-o.soo £rom the
mouth. Two ovaries and two titeri are present; the vulva is
posterior to the middle o£ the body. The eggs are 35-J7 X
59-66 microns; the eggs contain fully developed embryos.
MALES: Bro1~ in color, 3.0o-4.oo in length, maximum width
1) 14 0.224-0.290. Cuticle and mouth parts as in the females,
but lateral allae narrow·er. Pharynx 0.055-0.066; esophagus
0.454-0.590 in length; esophageal bulb 0.077-0~097 X 0.090- 0.110. Anus•tail distance 0.120-0.200; 17-18 pairs of
h---- preanal, one pair paranal, and two pairs of postanal all of §" ___ _
the rosette type, and three pairs of preanal and postanal
papillae of the simple type are also present. Spicules two,
each 0.480-0.550 in length; the gubernaculum is 0.1h3-0.170
------'-----~1-on-g·-.------~~~·--·_-·_.. c.
The genus cosmocercoides '\vas erected by l'lilkie
(1930) and characteriz~d as follows: Lateral allae present
or absent, mouth bound by three small lips, esophagus with
a short pharynx, excretory pore anterior to the level of
-~------the esophageal bulb, two spicules and a gubernaculum pre------
sent, and 11-20 pairs of prea~al and three pairs of post-
anal rosette papillae present in the male.
In addition to £• dukae (Holl, 1928) Travassos, 1931
-----~ six other species have been described from the digestive
tract of amphibians and reptiles: £• tridens Wilkie, 1930,
from T.Ylotryton in Japan, £• nulchra 1Vilkie, 1930 from ~ in Japan and China, £• rustica (Kreis, 1932) Chitwood, 1933
from Poligyra in U.S.A. £• skrjabini (I'tvanitzky, 1940) Skjr.,
Schilm.lob, and Hosg, 1951 from ~ in Russia, £• bufonis
Karve, 1944 from~ in India, and £• rickae Ogden, 1966
from Ceratophora in Ceylon. I accept Q• ~abilis as a
synonym of' £• dukae (Holl, 1928) Travassos, 1931, as 15 . suggested by Yamaguti (1961). I am also in agreement with
Ogd<~n (1966) that Q. multipapillata Khera, 1958 is a synonym
~ of £• bufonis Karve, 1944. r;------~-.. --~ t~-~-~~c:·· DibulbiGer caballeroi. sp. nov. ~=-- PLATE III Figs. 4, 5, and 6
TABLE II
Host: Dicamptodon ensatus (Eschscholtz) (1/l) k-----..;JS~i-_tt'."e"..':i______.L-"a~r"'-:2g-"'e'____Ji~n~t..,_e""-E's__~-t..Ji~n!'e~------:-----1t~~~~~ Locality: Guerneville
Description and measurements based on 15 spec-
imens; six females and nine males:
FE!-iALES: Whitish in color, 5.80-7.20 in length, 0.143-
0.275 in width at the level of the vulva; the mouth is
bound by three lips bearing two buccal papillae each; next ------to the mouth there. is a muscular pharynx 0.060-0.075; the
esophagus con~ists of three parts; the first portion of
the esophagus measures 0.720-0.810, the.second part is a
----- ·-- - - prebulb 0.077-0.088, and the third part, the esophageal --·-----
bulb, 0.099-0.120 X 0.110-,1J2. The anus is located near
the posterior end; anus-tail distance 0.242-0.275. Ovaries
and uteri paired, extend from the esophageal bulb to near
posterior end, 0.750-0.800 from the tip of the tail. Vulva
posterior to.-the·middle of the body. The eggs are 33-38 X
44-55, covered.with a smooth shell. :::-- MALES: Total length 4. 70-6.70; maximum w'id th 0. 138-0.280;
the mouth· parts are the same as in the female·•. The pharynx 16 is 0.055-0.070; the esophagus 0.665-0.74·0; the prebulb
0.067-0.081 and th• e~ophageal bulb 0.088-0.110 X 0.110-
0.133. Anus-tail distance 0.110-0.165. The copulatory organ consists of' tw·o equal spicules, each 0.300 long with
f'ine transverse striations, heavily chitinized and barbed,
and the gubernaculum o.4oo also chitinized and with f'ine
transverse striations. Six pairs of' papillae are present,
three preanal and three postanal.
~------~--~------~--~------~-----~ 1-- g=_-_-=-=-= -= Only one species of' Dibulbiger has been described
to date •. The genus Dibulbiger was erected by Caballero
(1935) to receive a new s~ecies Dibulbiger longispiculis
from ~ montezumae in }:Iexico.
Q• caballeroi may be distinguished from Q. longi
spicul~ by its smaller size (f'emales: 5.80-7.20 compared ------to 12.65-12.76; males: 4.70-6.70 compared to 10.45),
paired number of buccal papillae, fewer number of anal
papillae (three pairs of' preanal and postanal papillae
compared to four pairs of' preanal and three pairs of' post-
anal papillae) and anatomical dif'ferences in the copulatory
organ (spic1ues finely striated, barbed, and shorter than
the gubernaculum compared to smooth spicules which are
longer than the gubernaculum).
This species has been named in honor of. the late
Dr. Eduardo Caballero y Caballero, in recognition of' his
contributions to the f'ield of parasitology. 17 Pseudaplectna cablei sp. nov.
