© Institute of Parasitology, Biology Centre CAS = 9*&G G% **; doi: &* &88&&H 9*&G **; http://folia.paru.cas.cz
Review A synoptic review of Promonobothrium Mackiewicz, 1968 (Cestoda: Caryophyllidea), parasites of suckers (Catostomidae) in North America, with description of two new species
Mikuláš Oros& 9, Jan Brabec9, Roman Kuchta9, Anindo Choudhury3 and Tomáš Scholz9
1 " # M " # 5 6 < 9 = N " 5 # D3 " 5 6 < 3 D " Q Q 6 D R N
Abstract: Monozoic cestodes of the recently amended genus Promonobothrium V # W &EG; X Y @ # X Y Q " W W [ 6 > " [ X ?VY ?" type and voucher specimens from museum collections and newly collected material of most species indicated the following valid nom@ inal species: Promonobothrium minytremi V # W &EG; X Y< P. ingens X\ &E9'Y< P. hunteri XV # W &EG%Y< P. ulmeri X V # W &EGGY< P. fossae XR &E'8Y P. mackiewiczi XR &E'8Y Rogersus R &E;* with its only species R. rogersi is transferred to Promonobothrium based on morphological and molecular data. Promonobothrium cur- rani sp. n. and P. papiliovarium sp. n. are described from Ictiobus bubalus X5 [ ] Y Ictiobus niger X5 [ ] Y Erimyzon oblongus (Mitchill), respectively. The newly described species can be distinguished from the other congeners by the morphology of the > " " " " V > 6 ] 6 6 5Q X DQ DQ Y [ " # [ Promonobothrium based on morphological characteristics is provided. Keywords: ? [ " > W Q 5 [ #
Systematic research on caryophyllidean cestodes in XM "@\ " 9*&* # " 9*&& Q 6 W \ R X&E%*Y > 6 9*&9Y 6 @ monograph of the group and reached its highest intensi@ X 6# 9*&8 "@ &EG*R &E'*R W " " 9*&8 \ " 9*&+Y _ V # W V _ N 5 _ ? \ R @ Based on morphological and molecular data, Scholz liams described a number of caryophyllidean species and X9*&+Y [" Q " proposed several new genera (see references in Hoffman placed in Monobothrium D &;G% &EEEY V # W 6 @ monotypic Promonobothrium V # W &EG; \ W@ tant contributions to the current knowledge of this enig@ " > 6 6 W XV # W &E'9 &E;& > &EE8 9**%Y 6 study on amended Promonobothrium are presented and &E;*R X # [ " @ 9*&8Y W 6 3 @ tion, two new species of Promonobothrium are described X\ 9*&8Y from smallmouth buffalo Ictiobus bubalus X5 [ ] Y This long period, i.e. almost three decades, of neglect and black buffalo Ictiobus niger X5 [ ] Y V @ 6 Q 6 pi, and from eastern creek chubsucker Erimyzon oblongus caryophyllidean cestodes represent an interesting model XV Y Q N " @ for studies of unusual molecular and cytogenetic phenome@ relationships of species of Promonobothrium are assessed.
" # %& %'* *+ 5 # D3 " 5 6 789* %; +% &* %+&< = > 789* %; +% &* %;;< ?@ B Zoobank number for article: 6 # 6 =&+D+DD@8 +*@8''D@E;8=@; %*&&' &%
This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. &* &88&&H 9*&G **; Oros et al.: Synopsis of Promonobothrium (Cestoda)
Table 1. List of specimens of species of Promonobothrium V # W &EG; ]
# Specimen Host XN Y Voucher DQ H DQ Promonobothrium currani sp. n. Ictiobus niger X5 [ ] Y Chotard Lake, Mississippi @8&* MNGG++G%HMNGG++'8 Promonobothrium currani sp. n. Ictiobus bubalus X5 [ ] Y Chotard Lake, Mississippi N 98+ MNGG++'*HMNGG++;& Promonobothrium hunteri Hypentelium nigricans R W 5 " R @899 MNGG++G8HMNGG++'+ XV # W &EG%Y (Lesueur) Promonobothrium hunteri Moxostoma poecilurum Jordan 5 " V @8GG MNGG++G+HMNGG++'G Catostomus commersonii D # # R D &%+H*E@ MNGG++G&HMNGG++'9 Promonobothrium hunteri (Lacepède) Promonobothrium ingens Moxostoma anisurum R 5 " R D &&+H*E@ MNGG++G*HMNGG++'& X\ &E9'Y X5 [ ] Y Promonobothrium minytremi Minytrema melanops R 5 " R @8*G MNGG++G9HMNGG++'% V # W &EG; X5 [ ] Y Promonobothrium minytremi Minytrema melanops 5 " V N &';6 MNGG++GGHMNGG++'' Promonobothrium minytremi Minytrema melanops 5 " V N 9*E MNGG++G;HMNGG++;* Promonobothrium minytremi Minytrema melanops 5 " V N 9*E @ MNGG++GEHMNGG++'E Promonobothrium rogersi 5 " V N &E& MNGG++G'HMNGG++'; XR &E;*Y 6 Carpiodes velifer X5 [ ] Y @Q M ] D Q = 3 HHWWW W 6 H >< N Q D Q M ] [ D
MATERIALS AND METHODS " W " X9**EY Specimens studied " W 3 X9**;Y Material of the following species was studied: Promono- Deposition of specimens bothrium minytremi V # W &EG; X Y< P. in- Specimens studied are deposited in the following collections: gens X\ &E9'Y< P. hunteri XV # W &EG%Y< P. ulmeri the Helminthological Collection of the Institute of Parasitology, X V # W &EGGY< P. fossae XR &E'8Y< 5 # D@ P. mackiewiczi XR &E'8Y< Rogersus rogersi R 3 " 5 6 X Y \ R V @ &E;* Q W W [ > 6 N " Q 6 # V N X 5 M V x Y 6 Q X\RVY Q \ V NM XQ\VNMY V > R Q V Q \ XQVQ\Y N 9*&* 9*&9 9*&% R D N Morphological evaluation Molecular data generation and analyses Tapeworms collected recently by the present authors were pro@ @ W W [> 6 6 x X9*&*Y \ EG @ X live tapeworms were isolated from the host intestine and rinsed W [> W 6 6 " " in saline. Hot (almost boiling) 4% neutral buffered formaldehyde " Y 6 ] > DQ solution (= 10% buffered formalin) was added into the beaker or X 6 & Y DQ " W W 6 9M% W # @ 5 [ 6 5Q X D@ worms were transferred to 70% ethanol before further processing. Q DQ Y ] W 6 W W V R @ X9*&9Y ] W 6 @ mine solution, dehydrated through graded ethanol series, cleared " ' X HHWWW H M 9*&9Y in clove oil (eugenol) and mounted in Canada balsam as perma@ @ ?@ Q @ nent preparations. Line drawings were made using a drawing at@ V == XM 9*&%Y DV +*** W 6 W > Q # V W # 6 ] % ; # X Y "@ Phylogenetic relationships were estimated from a concatenat@ en in micrometres unless otherwise indicated. Pieces of worms > # XVY @ 6 W 6 [ W > @ X Y [ " &9M&+ 3 W R R > @ were selected using the small sample size corrected Akaike in@ stained with 1% eosin B following recently updated protocols = & & * X 9*&9Y Xx 9*&*Y The best ML estimate was obtained from 100 searches in the pro@ Several specimens were prepared for scanning electron mi@ 5 " 9 *& X W # 9**G< = 8 X ?VY W 6 x [ Y V 6 W 6 X9*&*Y _ _ V G+&* ? V @ + &** 6 5 ?V W 6 W V " % 9 X5 ] from Erimyzon oblongus W # W @ @ 9*&9Y 8 V % 8 9* @ ed specimens that were remounted from Canada balsam. Scan@ lion generations, sampling tree topologies every 1 000th genera@ ning electron micrographs of all but two species (P. fossae and tion, and using the same partitioning scheme and model as in the P. mackiewiczi) of Promonobothrium W ?V 6 @ V @ W * 9+
