PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 109(4):653-663. 1996 A review of the taxonomic nomenclature and a checklist of the species of the Branchiobdellae (Annelida: Clitellata) Stuart R. Gelder Department of Science, University of Maine at Presque Isle, 181 Main Street, Presque Isle, Maine 04769-2888, U.S.A. Abstract. — The taxonomic nomenclature of the branchiobdellidans is re- viewed. This includes the higher rankings and the numerous generic names assigned during the first half of the 19th century, particularly the use of Bran- chiobdella for species of the marine leech, Branchellion. The current taxonomic situation is summarized to high light where inconsistencies and omissions exist in the data. The class Branchiobdellae consists of one order, Branchiobdellida, with five families containing a total of 21 genera and 149 species. The binomia are given with any junior synonyms, and these are followed by the citation in which the rank was reduced to junior status. In addition, the valid taxon Sin- odrilus heterorchis has been placed in incertae sedis until the entry point of the sperm ducts into the glandular atrium can be determined, two species are considered nomen inquirenda and three others nomen nudem. Branchiobdellidans are leech-like sym- bionts found mainly on astacid crayfish in the Holarctic. The taxon is monophyletic and is considered to be independent of, and equivalent in rank to, the clitellate Oligo- chaeta and Hirudinea (Holt 1989a, Gelder & Brinkhurst 1990, Brinkhurst & Gelder 1991). Currently, the Clitellata is consid- ered to be a subphylum, and so the three taxa just mentioned have the assigned rank of class. Thus, the higher taxonomic rank- ings of the branchiobdellidans are, the class Branchiobdellae Kasprzak, 1984, which consists of one order, Branchiobdellida Holt, 1965a, with five famihes (Holt 1986), Branchiobdellidae, Bdellodrilidae, Cambar- incolidae, Caridinophilidae, and Xironodril- idae. Historical review of the higher ranks. — The branchiobdellidans were raised to fam- ily rank by Grube (1851) with the name Branchiobdellea. Subsequently, Vejdovsky (1884) reduced "Branchiobdellidae aut." to a junior synonym of his proposed Discod- rilidae. Ludwig (1886:786) did not accept Vejdovsky 's recommendation, but changed the suffix used by Grube (1851) to read Branchiobdellidae. This spelling has contin- ued to be accepted as it conforms to the recommendations of the International Code of Zoological Nomenclature for construct- ing a family name. The taxonomy of the branchiobdellidans has undergone a number of revisions, and this has resulted in some of the names and ranks becoming superfluous. The list below gives all of the names and their ranks as they have appeared in the literature. Higher Taxonomic Ranks and Names Proposed for Branchiobdellidans Class Branchiobdellae Kasprzak, 1984:217 Subclass Branchiobdellida Stuart, 1982:43 Branchiobdellidea Sawyer, 1986:647 Order Branchiobdellida Holt, 1965a:30 Superfamily Branchiobdelloidae Hall, 1914:190 Family Branchiobdellea Grube, 1851:114 Discodrilidae Vejdovsky, 1884:38 Branchiobdelhdae Ludwig, 1886:782 Bdellodrilidae Holt, 1986:668 654 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Cambarincolidae Holt, 1986:692 Caridinophilidae Holt, 1986:692 Xironodrilidae Holt, 1986:691 Subfamily Branchiobdellinae Goodnight, 1940:27 Cambarincolinae Goodnight, 1940:29 "Branchiobdell-" for non-branchiobdel- lidans. — The stem "branchiobdell-" was used by Rudophi in an unpublished collec- tion name for a marine leech with paired, external gills, Branchiobdellion Torpedinis. The new species was described and named Branchellion torpedinis Savigny, 1820. Un- fortunately three junior synonyms were then created, Hirudo {Branchiobdelld) Ru- dolphii Blainville, 1827, Branchiobdella torpedinis Blainville, 1828, and Hirudo {Branchiobdella) torpedinis Gervais, 1836. These misuses of ''Branchiobdella" for Branchellion ceased with Diesing (1850) and the synonym status of B. torpedinis was reviewed by Blanchard (1894). Other suf- fixes to the stem have been found in the literature by the author, however, they have not been cited or discussed as this would have added nothing to the presented clari- fication of these misuses. Historical review of genera and spe- cies. — The first report of a worm that can be recognized as a branchiobdellidan was made by Roesel (1755) when he reported a worm living on the surface of the crayfish, Astacus Fluviatilis. Although the Roesel reference has often been cited in historical reviews of branchiobdellidans (Sawyer 1986, and others), no tangible information has been reported, and the page and figure numbers cited were usually incorrect. Therefore, a translation of the anatomical description and location of the symbiont on the host is given (Roesel 1755:327, tab. LIX, figs. 19-22). "25. With regard to the remaining figures in tab. LIX, particularly [figs] 19, 20, 21, 22. They show us a kind of worm that re- ceives its nourishment from the "river crab" especially in December and January. Figure 19 depicts them [the worms] in their natural maximum size [about 8mm long] — smaller versions can also be found. Their movement corresponds to the movement of a leech. Here [Fig. 19] I portray them in the positions they take up. Their basic color is a shiny brownish yellow; the head is their most supple member, it is a little pointed in shape and has a pair of black