© Institute of Parasitology, Biology Centre CAS =9*&GG%**; doi: &*&88&&H9*&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"#56< 9=N" 5# D3"56< 3D"QQ6DRN Abstract: Monozoic cestodes of the recently amended genus Promonobothrium V#W⪚XY@ #XYQ"WW[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 minytremiV#W⪚XY<P. ingens X\&E9'Y<P. hunteri XV#W&EG%Y< P. ulmeri XV#W&EGGY<P. fossae XR&E'8YP. mackiewicziXR&E'8Y RogersusR&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 bubalusX5[]YIctiobus niger X5[]YErimyzon oblongus (Mitchill), respectively. The newly described species can be distinguished from the other congeners by the morphology of the >""" "V>6]665Q XDQDQY["# [Promonobothrium based on morphological characteristics is provided. Keywords:?[">WQ5[# Systematic research on caryophyllidean cestodes in XM"@\"9*&*#"9*&& Q6W\RX&E%*Y> 69*&9Y6@ monograph of the group and reached its highest intensi@ X6#9*&8"@ &EG*R&E'*RW" "9*&8\"9*&+Y _V#WV_N5_?\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 MonobothriumD&;G% &EEEYV#W6@ monotypic Promonobothrium V#W ⪚ \W@ tant contributions to the current knowledge of this enig@ " > 6 6WXV#W&E'9&E;& > &EE8 9**%Y 6 study on amended Promonobothrium are presented and &E;*R X #["@ 9*&8YW63@ 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[]YV@ 6Q6 pi, and from eastern creek chubsucker Erimyzon oblongus caryophyllidean cestodes represent an interesting model XVYQN"@ for studies of unusual molecular and cytogenetic phenome@ relationships of species of Promonobothrium are assessed. "#%&%'**+5# D3"56789*%;+%&*%+&<=>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&&H9*&G**; Oros et al.: Synopsis of Promonobothrium (Cestoda) Table 1. List of specimens of species of PromonobothriumV#W⪚] # Specimen Host XNY Voucher DQHDQ 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 N98+ MNGG++'*HMNGG++;& Promonobothrium hunteri Hypentelium nigricans RW5"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 R5"R D&&+H*E@ MNGG++G*HMNGG++'& X\&E9'Y X5[]Y Promonobothrium minytremi Minytrema melanops R5"R @8*G MNGG++G9HMNGG++'% V#W⪚ X5[]Y Promonobothrium minytremi Minytrema melanops 5"V N&';6 MNGG++GGHMNGG++'' Promonobothrium minytremi Minytrema melanops 5"V N9*E MNGG++G;HMNGG++;* Promonobothrium minytremi Minytrema melanops 5"V N9*E@ MNGG++GEHMNGG++'E Promonobothrium rogersi 5"V N&E& MNGG++G'HMNGG++'; XR&E;*Y6 Carpiodes velifer X5[]Y @QM]DQ=3HHWWWW6H><NQDQM][D MATERIALS AND METHODS "W"X9**EY Specimens studied "W3X9**;Y Material of the following species was studied: Promono- Deposition of specimens bothrium minytremi V#W ⪚ X Y< P. in- Specimens studied are deposited in the following collections: gensX\&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. mackiewicziXR&E'8Y<Rogersus rogersiR 3"56XY\RV@ &E;*QWW[> 6N"Q6#VN X5MVxY6Q X\RVYQ\VNMXQ\VNMY V > R Q V Q \ XQVQ\Y N9*&*9*&99*&% RDN Morphological evaluation Molecular data generation and analyses Tapeworms collected recently by the present authors were pro@ @WW[> 66xX9*&*Y\ EG@X live tapeworms were isolated from the host intestine and rinsed W[>W66"" in saline. Hot (almost boiling) 4% neutral buffered formaldehyde "Y6]>DQ solution (= 10% buffered formalin) was added into the beaker or X6&YDQ "WW69M%W#@ 5[65QXD@ worms were transferred to 70% ethanol before further processing. Q DQY ] W 6 WWVR@ X9*&9Y]W6@ mine solution, dehydrated through graded ethanol series, cleared "'XHHWWWHM9*&9Y in clove oil (eugenol) and mounted in Canada balsam as perma@ @?