PLATE IV Figs. 1, 2, and 3
Table III
Host: Ensatina £• eschscholtzii Gray (5/52), Aneides ltigubris (Hallow·ell) (1/7), A. :flavipunctatus ·Strauch ( 2/7)
Site: Intestine
Locality: Oakland
and :five males:
FEMALES: Slender, 2.00-4.00 in length; 'tvidth at the level
o:f the vulva 0.085-0.100; cuticle rather thin with :fine
transverse striations, . covered 'tvi th numerous "warts." The mouth is bound by six small iips, the esophagus is 0.171-
0~200; an esophageal bulb is present, 0.028-0.044 X 0.033- '""";---- o.o66; anus-tail distance 0.132-0.187. Ovaries two, vulva posterior to the middle o:f the body; the eggs are 30-38 X
52-66 covered with a smooth shell.
---~- MALES: Total length 2.70-3.35; maximum width 0.079-0.115. Cuticle and mouth parts as in the :fem.ales; the esophagus
is 0.095-0.185; the esophageal bulb is 0.030-0.034 X 0.044- o.oso. Anus-tail distance o.050-o.o8o. T'tvo spicules are present, 0.065-0.078; gubernaculum short and broad 0.030-
0.035 caudal allae well developed supported by seven pairs o:f postanal papillae.
Only one species o:f Eseudaplectana , ~· papillocauda
(1valton, 1940) Yamaguti, 1961 from the land form of the 18 salamander Desmoe;nathus f'uscus has been described to date.
Due to lack of f'emales in his collection, Walton (1940) did not erect a new genus to accomodate this parasite• Yamaguti ~--- q-·--·--~ (1961), however, established the genus Pseudaploctana
~~----
c;~~~ f'or this species. E;---
~ ~· cablei can be distinguished f'rom ~· papillocauda ~~- by its larger size (2.99 compared to 2.05; only males compared), f'ewer lips (six lips compared to three inner and
(0.065-0.078 compared to 0.120-0.128), and position of' the anal papillae; all seven pairs are postanal compared to two pairs of' preanal and f'ive pairs of' postanal papillae.
1val ton ( 1940) did not describe any body "warts" in .!!• papillocauda.
',I'his species has been named in honor of' Dr. Haymond :::;: ___ :____:_::-_-=--=--
M. Cable for his contributions to the f'ield of' parasitology.
Pseudaplectana waltoni sp. nov.
PLATE IV Figs. 4, 5, 6 ---- TABLE IV
Host: Batrachoceps ~· attenuatus (Eschscholtz) (4/250)
Site: Intestine
Locality: Santa Barbara
Description and measurements based on seven speci- mens; f'our females and three males:
FEMALES: Slender, 2.00-2.20 in length, width at the level of' the vulva o.o4o-0.070; cuticle thin with very f'ine 19 longitudinal striations. The body is studded •·dth numer-
ous 11 '\varts •" The mouth is bound by six small lips; the.
esophagus is 0.130-0.165 in length; an esophageal bulb is ~- ~~~ ~-~-· ~----'c-' present, 0.025-0.035 X 0.039-0.049; the intestine is tubular, §-=- - - ~-----
Q extends to near the posterior end; anus-tail distance 0.070- ~----- "' 0.170. The posterior end gradually tapers into a f'ine
pointed tail. Ovaries two, vulva slightly posterior.to the
middle of' the body; the eggs are 33-35 by 44-52, arranged
------!--~----~1--=n:---:a::---=s~i-=n~g=-=-l,-:;:e----=r:::-:o::-:w:-:-,----=a:-:n:-:d;,--::a-=r:-:e=--:.i=n---..tuhc:-e-=--=m=-=o=-=r=u:-:--,l-=a---=s:-:t.-:-.a=--=g:-:e:-.------~------'-'--'
}ULES: Smaller than the f'emales, 1.10-1.60 in length, max-
imum w·idth 0.039-0.061; cuticle and mouth parts are the same
as in the f'emales; the esophagus is 0.120-0.160 in length,
the esophageal bulb is 0.022-0.030 X 0.037-0.0J9; anus-tail
distance 0.045. Spicules two, 0.035 in length, caudal allae
well developed, supported by seven pairs of papillae, two ---- E preanal and five postanal.
E.• "ival toni may be distinguished from £• papillocauda ------by its smaller size (only males are compared; 1.100-1.600
compared to 2.050), fewer lips (six lips compared to six
outer and three inner lips), shorter spicules (0.0.35 comp
ared to 0.120-0.128), lack of a pharynx, and the presence
of warts all over the body.
£• waltoni dif'f'ers from £• cablei in size {male&;
1~100-1.600 compared to 2.700-3 • .350; females 2.00-2.200
compared to 2.0o-4.oo), shape of' the spicules (slightly
hamate compared to grooved spicules) and distribution of' 20 anal .papillae (two pairs of preanal and five pairs of postanal papillae compared to seven pairs of po~tanal pap- illae). ~ ------This species has been named in honor o£ the late A. c. Walton in recognition o£ his work in the field of E------~ parasitology. ---
Rhabdias sp.-
PLATE V Figs. 3, 4 ~r ------Host: Taricha torosa (Rathke) (1/6)
Site: Lungs
Locality: Sacramento valley (exact location unknown)
Description and measurements based on two specimens:
FE}~\LES:(parthenogenetic): The worms are brownish in color
10.60-10.90 in length; 0.470-0.500 in width at the level of
------the vu.lva. Cuticle rather thin anteriorly and thicker post- E- eriorly. The mouth is bound by six round lips, each one with a buccal papilla; the esophagus is 0.390-0.400 in
length, club shaped with a constriction at the point where ---~- it joins the intestine; ·the intestine is tubular, black in color. and s"\.,.ollen distally; anus-tail distance 0.690-0.700.