eye-spots. The remaining joints following the head get pro- gressively thicker — right to the last joint [segment] — the sections [segments] as well as the dividing points between the sec- tions are more round than wide. On the back of the third, fourth and fifth segment — counting from the rear — one notices a black spot discernible through the skin. It is simply feces in the rectum. These worms can be found anywhere under the breast plate of the "river crab", particularly at the two sides from the head to the tail; es- pecially where they occur in considerable numbers. Roesel's figures 20, 21, 22 show these worms in the same position as in [Fig.] 19, but greatly enlarged [about 5X]. Whether these [worms] originate within the body of the "river crab" or outside of it, I cannot say, but I have observed that the "river crabs" are covered with many small eggs that stick together in clumps, at the time when they harbor these worms." Roesel's figures 20 to 22 have been re- drawn (Fig. la-c) and show branchiobdel- lidans in typical fixed positions. Figure la depicts the "head" followed by 11 seg- ments with the posterior disc or "sucker" forming the 11th. The other two specimens are sufficiently contracted to hide the pos- terior segments. The reported and drawn "pair of eye-spots" are precisely where the dark jaws should be in a branchiobdellidan. Due to the fragile condition of this rare book, the author was supplied with photo- copies of only a few requested pages. It is possible that Roesel (1755) made additional references to branchiobdellidans in other parts of this volume that were not available to the author. Reason for this belief comes from a subsequent work by Moquin-Tandon (1846) who refers to Roesel's finding very VOLUME 109, NUMBER 4 655 la Fig. 1. Three preserved worms from the "river crab," redrawn from Roesel (1755: Tab. LIX, figs. 20, 21, and 22, respectively) at approximately 6.5 X. small hirudineans, Branchiobdella astaci, which were parasitic in the gills of crawfish. An improved description of a "crab worm," Hirudo parasita was made by Braun (1805), followed by a second species, Hirudo astaci Miiller, 1 806. However, it was the detailed observations on Branchiobdella astaci Odier, 1823, and presumably the same name as used by Roesel that resulted in this being accepted as the type species for the newly created genus. For completeness, it should be noted that Vejdovsky (1884) did not agree with Odier and recognized B. par- asita Braun, 1805, as the type species. Unfortunately, in the early 1800's, Blain- ville was already incorrectly using Bran- chiobdella for the gilled leeches, as noted earlier. Therefore, to accommodate the bran- chiobdellidans, he created a new generic name Microbdella Blainville, 1827. This name was modified to Hirudo {microbdella) astaci by Gervais (1836:628), and later (Gervais 1845) to Microbdella astaci. The problem of consistency with the genus name continued when Vallot (1840) proposed As- tacobdella branchiate for B. parasita. In his checklist of worms, Diesing (1850) accepted the former generic name for his two new species, Astacobdella Roeseli Diesing, 1 850, and Astacobdella Abildgaardi Diesing, 1850. Continuing this trend, Astacobdella Philadelphica Leidy, 1851, became the first reported branchiobdellidan in North Ameri- ca. After 1851, all branchiobdellidans were placed in the genus Branchiobdella, until Moore (1895 a) recognized that specimens found in North America were sufficiently different from Branchiobdella to warrant forming a number of additional genera. The century closed with Voigt (1885) arguing that the four know species of Branchiobdel- la in Europe were variants of a single spe- cies, B. varians. Although a few of his con- temporaries used the name, it was not gen- erally accepted. The last monograph on the branchiob- dellidans was produced by Pierantoni (1912). Since then a number of reviews, checklists, and keys to species have been produced, but they have been restricted to one of the three geographical regions in the Holarctic where the symbionts are found. These regions are North America, from Panama to southern Canada inclusive (Hoffman 1963; Holt 1965a, 1968a, 1973a, 1974a, 1986), Europe, from the Atlantic coast to the Ural Mountains (Moszyriski 1938, Pop 1965, Boshko 1983), and eastern Asia, which include southeastern Russia, Korea, Japan, and China (Yamaguchi 1934; Liu 1964, 1984; Subchev 1986; Gelder 1987; Timm 1991). Therefore, a single checklist for the taxon is long overdue. The largest contribution to branchiobdel- lidan taxonomy has been made by Holt (see Holt & Opell 1993), whose primary focus was on North American species. His phi- losophy (Holt pers. comm.) was that if a group of individuals has a recognizable an- atomical structure, or a structure with a dif- ferent relative proportion to that in a de- scribed species, then the new group should 656 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON be described as a new species. This philos- ophy was also shared by Yamaguchi (1934). However, European workers such as Moszyhski (1938), Pop (1965), and Kar- aman (1967) recognized that such a level of difference warranted only subspecies rank. Therefore, this dichotomy in what consti- tutes a species in the branchiobdellidans needs to be addressed in the future. This checklist contains the binomial names and junior synonyms of all bran- chiobdellidans with the