a museum IIe nt BONNER ZOOLOGISCHE > MONOGRAPHIEN © Herausgeber: Zoologisches Forschungsmuseum Alexander Koenig, Bonn 2007 Andrei Y. Utevsky ANTARCTIC PISCICOLID LEECHES 10 mm Leibniz Gemeinschaft BONNER ZOOLOGISCHE MONOGRAPHIEN Schriftleitung / Editor Karl-L. Schuchmann Zoologisches Forschungsmuseum Alexander Koenig (ZFMK) Ornithologie Adenauerallee 160 D-53113 Bonn, Germany Email: kl.schuchmann.zfmk@uni-bonn.de Die Reihe BONNER ZOOLOGISCHE MONOGRAPHIEN (BZM) erscheint seit 1971 in regelmäßiger Folge. Publiziert werden Originalbeiträge in englischer Sprache. Die BZM nehmen vor allem solche Arbeiten auf, die zu umfangreich sind, um in der Zeitschrift BONNER ZOOLOGISCHE BEITRÄGE zu erscheinen. Schwerpunkte der BZM sind: Systematik, Taxonomie, Biogeographie, Anatomie und Evolutionsbiologie. Alle eingereichten Manuskripte werden begutachtet. Autoren werden gebeten, sich vor Manuskriptvorlage mit dem Schriftleiter in Verbindung zu setzen. Erwerb der BZM: Bibliothek, Zoologisches Forschungsmuseum Alexander Koenig (ZFMK), Adenauerallee 160, D-53113 Bonn, Germany. The peer-reviewed series BONNER ZOOLOGISCHE MONOGRAPHIEN (BZM) has existed since 1971. The BZM consist of original zoology papers too long for inclusion in our institute’s journal BONNER ZOOLOGISCHE BEITRAGE. Preferred manuscript topics are: systematics, taxonomy, biogeography, ana- tomy, and evolutionary biology. Manuscripts must be in American English. Authors are requested to contact the editor prior to manuscript submittal. The BZM may be obtained from: The Library, A. Koenig — Zoological Research Institute and Museum of Zoology, Adenauerallee 160, D-53113 Bonn, Germany. Author of this issue: Andrei Y. Utevsky Published: November 2007 Bonner zoologische Monographien Nr. 54, 80 pp. Editorial assistants of this issue: Angela Schmitz Ornés, Brian Hillcoat Cover: Moorebdellina biannulata, external and internal structure. A. Y. Utevsky. Price: EUR 36,-- TE ea \ HoOVIN 1A] VY Andrei Y Utevsky ANTARCTIC PISCICOLID LEECHES © Zoologisches Forschungsmuseum Alexander Koenig, Bonn , 2 : x. N i UTEVSKY, ANTARCTIC PISCICOLID LEECHI CONFENTS ASTER 9-65 ee a le ee en Eevee ta Ont EAST TS a a en ee ee Giosgumy ai Specie Were ee ee EEE WPMin Ghote Ieee mee N EEE boeee souks ei Pamlistony or the study of Antarctic leeches... 2... 2.0.2200 ce ec nennen nae san 3 Morphology and anatomy of piscicolid leeches ............020:0ccsvceecveceeceveuues Bropertonstorune body se ae eases ts +n sea wae on Cha haat See a te SELLER. 0 A erent EL GE ae nGn Ohad cade Abn Ge a ea ewe ees PD estes cacy Steril ee ens Ae, Ge EEE EEE aio wes [Rejgrrackinae nes ten oases aad gine wlnnn eB gs DOR HARES SRO ha Mae ee hylalle fej ay eye reds 5 co va ey ae ee en AR ee ae Bemalegreproduetyessystenn nun 2154.4 35%. 4 bas le ee en ae Cioallantie Sr a ee Secor rr ZaBiolomgompiseieolid: leeches 2). 0... 2.0. nee es ne Pde RRS DR@o || ccuoneindipreseryation nee an a 8. SYSUS TAU AT ee a eee ee SO 7 sinezuomtoniche Fat Piscieolidae gis... | asa oe su Ae ace aod Kia age Se ew Rai eye the Subfamilies of the Family Piscicolidae .....:2: ..0+0ecns sacs ea BoubtammyePomtobdellinae 2.2644. 04%s scan cae w nas nennen ana nah a Meese nn Key to the Antarctic Genera of the “Subfamily Pontobdellinae” ............:....22.2... ers nn ne ie ene ae Se ee ou A aS ek ES ES MET RN EL ENR PIL Sg TN ee er ee ee ee ee “Suiloiemnlt hy [Aleta ove lel tre | cia eta cete ener ee era ee ie aa aha Keyito.the Antarctic Genera of the “Subfamily Platybdellinae” ..........:2.......::..=- (GSMS JAI ee U ER ee GeitSPATTTORACH OS Bic pine Base Bon ais ae Es bale OE LS Oe RW She aa RRS GeMUSE I CUTOUACH ede at oe ped ho a ne Ceo SRE Se eS EO Gems PLO OLOU AEA” ok Biot uora 5 aed pend gts «HHA re Rive moe He See ea PSS Se Gene (NA IN tn Pe EO ae aa ee a oe ee EEE "Sulhlermily sera nero re ET oak boo ewe mes Cea ae TEE LETTER Kevatosthesantarctic Genera of the Subfamily Piscicolinae’ u... Sate es sans GEmUSElTichelobAeliinas: 2.2.20. 