The posterior end gradually tapers into a narrow tail.
Caudal allae narrow; 12 pairs (?) of postanal papillae are present. Ovaries and uteri paired; eggs 55-66 X 118-132, are covered with ~ thin shell and contain fully developed embryos.
The genus Rhabdias was erected by Raillet in 1916.
This genus contains over 50 described species most of which 21 are parasites of' reptiles and amphibians; o:f these 50
species, f'ive have been reported from salamanders: Lehmann
(1954) reported Rhabdias sp. f'rom Taricha torosa and in 1960,
he reported Rhabd:i.as sp. f'rom Dicamptodon ensatus in Calif'- tl=== ornia; Chitwood (1933) described E• plethodontis :from ~ -- _- ----·--· Plethodon cinereus in Virginia; Wilkie (1930) described E•
tokioensis :from the Japanese ne,.,t Diemyctylus pyrrhogaster j. in Japan, and Baer (1930) described E• escheri f'rom the
o:f specimens (2) encountered in this study the worm is not
named or compared with others.
Thelandros magnavulvaris (Rankin, 1937) Schad, 1963
PLATE V Figs. 1, 2
Synonym: Oxyuris magnavulvaris_Rankin, 1937
----- Host: Taricha torosa (Rathke) (1/6) L Site: Rectum
Locality: Santa Barbara Description and measurements based on six specimens, ------five females and one male.
FE~lALES: 2 • .50-3.10 in length, 0.220-0.300 in lvidth at the.
level of' the vulva. The cuticle is thick with transvers
striations, the mouth is bound by three small lips, the
esophagus is o.4oo-0.510; an esophageal bulb is present,
0.990-0.103 X 0.110-0.120; excretory pore posterior to the
esophageal bulb. The intestine is tubular, extends to near
the posterior end, anal-tail distance 0.2.50-0.300, the
posterior end tapers·gradually into a pointed tail. Ovaries ~-¥-··------·--."------
22 two, tubular extend:anteriorly to the esophageal bulb, loop
over and around the esophagus and then continue posteriorly
as oviduct and uteri to near the anal region. The two uteri
meet in the center of the body cavity to form a muscular
Hf---;--::---- ovijector which opens to the outside through a large portrud- ~ -- ing vulva just anterior to midbody. The eggs are 77-90 X
33-39, have a polar plug and are in the 2-, 4-, or 8.-cell stage
w·hen laid •
------~------MJ:kl:,]~-i-Sma-l-1-e-J:"'----t-n-en-t-rre-----Iema 1 e s , r-;-ro-irr-1 ength, maxi'_m_u_m______t-~:~~--C
width 0.099, cuticle and mouth parts are the same as in the
females.· Esophagus 0.198, esophageal bulb 0.057 X 0.065,
caudal projection 0.070; spicule 0.050 in length.
Thelnndros minutus Read and Amreim, 1952
PLATE VI Figs. 1, 2
Host: Batrachoceps .!!• illenuatus (Eschscholtz) (121/250) E
Site: Rectum
Locality: Oakland
Description and measurements based on 20 specimens; ------
FEl-IALES: 1.78-4.20 in iength, 0.300-0.600 in width at the
level of' the vulva. Cuticle thick with transverse striations.
The mouth is bound by three small lips; the esophagus is
0.130-0.303 in length; an esophageal bulb is present, o.o66-
0.074 X 0 0 077-0.110; anus-tail distance 0.088-0.110; the:··
posterior end gradually tapers into a pointed tail. Ovaries
two, uteri two, .meet in the center of the body cavity to
form a muscular ovijector which opens to the outside through
the vulva somewhat anterior to the middle of.the body. The 23 eggs are operculated 33~77 X 80-144 and are in the 2-, 4-, or 8-cell stage when laid.
MALES: Smaller than the f'emalesf 0.650-0.900 in length; ~n ------maximum width o.o66-0.095; cuticle thinner than that of' the B f'emales and cuticular strations less marked; mouth parts as in the f'emales; esophagus 0.077-0.100 long, esophageal bulb
0.035-0.045 X 0.035-0.052~ The caudal projedtion (tail) me~sures 0.046-0.050 in length. One pair of preanal and two
Thelandros salamandrae Schad, 1960
PLATE VI Figs. 3, 4
Synonym: Oxzuris dubia o£ Lehmann (1954, 1960) nee. Leidy, 1856
--- Host: Ensatina ~6 eschscholtzii Gray (32/52), !• ~· ;:::---- ;_- sierrae Storer (11/17), Aneides lugubris (Hallo-
well) (3/7), !• flavipunctatus (Strauch) (4/7)
Site: Rectum
~--~ Locality: Oakland and Arnold .