author and date of the original description. The endings of a number of species binomen have been changed since the type descriptions were pubhshed (see Holt 1973a, Holt & Opell 1993). However, Holt & Opell (1993) changed the specific name endings of three species of Cambarincola from "us" back to the original "a" without supporting the change. These reversals have not been ac- cepted in this checklist. The change of Cambarincola osceola to osceolia by Holt (1973:10) is not supported by the current International Code of Zoological Nomen- clature (ICZN, 1985) in the opinion of F. M. Bayer (pers. comm.), and so the former spelling is accepted. In order to make this checklist more user-friendly, some species citations are followed by an author's name and a date. This citation indicates the paper in which the preceding name was modified or changed. Researchers studying type de- scriptions will become aware of the occa- sional anomaly where information in the checklist differs from the original paper. Two of these are explained to prevent con- fusion. The description of Branchiobdella balcanica sketi Karaman, 1967, was repub- lished as a new subspecies by Karaman (1970), but the former is the authentic ci- tation. Some papers have a year printed at the head of the pages, but due to delays, the journals were actually published a year later than stated. The date of publication is the only correct one, for example; Moore (1894) has the previous year imprinted on its pages. Ellis (1918) reported a number of branchiobdellidans on Michigan crawfishes. and a number of the species cited were fol- lowed by, "m ed." The type descriptions of these new genera and species were pub- lished in Ellis (1919). The discrepancy in time was caused by an editorial delay of the manuscript with the new taxa, which did not effect the publication or distribution of the paper. The correct type citations for these taxa are Ellis (1919). Taxonomic keys require a number of ex- ternal and internal anatomical characters to be observed before the specimen can be identified. Many of the characters are not easy to observe, so there has been a temp- tation to use only the obvious characters of the external body shape, body length, and jaw structure. Such an approach invariably ignores the differing effect that fixation anomalies can produce on a soft body, and the apparently different morphology of the jaws when seen from various aspects. As a result of the variability of these few char- acters, 33 new species were described by Georgevitch (1955, 1957) that have since been reduced to junior synonyms of four species by Pop (1965). Sinodrilus heterorchis (Liu & Zhang, 1983) has a pair of separately opening an- terior nephridiopores which makes it a member of either the Branchiobdellidae or the Xironodrilidae. However, the type de- scription did not state where the vas defer- ens enters the glandular atrium; without this information, it is not possible to assign this genus to a family. Therefore, the valid ge- nus and species of Sinodrilus heterorchis has been placed in incertae sedis until the missing information is obtained. In addi- tion, two species are considered nomen in- quirenda and three other species nomen nu- dem in the checklist. Checklist Class Branchiobdellae Kasprzak, 1984 Order BRANCHIOBDELLIDA Holt, 1965a Family Branchiobdellidae (Grube, 1851) Ankyrodrilus Holt, 1965b Ankyrodrilus koronaeus Holt, 1965b Ankyrodrilus legaeus Holt, 1965b VOLUME 109, NUMBER 4 657 Branchiobdella Odier, 1823 Branchiobdella astaci Odier, 1823 = Astacobdella Roeseli (Diesing, 1850) = Branchiobdella astaci fluviatilis (Ostroumoff, 1883) = Branchiobdella astaci leptodactyli (Ostrou- moff, 1883) = Branchiobdella varians var. astaci (Voigt, 1885) Pierantoni, 1912 = Hirudo astaci (Muller, 1806) Vejdovsky 1884 = Hirudo (Microbdella) astaci (Gervais, 1836) Vejdovsky, 1884 = Malacobdella (Hirudo) Astaci (Gervais, 1836) Diesing 1850 = Microbdella astaci (Gervais, 1845) Vejdovsky 1884 Branchiobdella balcanica balcanica Moszyriski, 1938 = Branchiobdella pentodonta orientalis (Pop, 1965) Karaman 1967 = Xironogiton dolicoderos (Georgevitch, 1957) Pop 1965 Branchiobdella balcanica sketi Karaman, 1967 Branchiobdella cheni Liu, 1964 Branchiobdella digitata Pierantoni, 1906 Branchiobdella domina Timm, 1991 Branchiobdella hexodonta Gruber, 1883 = Branchiobdella dubia (Pierantoni, 1912) Pop 1965 = Branchiobdella ochridensis (Georgevitch, 1957) Pop 1965 = Branchiobdella varians var. hexodonta (Voigt, 1885) Pierantoni 1912 = Cambarincola pluridentata (Georgevitch, 1957) Pop 1965 Branchiobdella italica Canegallo, 1928 = Branchiobdella insolita (Moszyriski, 1938) Pop 1965 = Branchiobdella pentodonta var. italica (Cane- gallo, 1928) Pop 1965 Branchiobdella kobayashii Yamaguchi, 1934 Branchiobdella kozarovi Subchev, 1978 Branchiobdella macroperistomium Liu & Zhang, 1983 Branchiobdella minuta Pierantoni, 1912 Branchiobdella monodontus Liu & Zhang, 1983 Branchiobdella orientalis Yamaguchi, 1934 Branchiobdella parasita Henle, 1835 = Astacobdella branchialis (Vallot, 1840) Vej- dovsky 1884 = Branchiobdella anatis (Pierantoni, 1912) Pop 1965 = Branchiobdella bidens (Georgevitch, 1955) Pop 1965 = Branchiobdella decidonta (Georgevitch, 1957) Pop 1965 = Branchiobdella tridens (Georgevitch, 1955) Pop 1965 = Branchiobdella varians var. parasita (Voigt, 1885) Pierantoni 1912 = Cambarincola cylindrica (Georgevitch, 1957) Pop 1965 = Cambarincola