0 See hehe ee aaa eee Se ae Genser Ob dca rs. wen en oa ae Re ee ee GR NN ee RE RE MEO EAs ate ek RE EG Con eT DENE ee ee Te eee a ee ae Re (BERUSSINDLOLHENIOHAEN ARE Re Se UR RAD ON & Sees CR KS SOE 7, RUBIES: ee 1 RRO OE, Ae By Insel aes a eee re pcre a eee nae FOr SS a RIEEE N 6 vd un QS ph ed pc ee Oe Feen \A WA NO NO N NN ae a | WN NN > UTEVSKY, ANTARCTIC PISCICOLID LEECHES ABSTRACT Fish leeches (Piscicolidae) occur in continental fresh waters and in marine habitats. Some members of the family are restricted to the southern ocean only and have not been recorded from other parts of the world. This monograph is focused on this group of annelids to summarize the existing knowledge of Antarctic leeches. At present 21 species belonging to 13 genera and 3 “subfamilies” have been recorded from the Antarctic Seas. Most of them were characterized and illustrated using a uniform standard, which, in particular, includes the map of geographical distribution and a photograph. Based on the detailed descriptions, identification keys were composed. Host data were summarized in the list of all known hosts and tables with infestation rates. Key words: Antarctica, Hirudinea, Pisicolidae, Pontobdella, Moorebdellina, Megaliobdella, Epsteinia, Austro- bdella, Pleurobdella, Glyptonotobdella, Cryobdella, Trachelobdellina, Trulliobdella, Galatheabdella, Trachelobdella, Nototheniobdella, identification key. J\ BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 54/2007 ACKNOWLEDGMENTS I could not have written this monograph without the help of my colleagues and friends. My interest in leeches is due to V. M. Epshtein, the advisor of my PhD research. I am grateful to many zoologists and parasito- logists who collected and sent specimens of Antarctic leeches to me. They include A. F. Pushkin, A. B. Neelov, I. S. Smirnov, E. N. Gruzov, S. Potin, B. I. Sirenko (Zoological Institute, St. Petersburg, Russia), V. V. Avdeev (Pacific Research Institute of Fisheries and Oceanogreaphy, Vladivostok, Russia), A. A. Kovaleva, G. N. Rodiuk (Atlantic Pacific Research Institute of Fisheries and Oceanogreaphy, Kaliningrad, Russia), V. N. Ljadov (Research Institute of Biology of Southern Seas, Sevastopol, Ukraine). H. H. Janssen gave me a specimen of G/yptono- tobdella antarctica and his informative letters helped me in understanding the biology of Antarctic leeches. Stephano Chiaparelly (University of Genoa, Italy) gave me a very rich collection of Antarctic leeches with unique specimens of Megaliobdella szidati. | am very grateful to those who have given permission to repro- duce copyright material: R. T. Sawyer (Biopharm (UK) Ltd.), E. M. Burreson (Virginia Institute of Marine Science, Gloucester Point, USA) and American Geophysical Union through Michael Connolly. Special thanks go to V. V. Murina (Research Institute of Biology of Southern Seas, Sevastopol, Ukraine) and A. V. Korniushin (Schmalhausen Institute of Zoology, Kyiv, Ukraine) for useful criticisms and a good piece of advice. I am extremely grateful to V. M. Nazarov, former Head of the Department of Zoology and Animal Ecology, V. N. Karazin Kharkiv National University, Kharkiv, for his support in writing this book. My brother, Serge Utevsky (Department of Zoology and Animal Ecology, V. N. Karazin Kharkiv National University, Kharkiv, Ukraine), an expert in Arctic leeches, my wife Olga Utevskaya (Department of Genetics and Cytology, V. N. Karazin Kharkiv National University, Kharkiv, Ukraine) read drafts of the text and provided valuable criticisms. My mother, Lidiya Utevskaya, my son, Yuriy, gave