Description based on 150, measurements on 23 speci- mens, 17 f'emales and six males:
FE~L\LES: 3.90-6.80 in length, 0.350-0.600 in width at the level of the vulva. The cuticle is thick with transverse striations. 'rhe mouth is bound by three small lips; esophagus o.4oo-0.650 in length, esophageal bulb present,. o.aoo-0.110 X 0.120-0.154; excretory pore posterior to the esophageal bulb; anus-tail distance o.4oo-0.520; the post- 24 erior end of' the worm gradually tapers into a pointed tail.
Ovaries two, tubular; uter~ meet in the center of the body
cavity to f'orm a muscular ovijector which opens to the out- ·~------['j··· side through the v-ulv-a just anterior to the middle of' the
h~--···_·_· body. The eggs are operculated, 40-66 X 88-118, and are in
the 2-, 4-, or 8-cell stage when laid.
HALES: Smaller than the f'ernales, 0.500-1.100 in length,
maximum width 0. 055-0. 130; cuticle thinner than tha·t of' the
~-----.f' ema-l-e~-anG!.-s-u-t-::ke-u-l-a-r-s-"b-r:J:.a-t:J:.or.rs-J:-e-s-s-:ta:rke-d-.~Tne mou tnhtl:aass~----~~===
three bilobed lips; esophagus 0•077-0.176 in length; the
esophageal bulb 0.037-0.066 X 0.039-0.066.; The caudal pro-
jection (tail) measures 0.079-0.090. One pair of preanal - - --- and two pairs of' postanal papillae are present. A single
spi~ule is present 0.037-0.060~
The genus Thelandros lvedl, 1862 is represented
today by at least 50 species most of' ,,hich are intestinal
parasites of' reptiles with the exception of three species ---- from salamanders: !• minutus, Read and Amreim, 1952 f'rom
Batrachoce~s ~· attenuatus in Calif'ornia; !• magnavulvaris
(Rankin, 1937) Schad, 1963 f'rom Desmognathu.s f'uscus, ll•
ochrophaeus carolinensis, Q .. P..!l~' Q. quadrimaculatus,
Eurycea bislineata wilderae, ~· guttolineata, Plethodon
cinereus, ~· glutinosus, Diemyctylus viridescens, in N.
Carolina, Aneides f'lavipunctatus in Calif'orniat !• salaman-
~' Schad, 1960 f'rom Aneides hardii in New Hexico, !•
lugubris, !• f'lavipunctatus, ~· ~· attenuatus in California, 25
A• ferreus, Ensatina ~· oregonensis in Oregon, !• ~· eschscholtzii in California, Plethodon elongatus, f•
stormi, ~· vehiculum in Oregon and ~· neomexicanus. in New
Hexico.. '!1vo species of Thelandros have been described from
frogs: I• espectatus Texeira and Freitas, 1962 from ~ spinulosus limensis in Peru and I• oswaldocruzi Travassos, 1925 from Hyla mesouhaea in Brazil.
! ______;;:;; ·-- -- ~- "--··-·
:::
-----
r:--=---:.
---- SUMMARY
t-~ Between September 1975 and April 1977, 340 sala- Dg h manders belonging to seven species, Aneides f'lavipunctatus r===
!··c . (7), !• lugubris (7), Batrachoceps attenuatus attenuatus !t-'-==-== ( 250), :E~nsa tina eschschol tzii eschschol tzii (52), .fd• .!•
sierrae (17), Taricha torosa (6), and Dicamptodon ensatus
( 1), w·ere examined f'or helminths and f'ound to harbor 10
Cestoda: Distoichometra ensatinae sp. nov. f'rom Ensatina ~· eschscholtzii and ,:m. !!• sierrae; and Baerietta diana {Helf'er,
1948) Douglas, 1958 f'rom Batrachoceps ~· attenuatus.
Nematoda: Dibulbiger caballeroi sp. nov. f'rom Dicamptodon
ensatus; Pseudanlectana cablei sp. nov. f'rom Ensatina e. ------. - eschscholtzii, ~eides f'lavipunctatus, and !• Jugubris;
Pseudaplectana waltoni sp. nov. f'rom Batrachoceps ~· atten•
~~; .:...C:.l.o~siZ.jmu,lolojou.cwe-...r~oo.~oocdoo'""'i~d...... ,e,loil,s duku (Roll, 1928) Travassos, 1931 f'rom,Taricha torosa; Rhabdias sp. f'rom Taricha torosa;
'!,helandros salamandrae (Lehmann, 1954) Schad, 1960 f'rom·
Ensatina ~· eschscholtzii, Aneides f'lavipunctatus and !• lugubris; 1> magnavulvaris (Rankin, 1937) Schad, 1963 f'rom
Taricha taros~; and 1"> minu~ Read and Amreim, 1952 f'rom
Batrachoceps ~· attenuatus.
~------A new host record is established f'or Thelandros salamandrae from Ensatina .!• sierrae. A host-parasite list covering '\vork in California, Oregon, and British Columbia is also included.
26 HOST-PARASITE LIST
Family Plethodontidae
Aneides flavipunctatus, the black s~lamander 1 Thelandros salamandrae Schad, 1960 2 Thelanclros magnayulyarjs (Rankin, 1937) Schad, 1960 3- Pseudaplectana cablei sp. nov.
Aneides lu,g-ubris, the arboreal salamander 1 Baerietta interauej des {Helfer, 19l.j.3) Yamaguti, 1------~129~59~----~------~------~=~ 2 Thelandros salamandrae Schad, 1960 3 Pseudaplectana cablei sp. nov.