odontias (Georgevitch, 1955) Pop 1965 = Hirudo Parasita (Braun, 1805) Henle 1835 = Pterodrilus aliata (Georgevitch, 1957) Pop 1965 = Pterodrilus bidens (Georgevitch, 1955) Pop 1965 = Pterodrilus dentata (Georgevitch, 1957) Pop 1965 = Pterodrilus karamani (Moszyriski, 1938) Pop 1965 = Pterodrilus megas (Georgevitch, 1955) Pop 1965 = Pterodrilus megodont (Georgevitch, 1955) Pop 1965 = Pterodrilus prion (Georgevitch, 1955) Pop 1965 = Xironogiton bidens (Georgevitch, 1955) Pop 1965 Branchiobdella pentodonta Whitman, 1882 = Bdellodrilus hexodonta (Georgevitch, 1957) Pop 1965 = Branchiobdella capito (Georgevitch, 1955) Pop 1965 = Branchiobdella cordis (Georgevitch, 1955) Pop 1965 = Branchiobdella karamani (Georgevitch, 1957) Pop 1965 = Branchiobdella segmentata (Georgevitch, 1957) Pop 1965 = Branchiobdella septadonta (Georgevitch, 1957) Pop 1965 = Branchiobdella unidonta (Georgevitch, 1957) Pop 1965 = Branchiobdella varians var. pentodonta (Voigt, 1885) Pierantoni 1912 = Cambarincola dojranensis (Georgevitch, 1955) Pop 1965 = Cambarincola gastrax (Georgevitch, 1955) Pop 1965 = Cambarincola hamata (Georgevitch, 1957) Pop 1965 = Cambarincola odontias (Georgevitch, 1955) Pop 1965 = Xironodrilus crassus (Georgevitch, 1955) Pop 1965 = Xironodrilus tetradonta (Georgevitch, 1955) Pop 1965 = Xironogiton dilatatus (Georgevitch, 1955) Pop 1965 = Xironogiton dolicoderos (Georgevitch, 1955) Pop 1965 = Xironogiton latus (Georgevitch, 1957) Pop 1965 658 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON = Xironogiton tridens (Georgevitch, 1957) Pop 1965 Branchiohdella teresae Subchev, 1986 Cirrodriliis Pierantoni, 1905 Cirrodrilus aequiannuhis (Liu, 1984) = Stephanodrilus aequiannulus Liu, 1984 Cirrodrilus anodontus (Liu, 1964) = Stephanodrilus anodontus Liu, 1964 Cirrodrilus aomorensis (Yamaguchi, 1934) = Stephanodrilus {St.) aomorensis Yamaguchi, 1934 Cirrodrilus breviformis (Liu & Chang, 1964) = Stephanodrilus breviformis Liu & Chang, 1964 Cirrodrilus chosen (Yamaguchi, 1934) = Stephanodrilus (St.) chosen Yamaguchi, 1934 Cirrodrilus cirratus Pierantoni, 1905 = Ceratodrilus cirratus (Yamaguchi, 1932a) Holt 1967a = Stephanodrilus (Ceratodrilus) cirratus (Pieran- toni, 1905) Yamaguchi 1934 Cirrodrilus ezoensis (Yamaguchi, 1934) = Stephanodrilus (St.) ezoensis Yamaguchi, 1934 Cirrodrilus Jimbriatus Timm, 1991 Cirrodrilus heteroglandularis (Liu & Chang, 1964) = Stephanodrilus heteroglandularis Liu & Chang, 1964 Cirrodrilus homodontus (Yamaguchi, 1932b) = Cambarincola homodontus Yamaguchi, 1932b Cirrodrilus inukaii (Yamaguchi, 1934) = Stephanodrilus {St.) inukaii Yamaguchi, 1934 Cirrodrilus japonicus (Pierantoni, 1912) = Stephanodrilus japonicus Pierantoni, 1912 Cirrodrilus kawamurai (Yamaguchi, 1934) = Stephanodrilus (St.) kawamurai Yamaguchi, 1934 Cirrodrilus liaoningensis (Liu & Chang, 1964) = Stephanodrilus liaoningensis Liu & Chang, 1964 Cirrodrilus makinoi (Yamaguchi, 1934) = Stephanodrilus {St.) makinoi Yamaguchi, 1934 Cirrodrilus megalodentatus (Yamaguchi, 1934) = Stephanodrilus {St.) magalodentatus Yamagu- chi, 1934 Cirrodrilus minimus (Liu, 1964) = Stephanodrilus minimus Liu, 1964 Cirrodrilus nipponicus (Yamaguchi, 1932c) = Carcinodrilus nipponicus Yamaguchi, 1932c Cirrodrilus peristomalis (Liu & Chang, 1964) = Stephanodrilus peristomalis Liu & Chang, 1964 Cirrodrilus pugna.x Timm, 1991 Cirrodrilus quadritentacularis (Liu, 1984) = Stephanodrilus quadritentacularis Liu, 1984 Cirrodrilus sapporensis (Pierantoni, 1906) = Stephanodrilus sapporensis Pierantoni, 1906 Cirrodrilus suzukii (Yamaguchi, 1934) = Stephanodrilus {St.) suzukii Yamaguchi, 1934 Cirrodrilus uchidai (Yamaguchi, 1932a) = Ceratodrilus Uchidai Yamaguchi, 1932a = Stephanodrilus {Ceratodrilus) uchidai (Yama- guchi, 1932a) Yamaguchi, 1934 Xironogiton Ellis, 1919 Xironogiton cassiensis Holt, 1974a Xironogiton fordi Holt, 1 974a Xironogiton instabilis (Moore, 1894) Holt 1974a = Bdellodrilus instabilis (Moore, 1 894) Pierantoni 1912 = Branchiohdella instabilia (Moore, 1894) Ellis 1919 = Xironogiton instabilius instabilius (Goodnight, 1940) Holt 1974a = Xironogiton instabilius oregonensis (Good- night, 1940) Holt 1974a = Xironogiton oregonensis oregonensis (Ellis, 1919) Holt 1974a = Xironogiton oregonensis pectinatus (Ellis, 1919) Holt 1974a Xironogiton kittitasi Holt, 1974a Xironogiton occidentalis Ellis, 1919 Xironogiton victoriensis Gelder & Hall, 1990 Family Bdellodrilidae Holt, 1986 Bdellodrilus Moore, 1895b Bdellodrilus illuminatus (Moore, 1894) = Branchiohdella illuminata (Moore, 1894) Cronodrilus Holt, 1968b Cronodrilus ogygius Holt, 1968b Hidejiodrilus Gelder & Brinkhurst, 1990 Hidejiodrilus koreanus (Pierantoni, 1912) = Stephanodrilus koreanus (Pierantoni, 1912) Uglukodrilus Holt, 1989b Uglukodrilus hemophagus (Holt, 1977a) = Adenodrilus hemaphagus Holt, 1977a Family Caridinophilidae Holt, 1986 Caridinophilus Liang, 1963 Caridinophilus unidens Liang, 1963 Family Cambarincolidae Holt, 1986 Cambarincola Ellis, 1912 Cambarincola acudentatus Holt, 1973a Cambarincola alienus Holt, 1963 Cambarincola barbarae Holt, 1981a Cambarincola bobbi Holt, 1988a Cambarincola branchiophilus Holt, 1954 Cambarincola carcinophilus Holt, 1973a Cambarincola chirocephalus Ellis, 1919 = Cambarincola {Coronata) chirocephala (Ellis, 1919) Goodnight 1940 Cambarincola demissus Hoffman, 1963 Cambarincola dubius Holt, 1973b Cambarincola ellisi Holt, 1973a VOLUME 109, NUMBER 4 659 Cambarincola fallax Hoffman, 1963 