= 9*&G G% **; 9 &8 &* &88&&H 9*&G **; Oros et al.: Synopsis of Promonobothrium (Cestoda)
Fig. 1. Scanning electron micrographs of species of Promonobothrium X M_ M > M M M [ Y A M Promonobothrium minytremi V # W &EG; Minytrema melanops X5 [ ] Y< B M P. ingens X\ &E9'Y Carpiodes cyprinus X Y< C M P. hunteri XV # W &EG%Y Hypentelium nigricans X Y< D M P. ulmeri (Calentine et V # W &EGGY Minytrema melanops X 9*&+Y< E, F, K, L M P. currani sp. n. from Ictiobus bubalus X5 [ @ ] Y< G, H M P. rogersi XR &E;*Y 6 Carpiodes velifer X5 [ ] Y< I, J M P. papiliovarium sp. n. from Erimyzon oblongus (Mitchill).
RESULTS Promonobothrium V # W &EG; Promonobothrium W E Synonyms: Monobothrium D &;G% partim (spe@ P. minytremi (type species), P. ingens, P. hunteri, P. ul- Q Y Rogersus R &E;* meri, P. fossae, P. mackiewiczi, P. rogersi (for Rogersus (new synonym) rogersi) and two new species described below. A synop@ Type species: Promonobothrium minytremi Mackiewicz, sis of these species (listed chronologically after type spe@ &EG; Y W > [ " hosts, life cycle and geographical distribution is provided Amended generic diagnosis of Promonobothrium: 6 W W W > Caryophyllidea: Caryophyllaeidae. Body elongated to ?V " W [ @
= 9*&G G% **; Page 3 of 14 &* &88&&H 9*&G **; Oros et al.: Synopsis of Promonobothrium (Cestoda) itriches. Inner longitudinal musculature well developed. " " X[ ; V # W > @ &EG;Y 5 P. minytremi from W " > Q # @ M. melanops are identical with those described by Mack@ @ " @ W X&EG;Y ?> " x" \@ 6 \ @ 6 Promonobothrium ingens X\ &E9'Y Scholz, Vitelline follicles medullary, lateral and median. Postovari@ x 6 R 6 9*&+ 6 N = & @ V N Synonym: Monobothrium ingens \ &E9' ? 6 T y p e h o s t : Bigmouth buffalo Ictiobus cyprinellus* (Valenci@ # X Y Q ennes) (Ictiobinae). V # W X&EG; &EE8Y Additional hosts: Quillback Carpiodes cyprinus* (Lesu@ X9*&+Y Y X 6 Y X W Y R # Catostomus commersonii (Lacepède) (Catostominae), Smallmouth buffalo Remarks. Promonobothrium was erected by Mackie@ Ictiobus bubalus X5 [ ] Y X 6 Y " W X&EG;Y P. minytremi Moxostoma anisurum X5 [ ] Y X Y X W from Minytrema melanops X5 [ ] Y 5 " host record). x# N @ # V N X 9*&+Y P. minytremi appears in a mono@ D 6 N M V Q W # x x# @ phyletic lineage comprising caryophyllidean species from R Q 6 V > x " in Monobothrium. Therefore, these species have been I. cyprinellus X 6 ;9& ; # W Y N transferred to the originally monotypic Promonobothrium. X @+;%Y Moreover, Rogersus R &E;* W Q W x is transferred to Promonobothrium as a new combination, C. cyprinus XD ;EH*E M Y # R 6 and two new species of Promonobothrium are described R < M. anisurum XD &&+H*EY R herein, thus increasing the total number of species in the 5 " R N 9**E X @+;%Y genus to nine. \ X&E9' &E%*Y X&E89Y X&EG'Y R X&E';Y Promonobothrium minytremi V # W &EG; M Remarks. \ X&E9'Y W > type species = & Q Type and only host1 : Spotted sucker Minytrema melan- America, described three new genera and four new species, ops* X5 [ ] Y X Y including P. ingens (as Monobothrium ingens) from Ictio- 5 " " _ bus cyprinella (= I. cyprinellus) (Valenciennes) in Lake x# N Pepin, Minnesota. This author also amended the generic D 6 N M V x# diagnosis of Monobothrium, which, at the time, contained Carolina. ? M. wageneri Q 6 &E99 V x XN Q Tinca tinca X Y \ @ '*E%*Y W " 5 " " xM logical characteristics of these species from suckers (Cato@ N XN Q '*E%&Y Y Q > W Q W ? @ " > M. melanops X [ 6 N &'; N 9*EY vesicle (absent in M. wageneri) and absence of postovarian 5 " V X @++9Y " X\ &E9' &E%*Y from M. melanops XN 9;% M Y @ Two other suckers, namely C. commersonii (Lacepède) 5 " N 9*&9 X @++9Y and I. bubalus " 6 [ " 5 V # W X&EG;Y P. ingens X \ &EEEY " 6 [ 6 6 W Remarks. The type species of the genus was described in new hosts, namely C. cyprinus from # R 6 from Minytrema melanops 5 " x# @ and M. anisurum X5 [ ] Y R 5 " 6 N 6 R W W X >@ (M. melanopsY XV # W &EG; \ &EEE< Y [ P. ingens. Y V # W X&EG;Y @ larity of P. minytremi Q Mono- Promonobothrium hunteri XV # W &EG%Y bothrium which possess separate gonopores and lack pos@ x 6 R 6 tova5rian vitelline follicles. Promonobothrium minytremi 9*&+ = & was differentiated from these species of Monobothrium Synonym: Monobothrium hunteri V # W &EG% 6 6 " > Type host: Catostomus commersonii X &;*%Y
1 \ [ 6 # W #
= 9*&G G% **; Page 4 of 14 &* &88&&H 9*&G **; Oros et al.: Synopsis of Promonobothrium (Cestoda)