me the freedom and peace of mind to spend long hours in front of journals and computer screens. This research was supported in part by the National Antarctic Scientific Center of Ministry of Education and Science of Ukraine and the Society for Underwater Activity (Kharkiv, Ukraine). UTEVSKY, ANTARCTIC PISCICOLID LEECHES GLOSSARY OF SPECIAL TERMS Accessory connection [Ac]: transverse coelomic channel connecting the testicular lacunae with the dorsal and/or ventral lacunae Accessory glands [Ag]: glandular cells covering the external surface of posterior parts of the male reproductive system Body [B]: the trachelosome and urosome without suckers Clitellum [Cl]: a posterior part of the trachelosome with gonopores on the ventral surface Conductive tissue [Ct]: a part of the female reproductive system serving for storing and passing spermatozoids to ovisacs; present in form of a mass [Ctm] and/or cord a [Ctc] Copulatory area [Ca]: external or internal, the modified ventral surface of the clitellum around one or both gonopores, or modified internal surface of the copulatory bursa Copulatory bursa [Cb]: a nonpaired terminal part of the male reproductive system terminating at the male gonopore | Crop [C]: a part of the digestive system in the testicular region of the urosome Crop chambers [Cc]: dilations of the crop between testisacs Crop region [Cr]: a part of the urosome possessing the crop Dorsal lacuna [DI]: a longitudinal coelomic channel possessing the dorsal vessel [Dv] Ejaculatory ducts (atrium) [Ed]: parts of the male reproductive system between seminal reservoirs and the copulatory bursa including paired initial [Ip], and terminal [Tp] parts, and nonpaired common part [Cp] Esophageal diverticula [Epd]: lateral dilation of the esophagus Esophagus [Ep]: part of the digestive system located between proboscis and crop Eye-like spots [Els]: light-sensitive pigmented areas on the dorsal surface of the anterior sucker Eye [E]: several integrated optic cells in pigment cup forming small symmetrical pigment zones on the dorsal surface of the anterior sucker and first annuli of the trachelosome Intestine [I]: a part of the digestive system extending from the Crop to the Anus [A] Intestinal region [Ir]: a part of the urosome possessing the intestine, the rectum and posterior crop caeca Lateral lacunae [LI]: paired longitudinal coelomic channels located under the muscular layer on both sides of the body Lateral processes [Lp]: lateral dilations of the crop, intestinal chambers and posterior crop caeca Main connection [Mc]: a part of the coelomic system in the ganglion region [Gr] of somite Marginal flanges [Mf]: external laterally located fin-like folds Marginal lacunae [MI]: paired longitudinal subepidermal coelomic channels on both sides of the body Mouth-pore [Mp]: aperture through which the proboscis everts Ocelli [O]: small pigment cells on the dorsal surface of suckers, and the dorsum and the venter of the body Oviducts [Od]: parts of the female reproductive system communicating ovisacs with the vagina Ovisacs [Os]: specialized part of the coelomic system with oogenic tissue Papillae [Pp]: small tubercles on the dorsal surface of the anterior sucker and the body Posterior crop caeca (last crop diverticula) [Pcc]: elongated processes of the 6% crop chamber number directing posteriorly Preclitellum [Pc]: first 3 somites of the trachelosome between the anterior sucker and the clitellum region Proboscis [P]: an initial muscular part of the digestive system everting through the mouth-pore Pulsatile vesicles [Pv]: segmental subepidermal muscular coelomic dilations on both sides of the body Rectal dilation [Rd]: a dilation of the terminal part of the intestine (rectum [R]) Seminal reservoirs (epididymis) [Sr]: paired parts of the male reproductive system communicating vasa deferentia and initial parts of ejaculatory ducts Sucker (anterior [As], posterior [Ps]): muscular discs formed by some fused somites Tentacles [T]: large projections on the dorsal surface of the anterior sucker BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 54/2007 Testicular lacuna [Tl]: part of the coelomic system surrounding testisacs Testisacs [Ts]: isolated parts of the coelom filled by spermatogenic tissue, indicated the testicular region [Tr] of somite Trachelosome [T]: first 6 somites of the body including preclitellar and clitellar regions of the body Tubercles [Tub]: projections on the dorsal surface of the suckers and on the body Urosome [U]: a part of the body between the trachelosome and the posterior sucker Vagina [V]: a nonpaired structure of the female reproductive system communicating with oviducts and opening by the gonopore on the ventral surface of the clitellum Vasa deferentia [Vd]: a pair channel connecting a vasa efferentia with a seminal reservoir (a part of the male reproductive system) Vasa efferentia [Ve]: a channel connecting each testisac with a vasa deferentia (a part of the male reproductive system) Ventral lacuna [VI]: a longitudinal coelomic channel possessing the nerve cord [Nc] and the ventral vessel [Vv] 1. INTRODUCTION Leeches are fascinated animals, which are the most specialized annelids. Characteristic features of leeches are determined by the transition of their ancestors to temporary external parasitism on large animals. Although they are all popularly considered to be bloodsuckers, a large number of leeches are not par- asitic. Leeches of the Antarctic Seas are restricted to the family Piscicolidae, the order Rhynchobdellida. Ma- rine piscicolid leeches are ectoparasitic on fishes and very few species presumably parasitize on crustaceans and pycnogonids. The Antarctic Seas harbor the di- verse and relatively well known fauna of leeches ex- UTEVSKY, ANTARCTIC PISCICOLID LEECHES plored by the outstanding zoologists: C. Badham, A. Brinkmann, E.M. Burreson, R. Dollfus, V.M. Ep- stein, W.A. Harding, M.C. Meyer, J.P. Moore, L.R. Richardson, R.T. Sawyer, and L. Szidat. However, Antarctic piscicolids need further con- sideration. Their biology, taxonomy and phylogene- tic relationships have not been adequately studied. Until the present time there has not been any detailed monograph on known representatives of the Antarctic leech fauna. We hope that the review of present know- ledge of Antarctic leeches will be of value to marine zoologist and parasitologists and it will be favourable for progress in hirudinology (bdellology). 9 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 54/2007 2, HISTORY OF THES UDYOF ANTARCTIG LEECHES The first information on Antarctic fish leeches is due to W.A. Harding who published in 1922 a short paper based on an examination of leeches collected by the Terra Nova Expedition in 1911 (Harding 1922). All the leeches caught in the coastal waters of Victoria Land were found to belong to a new species. The au- thor assigned this species to a new genus and named it Cryobdella levigata. However, as early as 1916 C. Badham described the leech Austrobdella translucens which was found in fish-farming ponds and off the Southeast coast of Australia (Badham 1916). Sub- sequently, the British-Australian-New Zealand Ex- pedition (BANZARE) found these leeches in the vicinity of Kerguelen Island and in Antarctic coastal waters in 1930 (Moore 1957). In 1938 J.P Moore published information on collections of the Australasian Antarctic Expedition by Aurora (1911-1914). Sir Douglas Mawson, the supervisor of the Expedition, passed on the materials collected in Antarctic coastal waters, in the