Cambarincola floridanus Goodnight, 1941 Cambarincola goodnighti Holt, 1973c Cambarincola gracilis Robinson, 1954 Cambarincola heterognathus Hoffman, 1963 Cambarincola hojfmani Holt, 1973a Cambarincola holostomus Hoffman, 1963 Cambarincola holti Hoffman, 1963 Cambarincola illinoisensis Holt, 1982 Cambarincola ingens Hoffman, 1963 Cambarincola jamapaensis Holt, 1973a Cambarincola leoni Holt, 1973b Cambarincola leptadenus Holt, 1973b Cambarincola macrocephalus Goodnight, 1943 Cambarincola macrodontus Ellis, 1912 = Cambarincola {Cambarincola) macrodontus (Ellis, 1912) Goodnight 1940 Cambarincola manni Holt, 1973c Cambarincola marthae Holt, 1973b Cambarincola mesochoreus Hoffman, 1963 Cambarincola meyeri Goodnight, 1942 Cambarincola micradenus Holt, 1973a Cambarincola montanus (Goodnight, 1940) = Triannulata montana (Goodnight, 1940) Holt 1974b Cambarincola nanagnathus Holt, 1973a Cambarincola olmecus Holt, 1973a Cambarincola osceola Hoffman, 1963 Cambarincola ouachita Hoffman, 1963 Cambarincola pamelae Holt, 1984a Cambarincola philadelphicus (Leidy, 1851) = Astacobdella Philadelphica (Leidy, 1851) = Branchiobdella americana (Pierantoni, 1912) Holt 1967a = Cambarincola (Coronata) philadelphica (Lei- dy, 1851) Goodnight 1940 Cambarincola restans Hoffman, 1963 Cambarincola serratus Holt, 1981a Cambarincola sheltensis Holt, 1973b Cambarincola shoshone Hoffman, 1963 Cambarincola smalleyi Holt, 1964 Cambarincola speocirolanae Holt, 1984b Cambarincola steevesi Holt, 1973b Cambarincola susanae Holt, 1973a Cambarincola toltecus Holt, 1973a Cambarincola virginicus Hoffman, 1963 Cambarincola vitreus Ellis, 1919 = Cambarincola {Cambarincola) vitrea (Ellis, 1919) Goodnight 1940 Ceratodrilus Hall, 1914 Ceratodrilus orphiorhysis (Holt, 1960a) Holt 1988b Ceratodrilus thysanosomus Hall, 1914 = Cirrodrilus thysanosomus (Hall, 1914) Good- night 1940 Ellisodrilus Holt, 1960b Ellisodrilus carronamus Holt, 1988a Ellisodrilus clitellatus Holt, 1960b Ellisodrilus durbini (Ellis, 1919) = Pterodrilus durbini (Ellis, 1919) Holt 1960b Magmatodrilus Holt, 1967a Magmatodrilus obscurus (Goodnight, 1940) = Stephanodrilus obscurus (Goodnight, 1940) Holt 1967a Oedipodrilus Holt, 1967b Oedipodrilus anisognathus Holt, 1988a Oedipodrilus cuetzalanae Holt, 1984b Oedipodrilus macbaini (Holt, 1955) = Cambarincola macbaini (Holt, 1955) Holt 1969 Oedipodrilus oedipus Holt, 1967b Pterodrilus Moore, 1895a Pterodrilus alcicornus Moore, 1895a Pterodrilus annulatus Gelder, 1996 Pterodrilus cedrus Holt, 1968c Pterodrilus choritonamus Holt, 1968c Pterodrilus distichus Moore, 1895a Pterodrilus hobbsi Holt, 1968c Pterodrilus mexicanus Ellis, 1919 Pterodrilus missouriensis Holt, 1968c Pterodrilus simondsi Holt, 1968c Sathodrilus Holt, 1968b Sathodrilus attenuatus Holt, 1981b Sathodrilus carolinensis Holt, 1968b Sathodrilus chehalisae Holt, 1981b Sathodrilus dorfus Holt, 1977b Sathodrilus elevatus (Goodnight, 1940) = Cambarincola elevatus (Goodnight, 1940) Holt 1978 = Cambarincola {Cambarincola) elevatus (Good- night, 1940) Hoffman 1963 Sathodrilus hortoni Holt, 1973c Sathodrilus inversus (Ellis, 1919) = Cambarincola inversa (Ellis, 1919) = Cambarincola {Cambarincola) inversa (Ellis, 1919) Goodnight 1940 = Sathodrilus virgiliae (Holt, 1977b) Holt 1981b Sathodrilus lobatus Holt, 1977b Sathodrilus megadenus Holt, 1968b Sathodrilus nigrofluvius Holt, 1989b Sathodrilus norbyi Holt, 1977b Sathodrilus okaloosae Holt, 1973c Sathodrilus prostates Holt, 1973a Sathodrilus rivigeae Holt, 1988a Sathodrilus shastae Holt, 1981b Sathodrilus veracruzicus Holt, 1968b Sathodrilus villalobosi Holt, 1968b Sathodrilus wardinus Holt, 1981b Tettodrilus Holt, 1968b Tettodrilus friaufi Holt, 1968b Triannulata Goodnight, 1940 Triannulata magna Goodnight, 1940 Family Xironodrilidae Holt, 1986 Holtodrilus Gelder & Brinkhurst, 1990 660 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Holtodrilus tnmcotus (Liang, 1963) Xironodrilus Ellis, 1919 Xironodrilus appalachius Goodnight, 1943 Xironodrilus bashariae Holt & Weigl, 1979 Xironodrilus dentatus Goodnight, 1940 = Xironodrilus pulcherrimus dentatus (Good- night, 1940) Goodnight 1943 Xironodrilus formosus Ellis, 1919 Xironodrilus pulcherrimus (Moore, 1894) = Branchiobdella pulcherrima (Moore, 1894) El- lis 1919 Incertae sedis: Sinodrilus heterorchis (Liu & Zhang, 1983) Gelder & Brinkhurst 1990 Nomen inquirenda: Branchiobdella tetradonta Pierantoni, 1912 (Holt 1967a:7) Cambarincola okadai Yamaguchi, 1933 (Holt & Opell 1993:253) Nomen nudem: Astocobdella Abildgaard Diesing, 1850:434 — inad- equate description. Bdellodrilus manus n. sp. cited in Moore 1895a: 454 — no description. Branchiobdella chilensis Moquin-Tandon, 1846: 300 — no description. = Astacobdella chilensis Diesing, 1850 Branchiobdella auriculae Moquin-Tandon, 1846: 301 — no description. Acknowledgments Many thanks for the help, tolerance, and understanding shown by Greg Curtis and Virginia Fisher at the library. University of Maine at Presque Isle, Maine; Dr. Jon L. Norenburg, Cheryl Bright and William E. Moser at the National Museum of Natural History, Smithsonian Institution, Washing- ton D.C.; Valerie Wheat, American Muse- um of Natural History, New York; Skip Sterener, University of Nebraska State Mu- seum at Lincoln, Nebraska, U.S.A.; Dr. Marco Ferraguti, Dipartimento di Biologia, Universita degli Studi di Milano, Italy and Drs Joseph B. Jennings and Richard A. Ba- ker, University of Leeds, U.K., to my re- quests for assistance at the most inoppor- tune times. Thanks are also due the Dr. An- drea M. Gorman and Mark J. Wetzel for their constructive critism of the manuscript, and Andreas Schneider for translating the archaic German texts. Literature Cited Blainville, M. H. D. de. 1827. Sangsue, Hirudo. 