Additional hosts: Carpiodes cyprinus (Lesueur), Blue@ x# R head sucker Catostomus discobolus = V > x sucker C. latipinnis X Y Q H. nigricans XN Q G*'8&Y M. erythru- hog sucker Hypentelium nigricans* (Lesueur) (new host re@ rum XN Q G*'89Y " H. nigricans, IA (lo@ cord), Shorthead redhorse Moxostoma macrolepidotum (Lesu@ # W Y XN Q G*'8%Y " E. ob- eur) (Catostominae). longus W # Q XN Q '%'8&Y 9' " x D # D @ from E. oblongus Q X\RV %%9+% %%9+8 %%9+G # Q W # N %%9+'Y " M. macrolepidotum 5 @ D 6 M 6 < N M @ 5 " R N XN Q '8E&'Y M # V Q 6 # Q W # Q Q W Q D # " N R M. melanops XN 9&' M Y R R 5 " V N 9*&9 X @+*+Y V > x V # W X&EGGY C. commersonii # XN Q +E;G'Y R X&E'8Y V X&E'+Y voucher from C. commersonii 6 # Q XN Q +E;'&Y " X Y Remarks. Promonobothrium ulmeri was described as C. commersonii R X @+*+Y W " Monobothrium ulmeri from three species of catostomid from M. macrolepidotum Q? N X\RV 9*8++Y [ Hypentelium nigricans (type host), Moxos- Q W W toma anisurum and M. erythrurum. The new species was C. commersonii R W 5 " V 6 R Q 6 6 XD &H*E Y D # # D R of distinct loculi (vs > W @ XD &%+H*EY H. nigricans (new host culi in P. ingensY 6 X%+*M+'* Y R 5 " R N 9**E XD &+;H*E @ in P. ulmeri vs '*M&9* P. hunteri), different hosts, i.e. < @+*+Y species of Moxostoma 5 [ ] Hypentelium 5 @ V # W X&EG%Y X&EG'Y R X&E''Y ] P. ulmeri and Catostomus commersonii for P. hunteri. Remarks. This species was described as Monobothrium V # W X&EGGY hunteri 6 V # W X&EG%Y W # development of P. ulmeri from the release of eggs to the differences between the Palaearctic (M. auriculatum Ku@ formation of infective plerocercoids in the oligochaete in@ # " # &EG& W Caryophyllaeus M termediate hosts, Limnodrilus hoffmeisteri Claparède and 9*&+ M. wageneriY Q XM. ingens and Tubifex templetoni Southern, under laboratory conditions. M. hunteri, now in Promonobothrium M " W [ 6 >@ 9*&+Y Monobothrium in their morphology and perimental infection of L. hoffmeisteri X 9G '* host associations, thus indicating that the genus may be an > %'Y [ 6 > Promonobothrium hunteri &M9 " Promonobothrium fossae XR &E'8Y vitelline follicles in some cases, which are always associ@ x 6 R 6 9*&+ ated with the postovarian loop of the vitelloduct (Mackie@ Synonym: Monobothrium fossae R &E'8 W &EG%Y W " " T y p e a n d o n l y h o s t : Blacktail redhorse Moxostoma follicles were observed in one specimen collected from poecilurum Jordan (Catostominae). C. commersonii R W 5 " Hypentelium ni- N # # V gricans X Y W [ 6 N P. hunteri. D 6 N M 6 V > x Promonobothrium ulmeri (Calentine et Mackiewicz, M. poecilurum N # N XN Q &EGGY x 6 '98G%Y R 6 9*&+ = &D 8 = Q W Q Synonym: Monobothrium ulmeri Calentine et Mackie@ 5 R X&E'8Y< 6 W &EGG since its original description. Type host: Hypentelium nigricans (Lesueur). Remarks. See comments on P. mackiewiczi. ? # 6 # Erimyzon oblongus (Mitchill), Moxostoma anisurum X5 [ ] Y M. erythrurum X5 [ ] Y M. macrolepi- Promonobothrium mackiewiczi XR &E'8Y dotum (Lesueur), Minytrema melanops X5 [ ] Y X @ x 6 R 6 tostominae). 9*&+ Intermediate host: Limnodrilus hoffmeisteri Claparède Synonym: Monobothrium mackiewiczi R &E'8 Xx 6 [ Y T y p e a n d o n l y h o s t : Alabama hog sucker Hypentelium = # W 5 " \ etowanum (Jordan) (Catostominae). W N T y p e l o c a l i t y : Kowalinga Creek, Coosa Co., Alabama, D 6 N M W V Q
= 9*&G G% **; Page 5 of 14 &* &88&&H 9*&G **; Oros et al.: Synopsis of Promonobothrium (Cestoda)
N Remarks. The monotypic genus Rogersus is now trans@ D 6 N M 6 ferred to Promonobothrium as a new combination based V > W on morphological characters typical of Promonobothri- H. etowanum M W # N XN Q um W [ '98G+ '98GGY host, C. velifer, collected in Mississippi for this study, Q W Q fully correspond to those described as Rogersus roger- 5 R X&E'8Y< 6 si from M. poecilurum from the original description, for since its original description. W R X&E;*Y W Rogersus R acknowledge the differences in host and geography (but Remarks. Two caryophyllidean cestodes, Promon- both drainages are southern), but the specimens studied by obothrium fossae and P. mackiewiczi, were described as the present authors are morphologically indistinguishable Monobothrium fossae and M. mackiewiczi 6 R from those of R. rogersi > @ X&E'8Y Moxostoma poecilurum Jordan and Hypen- W 6 [ telium etowanum (Jordan), in Alabama, respectively. In on comparison of our specimens with types of this species, 6 [" Q including its holotype, and present evidence of some fea@ of Monobothrium (all currently in Promonobothrium) tures present in the type species that are absent in the orig@ was presented. The two species were differentiated from inal description. As shown for P. hunteri (see above) and each other by a smaller body and more testes in P. fossae > 6 " X9+EM%G8 vs &%;M&'E P. mackiewiczi), and W [ W 6 R. rogersi. in their manner of attaching to the host gut: several worms Rogersus rogersi W [ @ together in mucosal pits in the case of P. fossae vs [ 6 > W attachment of individual worms in P. mackiewiczi. Both 6 # X6 6 [ + &% V # @ species most closely resemble P. ulmeri in having a dig@ W &EE8 > Y W > # " " \ W " X= & \Y follicles, and differ only in a different number of testes (as > X= 8Y W G8% P. ulmeri vs %G8 &'E P. fossae and collected specimens from C. velifer demonstrated a lo@ P. mackiewiczi, respectively) and osmoregulatory canals > > W # (10 in P. ulmeri vs ; &G P. fossae and P. mackiewic- with a pair of median loculi, two lateral depressions and zi " Y X 6 & R &E'8Y W " > 6 Because of the lack of newly collected material of # @ # W 6 @6 P. fossae and P. mackiewiczi, including ethanol preserved Q R Q 6 R material, both species are provisionally retained as valid Q 6 6 " 6 [ X 9*&9Y V " with other congeners. W [ 6 6 > " X V # W &EE8Y 6 Promonobothrium rogersi XR &E;*Y 6 including holotype, as well as the newly collected material = & \ %? 8 ? X = %? 8?Y Synonym: Rogersus rogersi R &E;* > " for the species of Promonobothrium. ZooBank number for species: 6 # &'D**9 D@9 +E@89 ;@;;9 @&?&?D8&& D9* Promonobothrium currani sp. n. = &? = M 9 % 8 D Type host: Moxostoma poecilurum Jordan (Catostominae).