Durville Sea and Davis Sea to J.P. Moore. Two new species, Pontobdella rugosa and Oxytonostoma varituberculata, were found in the collections (Moore 1938). Moore’s descriptions lack information on the digestive and re- productive systems of these leeches. The author noted that it was necessary to verify their taxonomic status because members of the genus Pontobdella Leach, 1815 occur in the tropics while Oxytonostoma Malm, 1863 is restricted to the North Atlantic and Arctic. In addition, the collections involved several specimens of C. levigata Harding, 1922 from the Davis Sea. Moore commented on the problem of its taxonomic status and assigned this species to the genus Platybdella Malm, 1863. Leeches collected by the Norwegian Antarctic Ex- pedition (Norway, 1927-1928) were not examined by K. Brinkmann until 1948. He described two new species and genera, Trulliobdella capitis and Cryo- bdellina bacilliformis (Brinkmann 1948). The author provided rather adequate descriptions of external and internal characters, but commited a number of errors, resulting in confusion about these species and some others. Leeches collected by the British-Australian-New Zealand Expedition (BANZARE) in the Risser Lar- sen Sea and Commonwealth Sea and given by Sir D. Mawson and J. Harvey Johnson to J.P. Moore were described in 1957 as Pontobdella biannulata and Tra- 10 chelobdellina glabra (Moore 1957). These descriptions lack information on the digestive, reproductive, and coelomic systems. A short description lacking figures of internal characters of 7. glabra was provided by M.C. Meyer and E.M. Burreson (1990). In 1964 R. Dollfus described a new species as- signed to a new genus, Antarctobdella tcherniai (Doll- fus 1964/1965). The leech was collected by Paul Tchernia during the Commandant Charcot Expedi- tion in the vicinity of Heard Island in 1950. In 1965 L. Szidat described a new species, Ophthalmobdella bellisioi from the region of the South Shetland Islands (Szidat 1965). R.T. Sawyer described two new species and one new genus: Glyptonotobdella antarctica collected on isopods in the vicinity of the South Orkneys Islands in 1966 (Sawyer & White 1969) and Antarctobdella crozetensis collected in the vicinity of Marion Island in 1962 (Sawyer 1972). Owing to the Soviet Antarctic Expeditions of USSR (SAE), which have been regular since 1955, V.M. Epshtein described 4 new species and one new genus of leeches: Cryobdella antarctica collected near Mirnyi Station in 1956 (Epshtein 1970b); Pterobdellina australis collected by the 3rd SAE on diesel-electric ship Ob’, in coastal waters of Argentina (Epshtein 1970a) and by the Evrika 22nd expedition in 1987 near the South Orkney Islands; Trulliobdella alba collected by the 8th SAE on diesel- electric ship Ob’ in 1963 (Epshtein 1970b); Moorebdellina uschakovi collected by the Skiff IIT Ex- pedition in 1970 (Epshtein 1974). Since 1970 researchers have concentrated their attention not only on descriptions of new species, but also on general systematic problems. Epshtein established the subfamily Platybdellinae, considered patterns of the distribution of the subfamilies and revised the taxonomic status of a number of species: Pontobdella rugosa Moore, 1938 (Epshtein 1972) and Pontobdella biannulata Moore, 1957 (Epshtein & Okun’ 1991) were assigned to the genus Moorebdel- lina Epstein, 1972. Sawyer (1986) revised the world fauna of Hiru- dinea, including Antarctic fish leeches, in his three- volume monograph. He shares Epshtein’s system of the subfamilies but provisionally included Cryobdella antarctica in the genus Glyptonotobdella, proposed to consider species of the genera Trulliobdella, Cryo- bdellina, Antarctobdella, Ophthalmobdella as synonyms of Trulliobdella capitis and Moorebdellina