@Q@ nent preparations. Line drawings were made using a drawing at@ V==XM9*&%Y DV+***W 6W> Q#VW# 6] %;#XY"@ Phylogenetic relationships were estimated from a concatenat@ en in micrometres unless otherwise indicated. Pieces of worms >#XVY@ 6W6[W>@ XY[" &9M&+3WRR>@ were selected using the small sample size corrected Akaike in@ stained with 1% eosin B following recently updated protocols =&&*X9*&9Y Xx9*&*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 [YV6W6 X9*&*Y__VG+&*?V@ + &** 6 5 ?V W 6 WV"%9X5] from Erimyzon oblongusW#W@@ 9*&9Y8V%89*@ 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 PromonobothriumW?V6@ V@W*9+ =9*&GG%**; 9&8 &*&88&&H9*&G**; Oros et al.: Synopsis of Promonobothrium (Cestoda) Fig. 1. Scanning electron micrographs of species of PromonobothriumXM_M>MMM[Y AMPromonobothrium minytremi V#W ⪚ Minytrema melanops X5[]Y< B M P. ingens X\ &E9'Y Carpiodes cyprinusXY<CMP. hunteri XV#W&EG%YHypentelium nigricans XY<DMP. ulmeri (Calentine et V#W&EGGYMinytrema melanops X9*&+Y<E, F, K, LMP. currani sp. n. from Ictiobus bubalus X5[@ ]Y<G, HMP. rogersiXR&E;*Y6Carpiodes veliferX5[]Y<I, JMP. papiliovarium sp. n. from Erimyzon oblongus (Mitchill). RESULTS Promonobothrium V#W⪚ Promonobothrium W E Synonyms: MonobothriumD&;G%partim (spe@ P. minytremi (type species), P. ingens, P. hunteri, P. ul- QYRogersus 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@ ⪚ YW>[" hosts, life cycle and geographical distribution is provided Amended generic diagnosis of Promonobothrium: 6WWW> Caryophyllidea: Caryophyllaeidae. Body elongated to ?V " W [@ =9*&GG%**; Page 3 of 14 &*&88&&H9*&G**; Oros et al.: Synopsis of Promonobothrium (Cestoda) itriches. Inner longitudinal musculature well developed. ""X[;V#W > @ ⪚Y5P. minytremi from W">Q#@ M. melanops are identical with those described by Mack@ @"@ WX⪚Y ?> " x" \@6\@6 Promonobothrium ingens X\&E9'Y Scholz, Vitelline follicles medullary, lateral and median. Postovari@ x6R69*&+ 6N =& @VN Synonym: Monobothrium ingens \&E9' ?6 T y p e h o s t : Bigmouth buffalo Ictiobus cyprinellus* (Valenci@ #XYQ ennes) (Ictiobinae). V#WX⪚&EE8Y Additional hosts: Quillback Carpiodes cyprinus* (Lesu@ X9*&+Y YX6YXWYR#Catostomus commersonii (Lacepède) (Catostominae), Smallmouth buffalo Remarks. Promonobothrium was erected by Mackie@ Ictiobus bubalus X5[]Y X6Y " WX⪚YP. minytremi Moxostoma anisurum X5[]Y XY XW from Minytrema melanopsX5[]Y5" host record). x#N@ #VN X9*&+YP. minytremi appears in a mono@ D 6 NMVQW#xx#@ phyletic lineage comprising caryophyllidean species from R Q6 V > x" in Monobothrium. Therefore, these species have been I. cyprinellus X6;9&;#WYN transferred to the originally monotypic Promonobothrium. X@+;%Y Moreover, RogersusR&E;*W QW x is transferred to Promonobothrium as a new combination, C. cyprinusXD;EH*EMY#R6 and two new species of Promonobothrium are described R<M. anisurumXD&&+H*EYR herein, thus increasing the total number of species in the 5"RN9**EX@+;%Y genus to nine. \ X&E9' &E%*Y X&E89YX&EG'YRX&E';Y Promonobothrium minytremi V#W⪚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[]YXY 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. wageneriQ6&E99 V xXNQ Tinca tincaXY\@ '*E%*YW"5""xM logical characteristics of these species from suckers (Cato@ NXNQ'*E%&Y YQ>W QW ? @ " > M. melanopsX[6N&';N9*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"N9*&9X@++9Y and I. bubalus"6[" 5V#WX⪚Y P. ingensX\&EEEY"6 [66W Remarks.