\ [ # Carpiodes velifer* ZooBank number for species: X5 [ ] Y X 6 Y X W Y 6 # *;*; 8 @G+''@8 &@E @ %98+ %% & 5 " " 6 N Description X6 [" W @ D 6 N M 6 V from Ictiobus bubalus W @ I. niger V > \ XN Q M M 6 # V '8;+8Y XN Q '8;++Y M. poecilu- N Y W W # rum 5 " N 6 9& +M9+ * > W * ; X* ; Y Q W ? C. velifer X [ Q N &E& M N 9&G " @ W M N &;G@E+Y 5 " X= 9 Y " W V N 9*&9 XQ\VNM 9*&G 9 99 %@8 [ X= &Y @ @GE; \RV &*&E'E@;*Y W @ " 6 6 5 R X&E;*Y< [6 X= 8 Y x W [ 6 &* > " X= 8 Y
= 9*&G G% **; G &8 &* &88&&H 9*&G **; Oros et al.: Synopsis of Promonobothrium (Cestoda)
Table 2. Measurements of new species of Promonobothrium V # W &EG;
P. currani sp. n. P. currani sp. n. P. papiliovarium sp. n. Characters/Promonobothrium spp. Ictiobus bubalus X5 [ ] Y X Y Ictiobus niger X5 [ ] Y Erimyzon oblongus (Mitchill) present study (n = 5) present study (n = 1) present study (n = 7) Body length (mm) 21.5M25.0 @ 17.0M50.5 > W * ; * ; * ; > distinct, loculotruncate distinct, loculotruncate indistinct, digitiform length G8*ME;G 750 @ width G%*M;G8 '+9 9G9MG9% Q # W X3 Y %*GM+*& 407 %GEMGE8 Testis size &&'M&E& & &&9M&++ &8+M&'& & &9GM&8& &&8M9&9 & E8M9*8 number &''M9++ @ 9&9M9+% [ " X Y 2.1–2.6 2.6 1.1–6.3 > X Y 9.2–10.7 9.5 3.4M14.6 reach up to anterior part of CS anterior part of CS anterior part of CS [ X Y E +M&& ' E +M%% @ X Y oval oval spherical size %8%M8*8 & 9;'M8*& %9*M%E; 9'GM8E8 & 9G&M8E* > W 6 &H9 &H9 &H%*M&H9+ Ovary shape H H width +G%MGE* +;' 8%9MEGE length of ovarian arms '&8ME'& ''9M;&& GG;M& 88G width of ovarian arms &8*M9+* &;&M&;% &8GM%G8 length of anterior ovarian arms 246–445 264–287 364M1 001 length of posterior ovarian arms 152–316 224–229 86M326 Vitelline follicle size &*8M&+G & G'M&*8 &*GM&%9 & ''M&*8 ;*M&;E & +GM&%E > X Y ' &M; & G E 9 9M; % reach up to posterior part of CS posterior part of CS posterior part of CS N > X Y & 'M& ; & G & +M% + in relation to length of testicular area &H'M&HG @ &H;M&H' Size of intrauterine eggs 8;MG* & %'M8+ +*M+; & %%M%E 8%M+; & %9M%' ?> " present present present 5 long, narrow long, narrow long, narrow separate separate separate Postovarian vitelline follicles present present absent Values in bold indicate differences between the species.
> X [ + % V # W & G'M&*8 X&*GM&%9 & ''M&*8Y X +*Y [ &EE8 > Y W 6 6 ' &M; & XG EY G8*ME;G X'+*Y G%*M;G8 X'+9Y W W > > " @ shallow median loculi, two lateral depressions and with @ 6 " @ " > X= &? = % Y Q # @ X= 9 % Y " " %*GM+*& X8*'Y W and median, surrounding testes, postovarian follicles pres@ X= 8 Y &''M9++ 6 &8M&E X&GY 6 X= % Y X 8Y &&'M&E& & &&9M&++ X&8+M&'& N " 6 W " " @ & &9GM&8&Y X +*Y< 6 @ " >@ E 9M&* ' XE + Y >@ @ 3 " W 9 &M9 G X9 G Y " N W " " X= 9 % Y [ ? 6 " 8;MG* & %%M8+ " @ X= % Y @ X+*M+; & %%M%EY X %* W @ " %8%M8*8 & 9;'M8*& X%9* & %E;Y W @ mounts). 6 &H9 X&H9Y 6 W ?> Type host: Ictiobus bubalus X5 [ ] Y X " #@W W Ictiobinae). X= % 8DY V @ Other host: Ictiobus niger X5 [ ] Y X @ < 6 W E9M&+G X&*%Y X= &MY tiobinae). x" \@ X= % Y W S i t e o f i n f e c t i o n : Anterior intestine. ] " '&8ME'& X''9M;&&Y Type locality: # # 6 R < W " +G%MGE* X+;'Y 6 X%9 %8%%Q< E& *98;RY W slightly sinuous, forms with terminal part of uterus short V 5 " 6 V N uterovaginal canal opening by female genital pore poste@ x 5 " rior to male genital pore. Seminal receptacle absent. Vitel@ X%* +9&;Q< ;; 8;9+RY W " &*8M&+G V 5 " 6 V N
= 9*&G G% **; Page 7 of 14 &* &88&&H 9*&G **; Oros et al.: Synopsis of Promonobothrium (Cestoda)
# M I. bubalus, prevalence A B &** X9H9Y X GM9&Y< I. niger, prevalence 100% (1/1) (intensity of infection 1). D \ X W @ 9% V 9*&9 I. bubalus< [ Q N 98+ Y W XW @ mounts, from I. bubalus M Q N 9G* I. niger M Q N 988 M Y @G9E @ XW @ I. bubalus< [ Q N 9*G Y Q\VNM X 9*&G 9 99 &Y " X Y Q\VNM X 9*&G 9 99 9Y XW @ from I. bubalusY \RV X &*&E''Y " X Y \RV X &*&E';Y ? 5 6 x V @ > W XVx 5MY 9*&9 " 6 6 [ Remarks. Promonobothrium currani sp. n. is placed in Promonobothrium, as circumscribed recently by Scholz X9*&+Y because it possesses all but one diagnostic > W loculi and two lateral depressions, genital pores separate, \@ " > @ " " > " XV # W &EG; &EE8Y Promonobothrium is also 6 X = +Y The most obvious difference between P. currani and other species of Promonobothrium is in the presence of postovarian vitelline follicles in the new species and their 6 X= % Y W 6 > morphology (loculotruncate in the new species vs digiti@ @ Y X 6 & 9Y Promonobothrium currani is most similar to P. miny- tremi, but can be distinguished by morphology of the > W from the remaining body in P. minytremi X= & Y the anterior vitelline follicles and testes, which begin much > P. minytremi than in the W X [ &M% V # W &EG; 6 9 Y D [ " 6 different, but may occur in sympatry. The new species was found in two species of buffalo (I. bubalus and I. niger), whereas P. minytremi [ sucker (M. melanops).