uschakovi as a synonym of Moorebdellina rugosa. Meyer and Bur- reson’s revision of 1990 is based on an examination of the material collected by the Australian National Antarctic Research Expedition to Heard Island (1949-1950), Melbourne (ANARE); Virginia Insti- tute of Marine Science (1959-1967), Gloucester Point, Virginia (VIMS); USNS Eltanin cruises (1962-1968), R/V Hero cruises (1968-1982) and /s- las Orcadas supported by the National Science Fo- undation of U.S. and Argentine Navy; German An- tarctic Expedition (1977-1978), Hamburg (GAE) (Meyer & Burreson 1990). The authors proposed new synonyms for Antarctic leeches; described two new species Trachelobdella bathyraja from the S. Shetland Islands, Megaliobdella szidati from the Ross Sea and a new genus Megaliobdella; reported on new dis- tribution records for species previosly unknown in Antarctic Seas (Galatheabdella bruuni Richardson & Meyer, 1973; Pontobdella tasmanica Hickman, 1947) UTEVSKY, ANTARCTIC PISCICOLID LEECHI and provided new information on the distribution of other Antarctic species. A detailed study of external, internal characters, and of coelomic system in particular, in the majority of fish leeches recorded from the Antarctic Seas, as well as a description of 4 new species (Notorhenio- bdella sawyeri, A. Utevsky, 1993; Cryobdella ladovi Epstein & A. Utevsky, 1994; Cryobdella pallida A. Utevsky, 1996; Moorebdellina meyeri, 1997) led the author to undertake a new revision and write this book. Recent phylogenetic analyses have revealed the pa- raphyletic and/or polyphyletic nature of the traditio- nal piscicolid subfamilies. According to principles of the phylogenetic systematics the subfamilies Pontob- dellinae, Platybdellinae, and Piscicolinae are unnatural groupings with no formal status (Williams & Bur- reson 2006, Utevsky et al. 2007). However we still employ the traditional system (Utevsky 2005) because of its practical usefulness. The system seems quite morphologically grunted and convenient to purposes of identifying fish leeches by non-experts in the field. BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 54/2007 3. MORPHOLOGY AND ANATOMY OF PISCICOLID LEECHES The piscicolid leeches (Hirudinea: Piscicolidae) differ from other leeches by their suckers which are usually clearly marked off from the body. The anterior sucker consists of the fused prostomium and 5 somites. The body is divided into two distinct regions (Fig. 1A). The short anterior portion is called the trachelosome. The longer and wider posterior portion is called the urosome. The trachelosome involves the preclitellum and clitellum, each consisting of 3 somites (6-11). The urosome is subdivided into the crop region (7 somites, 12-18) and intestinal region (8 somites, 19-26). The posterior sucker is made up of 7 fused somites (27-33). Other features, which define leeches, concern de- tailed characters of the body and internal organs. The body and suckers may be described by a model of body shape. This model simulates the body as the horizontal and vertical projections of the system of 6 trapezoids: trachelosome as 2 trapezoids, urosome as 4 trapezoids; and suckers as ellipses (Fig. 1B). The first trapezoid usually lies between the site of attach- ment of the anterior sucker and the male gonopore. The second trapezoid usually extends from the male gonopore to the urosome. The first trapezoid of the urosome usually extends from the bound of the trachelosome and urosome to the level of the first pair of testisacs (to the second pair if the first pair are absent), the second trapezoid includes the rest of the crop region, the third trapezoid includes the urosome from the boundary of the crop and intes- tinal region to the site of termination of the poste- rior crop caeca