Batrachocegs ~· attenuatus' 1 Baerietta diana (Helfer, 1948) Douglas, 1958 2 Thelandros minutus Read and Amreim, 1952 3 Thelandros salamandrae Schad, 1960 4 Pseudaplectana waltoni sp. nov.
Ensatina e. sierrae 1 Thelandros salamandr~ Schad, 1960 2 Distoichometra ensatinae sp. nov.
Ensatina e. eschscholtzii, the monterey salamander · +Brach;coelium salamandrae (FrHlich, 1789) Stiles and Hassell, 1898 2 Baerietta diana (Helfer, 1948) Douglas, 1958 3 Thelandros salaman<;lrae Schad, 1960 4 Pseudanlectana cablei sp. nov. 5 Distoichometra ensatinae sp. nov •.
----
': _____ :____-:______::______Species reported f'rom Oregon are indicated by + sign, those from British Columbia by ++; all others are known in Cali fornia.
27 --~~------
28 HOST-PARASITE LIST (CONTINUED) Family Ambystomidae !!!!.bystoma gracile 1 "Megalodiscus america~ Chandler, 192.3 2++r·iegalodiscus mic1:2,12hagus Ingles, 1936 3++Cephalouterina dicamptodoni Senger and Macy, 195.3 4++Cyatocephalus truncatus Ballas, 1781 Ambystoma macrodactylum, the long-toed salamander 1 ++}legalodiscus microphaeys Ingles, 1936
2 Megalodiscus americanus Chandler, 1923 .=~~~
3 +Negalodiscus microphagus Ingles, 1936 ~-~~ 4 +Phvlodistomum singulare Lynch, 1936 5 +Euryhelmis pacif'icus Senger and Hacy, 1952 6 +Cephalouterina dicamptodonti Senger and Macy, 1953 7 Thelandros salamandrae Schad, 1960 ~--~~~~~~~~~~--< HhaDaJ.as sp. <=!" ~~--- 9 Gyrodactylus ensatus Mizelle et al., 1968 10 Dibulbiger caballero! sp. nov.
Family Salamandridae
Taricha torosa, the California Newt 1 +Halipegtis occidualis Staf'f'ord, 190~ 2 1·!et";alodiscus americanus Chandler 192.3 J Cosmocercoides dukae (Holl, 1928) Travassos, 1831 4 Bothriocephalus rartls Thomas, 1937 5 ThE,landros ma~navulvaris (Hankin, 19.37) Schad, 19bJ--· . 6 Rhabdias sp. of Lehmann (1954) 7 Capillaria sp. 8 Negalodiscus sp. 9 Rhabdias sp. (this paper) Taricha torosa sierrae 1 Thelandros salamandrae Schad, 1960
Taricha granulosa, the rough skinned newt · 1 Megalodiscu! americanus Chandler, 192.3 2++Cephaloutherina dicamptodonti Senger and Macy, 1953 J++}·legalodiscu~ microphagus Ingles, 19.36 4 +Brachycoelium salamandrae (Frtllich, 1789) Stiles and Hassell, 1898
----
Taricha sp. 1 Megalodiscus arnericanus Chandler, 1923 ·-
LITERATURE CITED !=;-- ,_,u------=-= Baer, J.G. 1930. Deux helminths nouveaux, parasites de Uraeotyphlus oxyurus (Gray), Gymniphione de l'Inde meridionale. Rev. Siusse Zool., 37(2):43-52. ·
Caballero, y C~ballero Eduardo~ 1935. Nematodes parasites de los batracios de Mexico. III. Cuarta contribucion al :~------~c-o~n-o-c~1miento de la parasi tologia de Hana mont-e-zum13:"e.-.------::==_--'--_--_--': I. Ann. Inst. Biol. Univ.·Nac. Hexic'O':""b'(2):103-117. ·
Chitwood, B.G. 1933. A revised classification of the Nematoda Jour. Parasitol. 20(2):131.
Dickey, L.B. 1921. A new amphibian cestode. Jour. Parasitol. 7, 128-136. -----
Douglas, L~T. 1957. Fertilization, maturation, and early cleavage in Baerietta desmognathi sp. nov. (Nematotae niid.ae). Jour. Parasitol. 4J(supp.):24. (Abst.)
• 1958. The taxonomy of Nematotaeniid cestodes. Jour. ------~}~~a-r-,-asitol. 44:261-273.
Efford, I.E., and K. Tsumura. 1969. Observation on the Biol ogy of' the trematode t-Jegalodiscus microphagus in amphib ians from Marion Lake, British Columbia. Amer. Midland
Naturalist, Vol. 82(1):197-203. - -
Helf'er, J.H. 1948. Two ne,., cestodes from salamanders. Trans. Amer. Hicr. Soc. 67:359-361.
Holl, F.J-. 1928. ~-10 new nematode parasites. J.E. :t-1itchell Scient. Soc~ 43:184-186.
HsU, H.F. 1935. Contribution a• 1 1 etude des cestodes de Chine. Rev. Suisse Zool. 42:477-570.
Johnston, B.H. 1962. Nematodes parasites of salamanders. Hanuscript Thesis, library, University of the Pacif'ic. Stockton, Ca.