Promonobothrium papiliovarium sp. n. = & _ 9 % D
Fig. 2. A M " W Promonobothrium currani sp. n. from ZooBank number for species: Ictiobus bubalus X Q\VNM < " WY< 6 # 8;; 8 8@E?8 @89G8@ E*E@ ? G'D%&8*; B M " W Promonobothrium papiliovarium sp. n. from Erimyzon oblongus (Mitchill) X @GE'< " Description X6 " W @ view). Abbreviations M < " M @ " < M @ < M < " M " < from Erimyzon oblongus). Body elongate, with narrow " M " " < M # 6 &' *M+* + W @
= 9*&G G% **; ; &8 &* &88&&H 9*&G **; Oros et al.: Synopsis of Promonobothrium (Cestoda)
A B C D E
Fig. 3. Line drawings of species of Promonobothrium V # W &EG; AMB M Promonobothrium currani sp. n. (holotype, IPCAS @G9EY Ictiobus bubalus X5 [ ] Y< CMD M Promonobothrium papiliovarium X @GE'Y Erimyzon oblongus XV Y< E M Promonobothrium rogersi XR &E;*Y 6 X N Q '8;++Y Moxostoma poecilurum _ M 6 W [ " < D ? M 6 > " Abbreviations M < " M " < M @ < " M > " < V M V < " M " < M < M < " M " < " M "
W X= 9 Y V > 6 receptacle absent. Vitelline follicles medullary, numerous, W * ; " @ "@ " 6 ;*M&;E & +GM&%E X +*Y [ W [ 6 9 9M; % @ W @ " 6 6 [6 > X= % Y > " > X [ + % V # W &EE8 @ 6 " > Y 6 9G9MG9% X= %DY " " W W " > X= & _ % Y surrounding testes, postovarian follicles absent. Q # %GEMGE8 W N " 6 W " " @ X= %DY 9&9M9+% 6 @ " &&8M9&9 & E8M9*8 X +*Y< @ > @ 3 W " 6 % 8M&8 G @ W " N W @ @ > & &MG % " ? 6 " 8%M+; & " [ " %9M%' X %* W @ Y @ X= %DY @ Type host: Erimyzon oblongus (Mitchill) (Catostomidae: 9'GM8E8 & 9G&M8E* " " W W @ Catostominae). 6 %+M8* 6 W ?> S i t e o f i n f e c t i o n : Anterior intestine. " #@W W # 5 '' " X= %DY V 5 R # Q N < 6 W ;8M9&9 X= %DY D \ X x" 6 \ @ X= %DY W @ 9% V 9*&9 W " X%G8M& **&Y X \RV %%9++Y N W X @ " X&8GM%G8Y W " @GE'Y 8%9MEGE 6 W @ ? minal part of uterus short uterovaginal canal opening by " X6 \ @ Y M 6 \ " @ female genital pore posterior to male genital pore. Seminal um = ovary.
= 9*&G G% **; E &8 &* &88&&H 9*&G **; Oros et al.: Synopsis of Promonobothrium (Cestoda)
A B C
D E F G
Fig. 4. Histological sections of species of Promonobothrium V # W &EG; A, D M P. currani sp. n. from Ictiobus bubalus (5 [ ] Y< B, E, G M P. rogersi XR &E;*Y 6 Carpiodes velifer X5 [ ] Y< C, F M P. ulmeri (Calentine et Mack@ W &EGGY Minytrema melanops X5 [ ] Y M M < DM M Abbreviations M @ < " M > " < M < M < M < M @ < M < M < M < " M "
Remarks. ?> Erimyzon meri< 6 6 6 @ oblongus [ Monobothrium ulmeri X\RV > \ W " 6 6 %%9++Y W W @ 6 " X6 \ @ vs @ " W " 6 \@ P. ulmeriY @ X vs oval in molecular analyses, the new species differs so conspicu@ P. ulmeri). Moreover, a postovarian vitelline duct is absent osly from the species currently placed in Promonobothri- in P. papiliovarium, whereas it is present in P. ulmeri. um that its validity is well supported. It belongs to Pro- W 6 [ Erimyzon monobothrium because of its overall morphology, i.e. an oblongus > 6 > [ 6 > W " > # Interrelationships of species of Promonobothrium > @ based on molecular data @ " " 6 D@ > " XV # W &EG; &EE8< Q XD&MD% Y DQ ] @ the present study). ported the monophyletic status of the genus Promonoboth- Promonobothrium papiliovarium sp. n. can be easily rium, as well as the monophyly of most of the species of distinguished from all congeners by the possession of the Promonobothrium (only P. rogersi and P. ulmeri were rep@ 6 \ @ " W 6 6 Y X= +Y much longer than posterior arms. All remaining species position of the genus Promonobothrium within caryophyl@ of Promonobothrium including P. currani sp. n. have an lideans, as estimated on the basis of current data, remains \@ " @ uncertain, however, the genus tends to form a lineage with ilar in length and shape, even though the ovary of P. rogersi Q may have the posterior lobes markedly shorter than the an@ the analysis (i.e. Glaridacris &E9* Hunterella X= %?Y V # W V &EG9Y Archigetes Leuckart, In addition, P. papiliovarium differs from the remaining &;'; @ @ @ ence employed. Similarly to the statistically weakly sup@ soventral view, which is spherical (oval in other species). ported phylogenetic position of Promonobothrium within Promonobothrium papiliovarium is most similar to P. ul- the Caryophyllidea, the interrelationships within the genus
= 9*&G G% **; Page 10 of 14 &* &88&&H 9*&G **; Oros et al.: Synopsis of Promonobothrium (Cestoda)
$ % $ $ " ! ! ! ! # " ! " !