or beginning of the rectal dilation, and fourth trapezoid extends to the site of attachment of the posterior sucker. Proportions of the body According to the body model of leeches (Fig. 1 B) may be: cylindrical (D/N = 1) or subcylindrical (1 < (D/N)<1.5) (round or ellipse transverse sections); narrow, long (7< (L/D2 < 17) or short (L/D>4 = 7); subcylindrical (1 <(D/N) < 1.5) or flattened (1.5 < (D/N) < 5); short and wide (3 < (L/D;) < 7). By length, leeches are can be classified into: small — up to 20 mm; not large — 21 — 30 mm; medium — 31- 60 mm: large — more than 60 mm (very large — more than 100 mm). 12 Suckers 1. Size of the anterior and posterior suckers: large C/Ds>>1; medium C’/D>=1; small C/D><1. 2. Mode of attachment to the trachelosome or uro- some: centrally (R/M=1); excentrically (R/M>1). The posterior sukers may face ventrally (Fig. 4A, B) or directly posteriorly (Fig. 4,C, D). 3. Characters of dorsal surface. The dorsal surface of anterior and posterior suckers may be smooth (Figs. 2 A, 4 D), with transverse furrows or may bear small tubercles, papillae or tentacles. The edge of the anterior sucker may be rough (Fig. 2 B), its dorsal surface may possess eyes, eye-like spots, ocelli (Fig. 3) or a number of papillae. The posterior sucker may possess a number of ocelli near the edge (Fig. 4 C). 4. Location of the mouth-pore. Mouth-pore centrally located (Fig. 5 A), or lies prior to the centre (Fig. 5 B) or at the anterior margin of the sucker (Fig. 5 C). Trachelosome The trachelosome like the urosome may be smooth or bear small and/or large tubercles and papillae,and marginal flanges. Some first annuli of the trachelosome may pos- sess eyes or ocelli. A more important feature of the trachelosome is the structure of the clitellum. The clitellum may be marked by deeper furrows or larger annuli. Annuli of the clitellum may differ from pre- clitellar annuli in their surface. It bears male and female gonopores, which may be surrounded by the external copulatory area. In different leeches, gonopores are separated by a species-specific number of annuli. The structure of the copulatory area is also an important character. The copulatory area usually surrounds the female gonopore or is situated near it and serves for attachment of a spermatophore, but the copulatory area may surround both the gonopores or the male gonopore (Fig. 6). Urosome In addition to tubercles, papillae and marginal flanges, the urosome may bear external lateral pulsatile vesicles and segmental ocelli, which are located on the dorsum and the venter (Fig. 7). The structure of the com- plete somite is of taxonomical significance. These somites are situated in the crop region of the urosome UTEVSKY, ANTARCTIC PISCICOLID LEECHI (Fig. 8). The ancestral somite is 3-annulate, includes Annulus A, may bear pulsatile vesicles and seg- annuli Aj, Az, and A3. The number of these annuli mental ocelli. The number of annuli per somite may may reduce (2-annulate somite) or increase by addi- _—_ decrease in the intestinal region. The location of the tional annulation. anus on the dorsum is taken into consideration. S ®) = | a ae m =e FIG. 1. Body model: A, subdivisions of the body; As, anterior sucker; T, trachelosome; U, urosome; Ps, posterior sucker; Pc, preclitellum; Cl, clitellum; Cr, crop region; Ir, intestinal region; B, dimensions of the body; L, common lenght; Li, lenght of trachelosome; L», length of urosome; $}-Ss, hights of trapezoids; d;-d-, width of bases of trapezoids; Cı, C2, lengths of suckers; C’), C2, widths of suckers; Ri, Ro, lengths of external parts of suckers; Mı, Mb, lengths of internal parts of suckers; Nj, No, maximal thikness of urosome and trachelo- some; D}, D>, maximal width of urosome and trachelosome. 