47: 205-273 in J. B. de Lamarck, Historic naturelle des Animaux sans Vertebres. Paris. . 1828. "Vermes'". 57:365-625 /« J. B. de La- marck, Historic naturelle des Animaux sans Vertebres. Paris. Blanchard, R. 1894. Sur les Branchellion des Mers d' Europe. — Bulletin de la Socicte Zoologique dc France 19:85-88. Boshko, E. G. 1983. Small worms dwelling on river crayfish of some water bodies in the Ukraine. — Vestnik zoologii 7:13-20. Braun, J. F. P. 1805. Systcmatische Bcschrcibung ei- niger Egelarten Gottlieb Wilhelm Miiller, Ber- lin, 75 pp. Brinkhurst, R. O., & S. R. Gelder. 1991. Annelida: Oligochacta and Branchiobdellida. Pp. 401-435 in J. H. Thorpe & A. P. Covich, cds.. Ecology and classification of North American freshwater invertebrates. Academic Press, New York. Canegallo, M. A. 1928. Una nuova specie di Bran- chiohdella-Branchiobdella italica. — Atti della Societa Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano 67:214- 224. Diesing, C. M. 1850. Systema Helminthum. vol. I., Academiae Caesareae Scientiarum, Vindobon- ae, 980 pp. Ellis, M. M. 1912. A new discodrilid worm from Col- orado. — Proceedings of the United States Na- tional Museum 42:481-486. . 1918. Branchiobdellid worms (Annelida) from Michigan crawfishes. — Transactions of the American Microscopical Society 37:49-51. . 1919. The branchiobdellid worms in the col- lections of the United States National Museum with descriptions of new genera and new spe- cies. — Proceedings of the United States Nation- al Museum 55:241-265. Gelder, S. R. 1987. Observations on three species of branchiobdellid (Annelida: Clitellata) worms from eastern Asia. — Hydrobiologia 155:15-25. . 1996. Description of a new branchiobdelli- dan species, with observations on three other species, and a key to the genus Pterodrilus (An- nelida: Clitellata). — Proceedings of the Biolog- ical Society of Washington 109:256-263. , & R. O. Brinkhurst. 1990. An assessment of the phylogeny of the Branchiobdellida (Anne- lida: Clitellata) using PAUP. — Canadian Journal of Zoology 68:1318-1326. , & L. A. Hall. 1990. Branchiobdellidans from southwestern British Columbia, Canada with a phylogenetic analysis of Xironogiton (Annelida: Clitellata) using PAUP. — Canadian Journal of Zoology 68:2352-2359. VOLUME 109, NUMBER 4 661 d( h ei Gervais, P. 1836. P. 627 in Guerin, ed., Dictionnaire pittoresque d'histoire naturelle, et des ph'eno- m'enes de la nature. Impressione de Cosson. Paris. . 1845. Patria zoologie, France, p. 572. Georgevitch, J. 1955. Sur les Branchiobdellides des ecrevisses du Lac Dojran. — Acta Musei Mace- donici Scientiarum Naturalium, Skopje 2(10/ 21):199-221. 1957. Les Branchiobdellides de Yougosla- vie. — Bulletin de I'Academie serbe des Sci- ences (new serise 18), Classe des Sciences mathematiques et naturelles 5:5-25. Goodnight, C. J. 1940. The Branchiobdellida (Oli- gochaeta) of North American crayfish. — Illinois Biological Monographs 17:1-75. . 1941. The Branchiobdellidae (Oligochaeta) of Florida. — Transactions of the American Mi- croscopical Society 60:69-74. . 1942. A new species of branchiobdellid from Kentucky. — Transactions of the American Mi- croscopical Society 61:272-273. . 1943. Report on a collection of branchiob- dellids. — Journal of Parasitology 29:100-102. Grube, A. E. 1851. Die Familien der Anneliden. — Verlag der Nicolai'schen Buchhandlung, Berlin, 160 pp. Gruber, A. 1883. Bemerkungen uber die Gattung Branchiobdella. — Zoologischer Anzeiger 6: 243-248. Hall, M. C. 1914. Descriptions of a new genus and species of the discodrilid worms. — Proceeding of the United States National Museum 48:187- 193. Henle, J. F G. 1835. Ueber die Gattung Branchiob- della und uber die Deutung der inneren Ges- chlechstheile bei den Annelidan und hermap- phroditischen Schnecken. — Archiv fur Anato- mic, Physiologic, und wissenschaftliche Medi- zin: 574-608. Hoffman, R. L. 1963. A revision of the North Amer- ican annelid worms of the genus Cambarincola (Oligochaeta: Branchiobdellidae). — Proceed- ings of the United States National Museum 1 14: 271-371. Holt, P C. 1954. A new branchiobdellid of the genus Cambarincola (Oligochaeta, Branchiobdellidae) from Virginia. — Virginia Journal of Science 5: 168-172. . 1955. A new branchiobdeUidae of the genus Cambarincola Ellis, 1912 (Oligochaeta, Bran- chiobdellidae) from Kentucky. — Journal of the Tennessee Acadamy of Science 30:27-31. . 1960a. The genus Ceratodrilus Hall (Bran- chiobdellidae, Oligochaeta) with the description of a new species. — Virginia Journal of Science 11:53-77. . 1960b. On a new genus of the family Bran- chiobdellidae (Oligochaeta). — American Mid- land Naturahst 64:169-176. . 1963. A new branchiobdellid (Branchiobdel- lidae: Cambarincola). — Journal of the Tennes- see Acadamy of Science 38:97-100. — . 1964. A new branchiobdellid (Annelida) from Costa Rica. — Tulane Studies in Zoology 12:1-4. — . 1965 a. The systematic position of the Bran- chiobdellidae (Annelida: Clitellata).— System- atic Zoology 14:25-32. — . 1965b. On Ankyrodrilus, a new genus of branchiobdellid worms (Annelida). — Virginia Journal of Science 16:9-21. — . 