Fig. 5. V > # 6 DQ 7 DQ 5 Promonobothrium V # W &EG; W X Y of species of Promonobothrium X Y " W DQ X 5 7 7 cY DQ X 5 7 7 cY X Y DQ X 5 7 Y DQ X 5 7 7 cY X Y< W Q " V 6 +* 6 6 * E 6 indicate number of substitutions per site. Species of Promonobothrium in bold. remain uncertain and prone to change depending on the W &EE8Y 6 W 6 X = + 6 P. currani [ 6 " " @ M M cles. Interestingly, P. hunteri and P. ulmeri, both usually Q M Y W " " X > W follicles are present in P. hunteri M V # W &EG%< @ DISCUSSION ent study), possess a postovarian loop of the vitelloduct Promonobothrium, for several decades a monotypic XV # W &EG%Y V " P. rogersi 6 < W 6 @ Q some specimens lack postovarian vitelline follicles, where@ X9*&+Y &M&8 " " [" Monoboth- XR &E;*< Y rium, to Promonobothrium based on morphological and [ " 6 molecular evidence. In the present paper, another species species of Promonobothrium in relation to this otherwise is transferred and two new species are described. In the 6 > Q W XV # W &EE8Y " 6 6 contain a comparable number of species, i.e. Isoglaridacris concluded that the presence or absence of postovarian vi@ V # W &EG+ XW &* 6 Y Biace- 6 # > @ tabulum \ &E9' XE Y X &E;G \ " 6 XQ Y &EEE 9*&9Y XV # W &EE8Y x The main novelty in the amended generic diagnosis pre@ suggest that this character should be used with caution sented in this study is that the species of Promonobothrium W Q either possess or lack postovarian vitelline follicles. The > @ presence or absence of postovarian vitelline follicles is re@ W [ > XV # W &E'9Y 6 X &E;G V #@ [ Xx 9*&*Y
= 9*&G G% **; Page 11 of 14 &* &88&&H 9*&G **; Oros et al.: Synopsis of Promonobothrium (Cestoda)
" > 6 integrative approach combining morphology, molecular within the most speciose genera, i.e. Khawia \ &E%+ analyses and ecological data on hosts and their distribu@ and Caryophyllaeus &'E* X x 9*&* 6 > XD 9**+ 9*&& 6# 9*&8 \ " #@ 9*&*Y 9*&+Y Among the species of Promonobothrium, different de@ X ?VY " @ [ > < P. minytremi " > W > from M. melanops and P. papiliovarium from E. oblongus in species of Promonobothrium. The type species P. miny- > W P. hunteri, P. ingens, P. ulmeri, tremi, along with P. rogersi and the newly described P. cur- P. currani and P. rogersi 6 6 X Y > rani > > Of these, P. ingens and P. rogersi are distributed across much wider than the neck, with a pair of shallow median both catostomine and ictiobine lineages of suckers, where@ W X= & ?MH). Other X Y > 6 @ > sociated with catostomines (as in P. ulmeri) or ictiobines the remaining body. Promonobothrium ingens possesses (as in P. currani). However, several host records have not > W @ @ 6 [ 6 @ ulate, whereas P. hunteri, P. papiliovarium and P. ulmeri [ X @ " > W 6 tion for host records). they differ from each other in the morphology of the repro@ The family Catostomidae is a diverse group of benthic " X # [ 6 WY W [ [ In contrast, the scanning electron micrographs have also Q W ' " " > W X_ # # &EE%Y @ studied species of Promonobothrium X = &Y W cording to the morphological, biochemical, early life his@ " [ W tory and molecular data, catostomids can be divided into host mucosa. four subfamilies: Catostominae, Cycleptinae, Ictiobinae [ V > XQ 9**GY [ X @ of phenotypic variability of the scolices at the interspe@ binae), including Ictiobus bubalus and I. niger that harbour [ " Promonobothrium, similarly as observed by Promonobothrium currani, inhabit pools, backwaters and X9*&&Y Khawia 5 main channels of small to large rivers in the Mississippi " " " [ 5 " 6 # V # phenotypic plasticity in the morphology of the anterior (Minytrema melanops, subfamily Catostominae) inhabits part of the body of Caryophyllaeus laticeps X &';&Y small to medium rivers over clay, sand or gravel, being as well as in congener C. brachycollis _ W # &E+% only occasionally found in large rivers and impoundments both parasites of freshwater cyprinids in the Palaearctic X &EE&Y X 6# 9*&8 \ " 9*&+Y According to recent phylogenetic analyses (Sun et al. > 9**' D 9*&* V 9*&9Y Ictio- > @ bus 5 [ ] Carpiodes 5 [ ] W 6 todes should be weighed on the basis of combined detailed to the monophyletic subfamily Ictiobinae, represent two morphological, molecular and ecological studies. # R " ?