Lawler~ H.J. 1939. A new cestode Cylindrotaenia guadrijugosa n.. sp. from ~ pipiens, '\vi th a key to Nema totaeniidae. Trans. Amer. :t>Iicr. Soc. 58:73-77. ------
29 30 Lehmann, D.L., 1954. Some helminths of the ·1vest Coast urodelos. Jour. Parasitol. 40(2):231.
• 1956. Some Helminths of Oregon urodelese Jour. ___P_a_r_a-si tol. 42:25. . ~- 11~~~~ ------• 1960. Some parasites of Central California amphi !'!c:-=·-.--. bians. Ibid. 45:10.
Lynch, J.E. 1936. Phyllodistomum singulare n. sp. a trema tode from the urinary bladder of Dicamptodon ensatus Eschscholtz, with notes on related species. Jour. Para sitol. 22:42-47.
Macy, R.W., W.A. Cook, and W.R. Demott. 1960. Studies on the life cycle of Halipegus occidualis Stafford, 1905 ______1------~·( '.Pr-ema:t-o·da-:-H-e-miur:td-a-1-)~.~N orthwe s t-sci-;-J-4: 1~1 -.------~-~----·'C.-=
:r.-rcCauley, J.E., and I. Pratt. 1959. The paramphistome ~egalodiscus microphagus Ingles, 1936 from the giant salamander, Dicamntodon ensatus Eschscholtz from Oregon. Jour. Parasitol. 45:614.
Nizelle, J.D., D.c. Kristy, and R.B. Burry. 1968. Studies ---- on monogenetic trematodes. XLI. Gvrodactylus ensatus sp. n. The first species of' the genus described from Amphibia. Jour. ~arasitol. 54:281-282.
, , and H.D. McDougal. 1969. Studies on mono------genetic trematodes. XLII. New species of Gyrodactylu! from Amphibia. Jour. Para~itol. 55(4):740-741. Ogden, C.D. 1966. On some nematodes -from reptiles, mainly from Ceylon~ Jour. Helminth. 15{1):81-90.
Read, C.P. and u. Amreim. 1952. Some new nematodes from ---- . -~-- -- · Southern California~ Jour. Parasitol. 38(5):379-384.
Senger, c.M. and R.1v. r.-racy. 1952. Helminths of Northwest mammals. Part III. The description of Euryhelmis paci ficus n. sp. and notes on its life cycle. Jour. Para sitol. 38:481-486.
---::---' • 1953. A new· digenetic trematode (Ceph- alouterina dicamptodoni n.g. and n. sp.:pleurogenetinal) from the giant salamander. Jour~ Parasitol. 39:353-355. ------·
Ulmer, J.H., and H.A. James .. 1976. Nematotaenoides ranae gen. et sp. n. {Cyclophyllidea). from the leopard frog (Rana pipiens) in Iot-la. Proc. Helminth. Soc. 1vashington 4JT2):185-199.
Walton, A.C. 1940. Some nematodes from Tennessee Amphibia. ----- Jour. Tennessee Acad. Sc. 15(4):402-405. .31
1ofardle, R.A., and J .A. McLeod. 1952. ~ zoology .£! Tape'lwrm~ The University of' Ninnesota Press, Minneapolis. " ------~--' and s. Radinovsky. 1974. Advances in the of Taneworms, 1950-1970. University of Ninnes<;ta ZOol~gy "~-- Press, Minneapolis. · !!c--~ I" Wilkie, J.s. 1930. Some parasitic nematodes f'rom Japanese w~~~ Amphib.ia. Ann. Mag. Nat. Hist. 6(10):606-614. ~··-·-- Yarnaguti, s. 1959. Systema Helminthum. Vol. ti. The cestodes of' vertebrates. Interscience, NeW fork.
---~-··· 1961. Systema Helmin~. Part II. ~nematodes of' vertebrates. Vol. III. Interscience, New York. j------~----~------~------~~~
------
------
------[i_D_!_'}I_I_lm~ 1[1j .l! ~~~--~ '!Ill
TABLE I
Measurements on Distoichometra ensatinae sp1 .• nov.