1967a. Status of the genera Branchiobdella and Stephanodrilus in North America with de- scription of a new genus (Clitellata: Branchiob- dellida). — Proceedings of the United States Na- tional Museum 124:1-10. — . 1967b. Oedipodrilus oedipus, n.g., n.sp. (An- nelida, Clitellata: Branchiobdellida). — Transac- tions of the American Microscopical Society 86:58-60. — . 1968a. The Branchiobdellida: epizootic an- nelids. — Biologist 50:79-94. — . 1968b. New genera and species of branchiob- dellid worms (Annelida: Clitellata). — Proceed- ings of the Biological Society of Washington 81:291-318. — . 1968c. The genus Pterodrilus (Annelida: Branchiobdellida). — Proceedings of the United States National Museum 125:1-44. — . 1969. The relationship of the branchiobdellid fauna of the southern Appalachians. Pp. 191- 219 /« Perry C. Holt, ed.. The distributional his- tory of the biota of the southern Appalachians, Part I: Invertebrates. Virginia Polytechnic Insti- tute, Blacksburg, Research Division Mono- graph, vol. 1. — . 1973a. A summary of the branchiobdellid (Annelida: Clitellata) fauna of Mesoamerica. — Smithsonian Contributions to Zoology 142:1- 40. — . 1973b. Branchiobdellids (Annelida: Clitella- ta) from some eastern North American caves, with descriptions of new species of the genus Cambarincola. — International Journal of Spe- leology 5:219-256. — . 1973c. Epigean branchiobdellids (Annelida: Clitellata) from Florida. — Proceedings of the Biological Society of Washington 86:79-104. — . 1974a. The genus Xironogiton Ellis, 1919 (Clitellata: Branchiobdellida). — Virginia Jour- nal of Science 25:5-19. — . 1974b. An emendation of the genus Trian- nulata Goodnight, 1940, with the assignment of Triannulata montana to Cambrincola Ellis 1912 (Clitellata: Branchiobdellida).— Proceed- 662 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ings of the Biological Society of Washington 87:57-72. — . 1977a. A gill-inhabiting new genus and spe- cies of the Branchiobdellida (Annelida: Clitel- lata). — Proceedings of the Biological Society of Washington 90:726-734. — . 1977b. An emendation of genus Sathodrilus Holt 1968 (Annelida: Branchiobdellida), with the description of four new species from the Pa- cific drainage of North America. — Proceedings of the Biological Society of Washington 90: 116-131. — . 1978. The reassignment of Cambarincola elevatiis Goodnight, 1940 (Clitellata: Bran- chiobdellida) to the genus Sathodrilus Holt, 1968. — Proceedings of the Biological Society of Washington 91:472-483. — . 1981a. A resume of the members of the ge- nus Cambarincola (Annelida: Branchiobdelli- da) from the Pacific drainage of the United States. — Proceedings of the Biological Society of Washington 94:675-695. — . 1981b. New species of Sathodrilus Holt, 1968 (Clitellata: Branchiobdellida) from the Pa- cific drainage of the United States, with the syn- onymy of Sathodrilus virgiliae Holt, 1977. — Proceedings of the Biological Society of Wash- ington 94:848-862. — . 1982. A new species of the genus Cambar- incola (Clitellata: Branchiobdellida) from Illi- nois with remarks on the bursa of Cambarin- cola vitreus Ellis, 1919, and the status of Sath- odrilus Holt, 1968. — Proceedings of the Biolog- ical Society of Washington 95:251-255. — . 1984a. A new species of the genus Cambar- incola (Clitellata: Branchiobdellida) from Cali- fornia. — Proceedings of the Biological Society of Washington 97:544-549. — . 1984b. On some branchiobdellids (Annelida: Clitellata) from Mexico with the descriptions of new species of the genera Cambarincola and Oedipodrilus. — Proceedings of the Biological Society of Washington 97:34-42. — . 1986. Newly established families of the order Branchiobdellida (Annelida: Clitellata) with a synopsis of the genera. — Proceedings of the Bi- ological Society of Washington 99:676-702. — . 1988a. Four new species of cambarincolids (Clitellata: Branchiobdellida) from the south- eastern United States with a redescription of Oedipodrilus macbaini (Holt. 1955). — Proceed- ings of the Biological Society of Washington 101:794-808. — . 1988b. The correct name of Ceratodrilus or- phiorhysis Holt, 1960 (Annelida: Branchiobdel- lida). — Proceedings of the Biological Society of Washington 101:308. . 1989a. Comments on the classification of the Clitellata.— Hydrobiologia 180:7-15. . 1989b. A new species of the Cambarincolid genus Sathodrilus from Missouri, with the pro- posal of a replacement name for Adenodrilus Holt, 1977 (Clitellata: Branchiobdellida).— Pro- ceedings of the Biological Society of Washing- ton 102:738-741. , & A. M. Weigl. 1979. A new species of Xir- onodrilus Ellis 1918 from North Carolina (Cli- tellata: Branchiobdellida). — Brimleyana 1:23- 29. , & B. D. Opell. 1993. A checklist of and il- lustrated key to the genera and species of the Central and North American Cambarincolidae (Clitellata: Branchiobdellida). — Proceedings of the Biological Society of Washington 106:251- 295. International Commission on Zoological Nomencla- ture. 1985. International code of zoological no- menclature adopted by the XX General Assem- bly of the International Union of Biological Sci- entists. International Trust for Zoological No- menclature, London, 338 p. Karaman, S. M. 1967. Branchiobdelldae Jugoslavije (Annelida: Clitellata). — Buletin i Punimeve Shkencore te Fakultetit Filozofik te Prishtines IV, 39-63. . 