> Pliovitellaria = &E+& subfamily Catostominae, phylogenetic position of M. mel- Wenyonia R &E9% W " " anops is not W " XD 9*&* middle of the body, the ovary of all caryophyllideans is sit@ V 9*&9Y V 6 Promonobothrium 6 XV # W &E'9 " 6 @ 6@ 9*&&Y W " @ family Catostominae that contains four tribes (Chen and ent in the same genus only rarely, such as in Isoglaridacris V 9*&9Y Promonobothrium are known XV # W &E'9Y Khawia, the species of which [ [" 6 \@ " @ X @ ? XMinytrema and Erimyzon), Ca@ Y " X 9*&&Y tostomini (Catostomus), Thoburniinae (Hypentelium) and present study, two different shapes of the ovary were also V > XMoxostomaY D @ recognised among species of Promonobothrium, i.e. the tion of hosts of both of these species of Promonobothrium \@ " [" 6 @ > @ \ @ " P. papiliovarium X= %DY tion of congeneric tapeworms in phylogenetically distantly 6 W " Xx 9*&* # W @ @ 9*&& 6# 9*&8 \ " 9*&+Y esis of the host and its parasite. and also demonstrated in species of Promonobothrium by # X > > " Promonobothrium postovarian vitelline follicles, shape of the ovary) are sub@ [ # Pro- 3 [ " 6 monobothrium recognised as valid is presented. The key is " [ " based primarily on the gross morphology and a few char@
= 9*&G G% **; &9 &8 &* &88&&H 9*&G **; Oros et al.: Synopsis of Promonobothrium (Cestoda)
> \ W " @ 7 Postovarian loop of vitelloduct present, postovarian vi@ [ 6 " [ " @ telline follicles may be present ...... cies diagnoses provided in the morphological descriptions ...... Promonobothrium hunteri of the respective species. M " " 6 " vitelline follicles absent ...... ; & > > # W than neck region ...... 9 ; x 6 ; 9+* @ M > > W tes, worms together in mucosal pits ...... only slightly wider than neck region ...... 4 ...... Promonobothrium fossae M x 6 &G &;* @ 9 W W @ tes, worms not in mucosal pits ...... most vitelline follicles ...... 3 ...... Promonobothrium mackiewiczi M " follicles or on same level (in some cases close posterior * because of the lack of newly collected material of P. fossae and P. mackiewiczi, the key differential characters used herein are based on to anteriormost vitelline follicles) ...... " 6 R X&E'8Y ...... Promonobothrium rogersi comb. n. Acknowledgements. V x 5 M 6 % " " 6 < Miny- de Buron and Vincent A. Connors (College of Charleston, South trema melanops ...... Promonobothrium minytremi Y " ? 5 6 x" X M " " < 5 6 V Y V @ Ictiobus spp...... Promonobothrium currani sp. n. X > # R D > Y " 6 9*&9 R 6 XQ \ V Y # @ 8 > W @ @ " 6 ] Promonoboth- late ...... Promonobothrium ingens rium ? \ 6 XN Q Y M > # ...... 5 6 5 X\RVY 6 from their collections. Two anonymous reviewers provided help@ + x" \@ W ] " @ ful and insightful comments that helped improve considerably ian arms ...... G ] V x Q M x" 6 \ @ W " @ 9*&% W 6 6 = 6 = " < W 6 " # 5 D " Erimyzon oblongus ...... X 3 @*G+%@&& *&'&@*EY ...... Promonobothrium papiliovarium sp. n. ? XQ 9H*&+EH&GY X5x G**''%88Y = X 3 Q +*+H&9H &&9Y W # W # W W # @ G 6 W X 6 +* Y # 3 5 D " x long, more than 300 testes ...... 6 ?5D= X V 9G99*&9**99Y X* 9Y ...... Promonobothrium ulmeri Q = N X W *;&;GEG M W X 6 9* *;&;;9%Y # W Q 6 @ mm), neck short, fewer than 300 testes ...... 7 = D "
REFERENCES
ǂǏLjljǂǎ 5 džǏǂǓDž ? &E89 5 @ ǂǍdžǏǕNJǏdž 5 &EG' " " # [ D 6 # @ W '9 8&;M898 _ &' 99M%; ǂǍdžǏǕNJǏdž 5 VǂDŽnjNJdžǘNJDŽǛ _ &EGG Monobothrium ulmeri ǂǓȁǤnj D xǓǐǔ V \ǂǏǛdžǍǐǗǤ DŽljǐǍǛ 9*&8 @ X Y Q @ typic plasticity in Caryophyllaeus brachycollis Janiszewska, tomidae. Trans. Am. Microsc. Soc. ;+ +&GM+9* &E+% X Y [ ª ljdžǏ R @_ VǂǚDždžǏ 5 9*&9 # X ;; &+%M&GG Cypriniformes: Catostomidae): further evidence of relationships ǂǛǔǂǍǐǗNJDŽǔǐǗǤ ? MǓǤțǐǗǤ@\ǓǐǎǂDžǐǗǤ ǓǂǃdžDŽ _ \ǂǏ@ " 6 @ 5 9 > %+;G ǛdžǍǐǗǤ xǓǐǔ V DŽljǐǍǛ 9*&8 \ 6 W @ &E+M9&* phology and molecular data: a case of the genus Caryophyllaeus ljdžǓǗǚ 9**E N [ X Y W [ @ R # @ = G& %8GM%+9 9**9M9**; = +G &EEM9%* ǓǂǃdžDŽ _ DŽljǐǍǛ MǓǤțǐǗǤ@\ǓǐǎǂDžǐǗǤ ǂǛǔǂǍǐǗ@ DǂǚǓǂǕ 9**+ W " > _ NJDŽǔǐǗǤ ? xǍǔǐǏ D 9*&9 6 @ ;+ 8*'M8&+ ar paralogs of commonly employed phylogenetic markers in the Dǐǐǔdžǚ V \ ǂǓǕ \ ǂNJǕǐlj M VNJǚǂ V 9*&* @ @ W [ X @ X Y _ 89 9+EM9G' Y 6 QD8HQD+ ] ǂNJǓǂ _ Q _džǏǔdžǏ M ǂǓǃdžǂǖ ? 9*&9 6 D @ V ?" +8 &*9;M&*%8 6 R R R 6 6 WWW @ ǓNJǎdžǔ 5 VNJǍǍdžǓ &E'+ W 6 &9H9*&+ creek chubsucker, Erimyzon oblongus XV Y Q @ lina. J. Parasitol. G& E'%ME'8
