94(3), 2008, pp. 716–726
American Society of Parasitologists 2008
A REVIEW OF SPECIES IN THE GENUS
Terry R. Haverkost and Scott L. Gardner
The Harold W. Manter Laboratory of Parasitology, University of Nebraska State Museum, Lincoln, Nebraska 68588-0514.
: Currently, there are 6 recognized species in the genus
. These parasites are found in the small intestinesof numerous species of marsupials throughout North and South America. Small mistakes in various classical taxonomic workshave given rise to recent and numerous misidentiﬁcations of these species. In this work, we examine a total of 99 specimensacross all species from museum collections in an attempt to determine informative taxonomic characters to distinguish thesespecies. Despite confusion in the literature, accurate identiﬁcation of these species can be achieved by observing the presence orabsence of oral and ﬂanking spines anterior to the oral sucker.
(Trematoda: Rhopalidae) occur in thesmall intestines of marsupials throughout the Nearctic and Neo-tropical regions. Six species are currently recognized as valid.Rudolphi (1819) described
Linnaeus, 1758 collected by Johann Natterer inBrazil. Diesing (1850) described
(Zimmermann, 1780) also collected byNatterer. Stiles and Hassall (1898) recognized that
was an occupied name, and renamed the genus
.Braun (1901) provided formal descriptions of both previouslydescribed species, and recognized and described another spe-cies,
, from the same material examined byDiesing (1850). Chandler (1932) described
from North America in
Kerr,1792. Kifune and Uyema (1982) described
(Linnaeus,1758) in Brazil. Finally, Rivallis et al. (2004) described themost recent species in this genus,
collected in Colombia.Skrjabin (1948) reproduced the descriptions of the species inthe genus and provided an identiﬁcation key to the known spe-cies of
in that work, it is apparent that the labels forthe ﬁgures referring to
were re-versed. Travassos et al. (1969) reproduced the work of Skrjabin(1948), along with the mislabeled ﬁgures. In the summary work of Yamaguti (1971), Figure 1651 of
is areproduction of
after Caballero (1946); this ﬁgureis in fact a representation of the currently recognized species
. In the same work (Yamaguti, 1971), Figure1650B of
, after Caballero (1946), but attributed toBraun (1901), also appears to be a drawing of
,but because the tentacles are covering the area where ﬂankingspines would be found, identiﬁcation to species is not possiblefrom the Figure (although it is likely
). Finally,Radev et al. (2005) offer original ﬁgures of
drawn from specimens stored in the Manter Laboratory of Par-asitology (HWML 0844, 22422, 22423). These specimens havebeen veriﬁed by 1 of us (T.R.H.) as
, but theﬁgures of these specimens in Radev et al. (2005) lack the de-ﬁning ﬂanking spines, making them easily mistaken for
.The species of
are some of the most distinctiveand easily recognized trematodes of mammals. However, the
Received 16 August 2007; revised 22 October 2007; accepted 25 Oc-tober 2007.
correct identiﬁcation of the species in this genus has remaineda difﬁcult task, given the ﬂaws in the above references and thelack of a thorough investigation of important taxonomic char-acters. Here, we intend to provide the most thorough investi-gation, to our knowledge, of the 6 currently recognized speciesin this genus.
MATERIALS AND METHODS
Specimens prepared and studied by us were stained in acetic Semi-chon’s carmine, dehydrated in an ethanol series, cleared in terpineol andxylene, and mounted in Canada balsam or gum Damar (Prichard andKruse, 1982). All other specimens studied were obtained from museumcollections including: the United States National Parasite Collection(USNPC), Beltsville, Maryland; the Harold W. Manter Laboratory of Parasitology (HWML), Lincoln, Nebraska; the Naturhistorische Muse-um Wein (NMW), Vienna, Austria; the Universidad Nacional Auto´no-ma de Me´xico (UNAM), Mexico City, Mexico; the Museo de HistoriaNatural at the Universidad Nacional Mayor de San Marcos (UNMSM),Lima, Peru; and the Kyushu University Museum (FUK), Fukuoka, Ja-pan. Due to the commonality of misidentiﬁed and mislabeled specimensthroughout the previous literature and in museum holdings, this paperdeals only with specimens that have been studied by the authors. Wedo not include a comparative table of measurements from previous stud-ies for this same reason. All measurements were taken with a ZeissUltraphot
microscope equipped with an ocular micrometer and arepresented to the nearest micrometer unless otherwise noted; ranges arefollowed by the mean and the number of characters studied (n) is givenif different from the number of specimens studied (N). Canonical dis-criminant analysis (CDA) was performed on the natural log transfor-mations of the 23 measurements summarized in Tables I and II exclud-ing egg length and egg width using PROC CANDISC in SAS (version6.12, SAS Institute, Cary, North Carolina).
(22 specimens total): HWML34950 (1specimen) from
, Paraguay; HWML70000(12 vouchers) from
, Santa Cruz Department, Bo-livia; HWML70002 (1 specimen) from
, Santa Cruz,Department, Bolivia; HWML70009 (3 vouchers) from
, Santa Cruz, Bolivia; HWML70013 (1 specimen) from
Berisso, Argentina; UNAM4081 (1specimen) from
sp., Veracruz, Mexico; USNPC14998(1 specimen) from unknown host, Panama; USNPC72792 (2vouchers) from
, El Tacal, Venezuela.
: V4677 (1 neotype, 1 voucher) from
(4 specimens total): UNAM1225 (2specimens) from
, Guazacapa, Guate-