Minimum X n Maximum I s
Total length 14 17.00 4o.oo 29.92 7.160 Scolex width •• 0.200 0.450 0.321 0.063 Diameter of suckers 13 . 0.090 . 0.120 .o .106 0.008 Length of neck 14 4.so. 6.80 . 5.)8 0.730 Ovary 13 0 • 0 55 X 0 • 0 60 0.088 X 0.114 o.o641X o.o87 0.011 X 0.016
Vitelline gland " 0.044 X 0.055 0.066 X 0.100 0.057 X 0.076 0.090 X 0.015 Testes 28 0.055 X 0.090 0.100 X 0.200 0.073 X 0.140 0.012 X 0.027 Cirrus sac 14 0.033 X 0.042 0.050 X 0.120 0.039 X 0.079 o.oo6 x o.o17
Last segment " 0.200 X 0.350 0.500 X 1.210 0.)16 X 0.717 o.o84 x 0.225 Paruterine organs 63 _0.050 X 0.050 0.110 X 0~120 0.082 ~ 0.089 0.010 X 0.011 Par11terine capsules 1 i 24 X 44 33 X 66 30.10 IX 51.50 4.340 X .8.410
Measurements in nun ·:exc·.ept for paruterine capsules
l.oJ N
,.J_J .E IIi:·: ,J i 'i]i!il :: II 1.···, I""' 0 I ~·:.".'11'1. I In, . . I i '11 •!"I
: ! I II' I !I 1111, I !1111 1. I I I II n ,_ _t,_,_I~ILH._l~-~"---~ '' "''''
·TABLE II
Measurements on Di bulbiger caballeroi sp. no~ir, I
Females n Minimum Maximum X s
Total length 6 5.80 7.20 6.81 I 0.508 k'lidth through 0.187 0.046 vulva " 0.143 0.275 o.o66 o.ooa Pharynx " 0.060 0.075
Esophagus It 0.720 0.810 0.781 0.024
Prebulb " 0.077 0.088 0.083 o.oo4
Esophageal bulb It 0.099 X 0.110 0.120 X 0.132 0.11} X ]0.123 0.008 X 0.008
Anus-tail 0.265 0.012 distance · " 0.242 0.275 2.480 X 3.780 Egg size " 33 X 44 38 X 55 35 X 48
Measurements in mm except for eggs
w w
::.J;,, .. 1~:11 .: .;: ,- I : 1., II :.If: I I I I I I I
I I r ~ '''''."""I I .___l!_ll__,_l._,_lm_H_l,__'_!_L_]I.J...""---'--'oi.;:;l---lllll·l.l.:.""".o..;.'-...;..=='-"'------A---.._-.11!..---...,;------'-----
TABLE II (CONTINUED)
Males n Minimum Maximum· X I s Total length 6 4. 70 6.70 .5.91 0.67.5
. 0.138 0.280 0.203 0.043 Maxi mum width. " 0.061 0.00.5 Pharynx " 0.0.5.5 0.070
Esoph~.gus .. 0.665 0.740 0.7)1 0.030
Prebulb II 0.067 0.081 0.072 o.oo4
Esophageal bulb II 0.088 X 0.110 0.110 X 0.133 0.10.5 X o.t17 ·o ~,o11 x o .011
Anus-tail· distance .. 0.110 0 .• 16.5 0.147 I 0.019 Spicules ·1 0.)00 - 0.300 0.400 Gubernaculum " 0.400 -
Measurements in mm
w ~
1.1·.: 1 ,.1: ,J, I~ I I I lll " il l]'~ . .' "'''"0"1""""' : ': ·i' : i ' ~r ,,. I ·!:.ji i. il'.:~~ r .·. 1.11!:: ;li': i . I I . ! I i •. '1'1 I I • I I Ill !I . I . '"'I""
TABLE III
·Measurements on Pseudanlec tana c::ablei sp. n~v •
Females n Minimum Maximum X I s .I Total length 8 2.00 4.00 3.00 0.794
Width through vulva 7 0.085 0.100 0.092 o.oo6
Esophagus · 4 0.171 0.200 p.189 0.013
Esophageal bulb " o.o28 x o~o44 0.033 X Q.066 0 • 0 3 5 ~: 0 • 0 60 0.003 X 0.011 Anus-tail distance 8 0.132 0.187 0.153 I 0.019
Egg size 5 JO X 55 38 X 66 JJ x 6o: 2 • 88q. X 4 • .50Q:
Measurements in mm except·.fcrr ~ggs
w V1
1 0 1:.1.! : J1l!ilr ! I :1 :c::·~-~---'II':. Im,_· ·~·--·1·1.lllll::e ... CIL .. ,L~il ... : '" .. , ·;r • : ..• 1 ·t. Ill • •. ~~ 'I. 11·1r 1r · : I . I :, :., 'I, ''II 111111 I 'II I l~li· •id I• "' d I II
TABLE III (CONTINUED) ·I I Maximum X s l'vla~es n . Minimum I Total length 5 2. 70 3·35 2.99 0.255 ,, Maxi mum width If 0.079 0.115 0.100 0.016
Esophagus 4 0.095 0.185 0.151 0.039
Esophageal_'bulb 2 0.030 X 0.044 0.034 X 0.050 0.032 X 0.047 o.oo2 x o.oo4
Anus-tail distance 5 0.050 0.080 o.o66 0.011
Spicules If 0.065 0.078 0.073 0.005
Gubernaculum 2 0.030 0.035 0.032 0.003
Measurements in'mm
w 0\
L ... I:I :II'LG 1.1 I I I J, I "''''"' ' ' :l::r .. ·:i '· ·1· :' I''I I' :,' , ..li ,I I I .•. ,i I I'·I ·. '·•, ,:1 1: . ,; , 'II"' .. '~"'Ir"'D"'i: I I: :I; :I "' 'I II; :'I il:l :I '!"''" I ,1, I I ,I, I I 'I' I II l!,n 1"1m J.,, 1L____u_c__ _, __,__J;::·"••••IIIIIm""'"""'-...._-...:.=""-_.. ______....._ _ _,__.lli._~------~------
'TABLE IV Measurements on Pseudaplectana waltoni sp. rov.