= 9*&G G% **; Page 13 of 14 &* &88&&H 9*&G **; Oros et al.: Synopsis of Promonobothrium (Cestoda)
\ǂǍdžǚ D 5 ǂǓLjdžǓ V 9*&8 W 5 [ caryophyllaeid (Cestoda: Caryophyllidea) from spotted suckers, = +' %'M8G Minytrema melanops (Catostomidae), in the Big Thicket nation@ ǂLjdž V ǖǓǓ V &EE& = = W = " > N ;& 9%M9G Q Q V > \ V \ \ǂǏǛdžǍǐǗǤ xǓǐǔ V ǂǓȁǤnj D VNJnjǍNJǔǐǗǤ D MNJǓNJǏ D V 8%9 DŽljǐǍǛ 9*&+ V W ǍdžNJNjdžǍ = _ǐǏDždžǍNJǖǔ N QǐǓǍNJǏDždžǓ ? QǚLjǓdžǏ xǙdžǍ@ redescription of Caryophyllaeus laticeps X &';&Y X @ ǎǂǏ DŽljǂǏDždžǓ ǖǏDžǃdžǓLj ljǐǍǍdžǔǔǐǏ V 9**; toda: Caryophyllidea) and characterisation of its morphotypes W ª " @ [ E* &''M&E* V ?" 8; %GEM%'& \ǐLJLJǎǂǏ &EEE Q W [ @ 5ǐǏǒǖNJǔǕ = džǔǍdžǏnjǐ V ǗǂǏ DždžǓ VǂǓnj ǚǓdžǔ D ? # 6 DǂǓǍNJǏLj \ǷljǏǂ ǂǓLjdžǕ NJǖ ǖDŽljǂǓDž V N " Q W # +%E \ǖdžǍǔdžǏǃdžDŽnj _ 9*&9 V % 9 [ @ \ǖǏǕdžǓ R &E9' Q Q logenetic inference and model choice across a large model space. America. J. Parasitol. &8 &GM9G G& +%EM+89 \ǖǏǕdžǓ R &E%* Q DŽljǂdžLJLJǏdžǓ _NJǓnjȳ V VǂljǎǐǖDž Q DŽljǐǍǛ 9*&& V && X&E9'Y &;G 5 " Wenyonia (Cestoda: Caryophyllidea) from Synodon- _džǏnjNJǏǔ 5 ? ǖǓnjljdžǂDž Q V &EE% = W = @ tis [ 'E ;%M&*' = V & *'E DŽljǍNJDŽnj@ ǕdžNJǏdžǓ ǕdžNJǏdžǓ = V džNJLJdžǓǕ ǕǂǖLJLJdžǓ MǂǕǐlj M ǕǂǏDžǍdžǚ D V 9*&% V == ] ljǓNJǔǕNJǂǏ ? ǓǐǛNJdžǓ 5 \ 9*&* " > alignment software version 7: improvements in performance and > 6 " 5 " 6 V ?" %* ''9M';* ? ++ 89&M8%; MdžǂǓǔdž V VǐNJǓ 5 RNJǍǔǐǏ ǕǐǏdžǔ@\ǂǗǂǔ ljdžǖǏLj DŽljǎNJDžǕ D &E;G 5 \ 6 # W [ V ǕǖǓǓǐDŽnj ǖǙǕǐǏ ǐǐǑdžǓ VǂǓnjǐǘNJǕǛ 5 5 = G'+ DǖǓǂǏ ljNJdžǓdžǓ ǔljǕǐǏ VdžǏǕNjNJdžǔ DǓǖǎ@ DŽljǐǍǛ ǓǂǃdžDŽ _ MǓǤțǐǗǤ@\ǓǐǎǂDžǐǗǤ xǓǐǔ V ǎǐǏDž 9*&9 > 6 ǂǛǔǂǍǐǗNJDŽǔǐǗǤ ? ?ǓǎǐǍdžǏnjǐ \ǂǏǛdžǍǐǗǤ 9*&& desktop software platform for the organization and analysis of 5 " Khawia spp. (Cestoda: Caryophyllidea), parasites of ] 9; &G8'M&G8E [ # [ @ MǓǤǍǐǗǤ@\ǓǐǎǂDžǐǗǤ ǕdžLJnjǂ _ ǑǂnjǖǍǐǗǤ V xǓǐǔǐǗǤ = +; &E'M99% V ǐǎǃǂǓǐǗǤ V \ǂǏǛdžǍǐǗǤ ǂǛǔǂǍǐǗNJDŽǔǐǗǤ ? DŽljǐǍǛ ljǐǖDžljǖǓǚ 9*&8 W [ DŽljǐǍǛ 9*&* @ " 6 & Q W ª _ &** 9GM8+ X &Y 9 6 ] " # W @ DŽljǐǍǛ xǓǐǔ V ljǐǖDžljǖǓǚ ǓǂǃdžDŽ _ RǂdžǔDŽljdžǏ@ DQ @ ǃǂDŽlj 9*&+ Q W W [ sis, the monozoic cestode of common carp. Int. J. Parasitol. 40: Monobothrium D &;G% Promonobothrium Mackie@ &'+M&;& W &EG; X Y ǂǏLJdžǂǓ 5 ǂǍDŽǐǕǕ \ǐ R ǖNJǏDžǐǏ 9*&9 @ molecular evidence. J. Parasitol. &*& 9EM%G [ 6 6@ ǑǂnjǖǍǐǗǤ V xǓǐǔǐǗǤ V VǂDŽnjNJdžǘNJDŽǛ _ 9*&& @ V ?" 9E ics and chromosomes of tapeworms (Platyhelminthes, Cestoda). &GE+M&'*& " '8 &''M9%* VǂDŽnjNJdžǘNJDŽǛ _ &EG% Monobothrium hunteri sp. n. (Cestoidea: ǖǏ \ ºNJdž º RǂǏLj R V NJǖ ǓdždžǓ ljǂǏLj Caryophyllaeidae) from Catostomus commersoni X Y M.M. 9**' " 6 @ X Y Q _ 8E [ 6 DQ ] '9%%'%* _ = '* 9E&M%*E VǂDŽnjNJdžǘNJDŽǛ _ &EG; W W RNJǍǍNJǂǎǔ D D &E'' Biacetabulum oregoni sp. n. (Cestoda: the spotted sucker, Minytrema melanops X5 Y X Y Caryophyllidea) from Catostomus macrocheilus. Iowa State J. J. Parasitol. +8 ;*;%;&% 5 +9 %E'M8** VǂDŽnjNJdžǘNJDŽǛ _ &E'9 X Y " W ?> RNJǍǍNJǂǎǔ D D &E'; # R @ %& 8&'M+&9 [ W _ 5 +& 8'&M8'' VǂDŽnjNJdžǘNJDŽǛ _ &E;& X Y " RNJǍǍNJǂǎǔ ? \ &E'8 W W Monobothrium (Cestoda: [ " &E &%EM9*G Y [ VǂDŽnjNJdžǘNJDŽǛ _ &EE8 x " N = &*% G&*MG&+ &;G% = M _ 5 X? Y RNJǍǍNJǂǎǔ ? \ &E;* Rogersus rogersi gen. et sp. n. (Cestoda: Keys to the Cestode Parasites of Vertebrates. CAB International, Caryophyllaeidae) from the blacktail redhorse, Moxostoma R 9&M8% poecilurum X_ Y Xx Y N VǂDŽnjNJdžǘNJDŽǛ _ 9**% X Y States. J. Parasitol. GG +G8M+G; " 8; &8%M&+8 ǘNJDŽnjǍ D _ 9**G @ QdžǍǔǐǏ _ 9**G = R _ R Q W 6 ] # G98 > # D N " > xǓǐǔ V DŽljǐǍǛ \ǂǏǛdžǍǐǗǤ VǂDŽnjNJdžǘNJDŽǛ _ 9*&* at Austin, 115 pp. > X Y
5 " %* Q " 6 9*&+ &+ = 6 9*&G 6 9% V 9*&G
Cite this article as: x V 6 _ M 5 9*&G " W Promonobothrium Mackie@ W &EG; X Y # X Y Q W W W = G% **;
= 9*&G G% **; Page 14 of 14