1970. Beitrag zur Kenntnis der europaischen Branchiobdelliden (Clitellata, Branchiobdelli- dea). — Internationale Revue der gesamten Hy- drobiologie 55:325-333. Kasprzak, K. 1984. The previous and contemporary conceptions on phylogeny and systematic clas- sifications of Oligochaeta (Annelida). — Anneles Zoologici (Warsaw) 38:205-223. Leidy, J. 1851. A description of Astacobdella phila- delphica. — Proceedings of the Academy of Nat- ural Sciences of Philadelphia 5:209. Liang, Y.-L. 1963. Studies on the aquatic Oligochaeta of China. I. Descriptions of new naids and bran- chiobdellids. — Acta Zoologica Sinica 15:560- 570. Ludwig, H. 1886. Dr. Johannes Leunis Synopsis der Thierkunds. Hahnbsche buchhandlung, Hanno- ver, vol. 2, 1232 pp. Liu, S.-C. 1964. A second report of Branchiobdellidae in Liaoning Province with descriptions of three new species. — Acta Zoologica Sinica 16:602— 610. . 1984. Descriptions of two new species of the genus Stephanodrilus from Northeast China and notes on St. trunctus Liang from Guagdong Province (Oligochaeta: Branchiobdellidae). — Acta Zootaxonomica Sinica 9:351—355. , & C.-A. Chang. 1964. On four new species of Branchiobdellidae from crayfish, Cambaro- VOLUME 109, NUMBER 4 663 ides dauricus (Pallas). — Acta Zoologica Sinica h 16:33-38. -, & D.-C. Zhang. 1983. Three new species of the genus Branchiobdella (Oligochaeta: Bran- chiobdellidae) from China. — Acta Zootaxon- omica Sinica 8:246-251. Moore, J. P. 1894. On some leech-like parasites of American crayfishes. — Proceedings of the Academy of Natural Sciences of Philadelphia p. 419-428. ^ . 1895a. Pterodrilus, a remarkable discodril- ^^L id. — Proceedings of the Academy of Natural Sciences of Philadelphia, p. 449-454. -. 1895b. The anatomy of Bdellodrilus illumi- natus, an American discodrilid. — Journal of Morphology 10:497-540. Moquin-Tandon, A. 1846. Monographic de la famille des Hirudinees. J.-B. Bailliere, Paris, 448 pp. Moszynski, A. 1938. Oligochetes parasites de I'ecrev- isse {Potamobius astacus L.) de la Yougosla- vie. — Bulletin de la Societe Scientifique de Skoplje 18:69-75. Miiller, O. E 1806. Zoologica danica, sev Animalivm Daniae et Norvegiae rariorvm ac minvs no- torvm descriptiones et historia. Havniae. Odier, A. 1823. Memoire sur le Branchiobdella nou- veau genre d'Annelides de la famille des Hi- rudiner. — Memoires de la Societe d'Histoire Naturelle de Paris 1:69-78. Ostroumoff, A. 1883. Ueber die Art der Gattung Branchiobdella Odier auf der Kiemer der Flus- skrebs {Astacus leptodactylus Eschh.). — Zoolo- gischer Anzeiger 6:76-78. Pierantoni, U. 1905. Cirrodrilus cirratus n.g., n.sp. parassita dell' Astacus japonicis. — Annuario del Museo Zoologico della R. Universita di Napoli (new series) 1:1-3. 1906. Nuovi Discodrilidi del Giappone e del- la California. — Annuario del Museo Zoologico deUa R. Universita di Napoli (new series) 2:1-9. 1912. Monografia dei Discodrilidae. — An- nuario del Museo Zoologico della R. Universita di Napoli (new series) 3:1-27. Pop, V. 1965. Systematische Revision der europaisch- en Branchiobdelliden (Oligochaeta). — Zoolo- gische Jahrbuecher Abteilung fiir Systematik, Oekologie und Geographic der Tiere 92:219- 238. Robinson, D. A. 1954. Cambarincola gracilis sp. nov., a branchiobdellid oligochaete commensal on western crayfishes. — Journal of Parasitology 40:466-469. Roesel von Rosenhof, A. I. 1755. Monatliche heraus- gegebene Insektenbelustigungen. — Niirenburg, 3:1-120. Savigny, J.-C. 1820. Systeme des Annelides, Ordre 4, A. Hirudinees, pp. 105-120. Sawyer, R. T. 1986. Leech Biology and Behaviour. Clarendon Press, Oxford, 1065 pp. Stuart, J. 1982. Hirudinoidea. R 43 in S. P Parker, ed.. Synopsis and classification of living organisms. McGraw-Hill, New York. Subchev, M. A. 1978. A new branchiobdellid — Bran- chiobdella kozarovi sp.n. (Oligochaeta, Bran- chiobdellidae) from Bulgaria. — Acta zoologica Bulgarica 9:78-80. . 1986. On the Korean branchiobdellids (An- nelida: Clitellata) with a description of a new species — Branchiobdella teresae sp.n. — Acta zoologica Bulgarica 31:60-66. Timm, T. 1991. Branchiobdellida (Oligochaeta) from the farthest South-East of the U.S.S.R. — Zool- ogica Scripta 20:321-331. Vallot, J. N. 1840. Memoire sur V Astacobdella bran- chiale Actes de 1' Academic des Sciences de Bordeaux 2, 3. trim., p. 483. Vejdovsky, F. 1884. System und Morphologic der Oli- gochaeten.-F. Rivnac, Prague, 39 pp. Voigt, W. 1885. Untersuchungen iiber die Varietaten- bildung bei Branchiobdella varians. — Arbeiten aus dem zoologisch-zootomischen Institut in Wurzburg 7:41-94. Whitman, C. O. 1882. A new species of Branchiob- della. — Zoologisher Anzeiger 5:636-637. Yamaguchi, H. 1932a. On the genus Cirrodrilus Pier- antoni, 1905 with a description of a new bran- chiobdellid from Japan. — Annotationes Zoolo- gicae Japonenses 13:361—367. . 1932b. A new species of Cambarincola, with remarks on spermatic vesicles of some bran- chiobdellid worms. — Proceedings of the Impe- rial Academy, Japan 8:454-456. . 1932c. Description of a new branchiobdellid, Carcinodrilus nipponicus n.g. et n.sp. — Journal of the Faculty of Science Hokkaido University, series VI, Zoology 2:61-67. . 1933. Description of a new branchiobdellid, Cambarincola okadai n.sp., parasitic on Amer- ican crayfish transferred into a Japanese lake. — Proceedings of the Imperial Academy, Japan 9: 191-193. . 1 934. Studies on Japanese Branchiobdellidae with some revisions on the classification. — Journal of the Faculty of Science Hokkaido University, series VI, Zoology 3:177-219.