Females n Minimum Maximum X s
Total length 4 2.00 2.20 2.10 0.082
\vidth through · vulva · .. 0.040 0.070 0.060 0.014
Esophagus ... 0.130 0.165 0.150 0.017
Esophageal bulb " 0.025X 0.039 0 • 0 3 5 X 0 • 049 0.029 X 0.042 0 • 00 5 X 0 • 00 5
Anus-tail distance 3 0.070 0.170 .0.118 0.046
Egg size II 33 X 34 .· 35 X 52 43 X 481 :3.460 X 7.760
Measurements in mm except ·:ror eggs
...... w
••.1. I I ' i~ n; 'i " .. , . ·""'""'1~'""·'CLC." '" •· ...... I .I rI '':1"1 i ~~, ·11 111 : ,, I . : ! .• I I .,, , . 'I : I' I r , I ·r..... I 1,:: :, :1~1 :1 I I ! lll ! I!:·,:1 ·il I ' ! r1 I I I II I, I I ,~,_J,_dlli_j~J "'"
TABLE IV (CONTINUED)
Males n Minimum Maximum X s
Total length 3 1.10 1.60 1.JJ 0.252
Maximum width " 0.0)9 0.061 0.048 0.012
Esophagus " 0.125. 0.160 0.146 0.019
Esophageal bulb " 0.022 X 0.037 0.0)0 X 0.0)9 0.026 X ~).0)7 o.oo4 x o.oot
Anus-tail distance 1 - 0.045 0.045 Spicules " - 0.0)5 0.0)5 Gubernaculum " - O.OJO 0.0)0
Measurements in mm
·w 00
:I.JH.: •• i:J..,.il. :·. 1 ... , ll I ,,.iJ I. pu:r. ~ r:: II .. .:r. :·
[, I l'll'fl ~1"'1"1~ I ------·---··-·-·-··-····-· ------·-----··-·-·-·-----~-
39
PLATE I 6 ,~-- -·. == !-!=-=--== ~
Distoichometra ensatinae sp. nov.
~------~------~ ~-----~~
[j ------Fig. 1 Scolex
Fig. 2, J, and 4 Mature Proglottids
----- Fig. 5 Gravid Proglottid
'-:-=-----=--- ~---
-----
------
PLATE I
l I".. 3 3 J
Fig. l
l ~.. 3 j Fig. 3
l0.. 3 j !'1~~.~ PLATE II .~~.~
~=- ---=-== ~·~~~ Baerietta diana (Helfer, 1948) Douglas, 1958 6~--
Fig. 1 Scolex
------i===:~~
Fig. 4 Gravid proglottid
------
----~ .·"
PLATE II
Er. E U) 0 .... ·.. : .. .· .. · t.. . . [ . . ~ ... ~:: ...... ·. ·.:: ·.' ·. ·: ~ Fig. 1 .· ... a-:·. . . : ...... : : .. ·. ·.. ·. : :. ~ : ... ·. :. ::.: .·::.:. ·:.::.:::.:~.:. .. ·... ~ ,·
I E e (') 0
F;g.-L J· Fi9. 4)
~- -- - ['j---~~ !!!~ .. ~----'-,
L:_-__ PLATE III E- -____ c_:__ c ______:_:_ __:_ ___
Cosmocercoides dukae (Hall, 1928) Travassos, 1931
,;------~------~--~~-~~--
Fig. 1 Female anterior end
Fig. 2 ~1ale posterior end
~--- Fig. 3 Female posterior end
:;:;:---::-::------
Dibulbi!er caballeroi sp. nov.
Fig. 4 'Hale posterior end
Fig. 5 Female anterior end
Fig. 6 Female posterior end
----
------PLATE Ill !!·c~.~- f"··-
§ ______~_ ~---- w :..~ ---
--- -- ~------~~==~~- E '-';·~~~= E ~ J
-- ---
Fi 9. 1
---· ----_------··-·--
l E E E E <') 2 0 J J
fi9. 6 PLATE IV
Pseudaplectana cablei sp. nov.
1------~------~~-P,~~~== Fig. 1 Female anterior end
Fig. 2 Male posterior end
Fig. 3 Female posterior end
l-
~------
------
p_seud~lectana ,val toni sp. nov.
Fig. 4 Female posterior end
------·------·-· Fig. 5 Male posterior end
Fig. 6 Female anterior end
------PLATE IV
I ~------:------~·u<3 ------
Fi9. 2 lE E
-0
J = fl 9.1
· ..... ·.' . ·.·.·... . · .. ·
Fig. 6
, 47 ...
,.;:::;
~----
PLATE V 8 ~--=-=:~ ------
Thelandros magnavulvaris (Rankin, 1937) Schad, 1963
;~------~------~;"··· .... ~~
Fig. 1 Female (entire worm)
Fig. 2 Male (entire worm)
-----
~habdias sp.
Fig. Posterior end
Fig. 4 Anterior end
-----
-- -·-----
----- PLATE. V
E:1=:---==-;-:-----:;-;--:c !-~~ ,~.~;:;:' R------g______
"'~
lz 3 3
J o::---.=-----
3z 3 J
Fi9. 3
---··-- ~-~~-~ ··~·- ~------·--··
PLATE VI ~ -
Thelandros minutus Read and Amreim, 1951
~------·~~~ ...... ~-~~~- 9-- Fig. 1 Male (entire worm)
Fig. 2 Female (entire worm)
Thelandros salamandrae Schad, 1960
!::------
Fig. Female (entire ~orm)
Fig. 4 Male (entire worm)
------~------
------
- ~------=--
------
---- PLATE VI·
~ ~------
.. ___
~-:---:---_---::._:.-~=---: ;------~~---
------
------
-: /. Fi 9. -' ·:: f i 9. 1 .
\ ... . ::.
Fig. 3
------