13 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 54/2007 FIG. 2. Characters of dorsum of anterior sucker: Tub, tubercles; Pp, papillae; T, tentacles. FIG. 3. Location of eyes, eye-like spots, ocelli on anterior sucker: E, eyes; Els, eye-like spots; O, ocelli. FIG. 4. Surface and attachment of posterior suckers: Pp, papillae; O, ocelli. 14 onen ten nn pec py ee eer SE Rene LE FIG. 5. Location of the mouth-pore: Mp, mouth-pore; P, proboscis. Digestive system In piscicolid leeches, the proboscis is an eversible muscular tube lying within a proboscis cavity. It may extend from ganglion 1 to ganglion 5 of the ventral nerve cord. The proboscis is connected with the short esophagus, which often has diverticula located be- tween ganglia 3 and 5. The esophagus opens into the long crop. The crop usually possesses 6 or 5 cham- bers, but the crop may be tubular without distinct chambers. The crop chambers may be ovoid or dia- mond-shaped, with lateral processes of various form or without them. The posterior crop caeca are cau- FIG. 6. Ventrum of clitellum: Ca, copulatory area. UTEVSKY, ANTARCTIC PISCICOLID LEECHI dal processes of the posterior cham- ber of the crop. In the family Pis- cicolidae, the posterior crop caeca show different degrees of junction; they may be completely separated, nontotaly fused with various num- ber and size of fenestrae, or com- pletely fused together without fe- nestrae. The posterior crop caeca may be absent. The intestine has various numbers of lateral cham- bers (5 at most). They show diffe- rent degrees of developing and may have lateral processes of different structure. The posterior chamber may be modified into the “folded organ”. [he intestine is terminated by the rectum, which may be tu- bular or with a side dilation of vary- ing size (Fig. 9). Reproductive system Male reproductive system The piscicolid leeches have 4, 5, or 6 pairs of testi- sacs located between chambers of the crop. The exi- stence of 5 pair is caused by missing the Ist pair of testisac; in leeches with only 4 pairs of testisac the first two pairs are missing. A short vasa efferentia connects each testisac to a vas deferens, which extends anteri- orly on each side of the body. Each vasa deferentia opens into a seminal reservoir, which has various sha- pe and size. It may be straight, loop-like or very coiled and may be located between ganglia 6 and 7 or ex- tend to ganglion 12 of the ventral nerve cord. Semi- nal reservoirs empty into ejaculatory ducts. An eja- culatory duct is divided into initial, terminal and com- mon parts, which vary in form and size. Terminal parts and the single common part of ejaculatory ducts (atrium) are highly muscular and often covered by accessory glands, which open through walls into the lumen. The copulatory bursa varies in length and opens to the exterior through the male gonopore. Female reproductive system Female reproductive organs include a pair of ovisacs and the conductive tissue. The ovisacs may be straight or coiled, short or long. Some piscicolid leeches have conductive tissue, which may be absent in many genera. Ovisacs empty into oviducts, which lead to 15 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 54/2007 FIG. 7. External features of somites: Tub, tubercles; Pp, papillae; Py, pulsatile vesicles; O, ocelli; Mf margi- nal flanges. the nonpaired vagina. The vagina opens to the exterior through the female gonopore, which may be sur- rounded by the copulatory area. The conductive tissue connects the ovisacs with the copulatory area (which may be external or internal and located on the male bursa) if it is present. It may possess some parts such as reservoirs, cords and mass with an internal lumen. The conductive tissue acts as a pathway for sperma- tozoids from a spermatophore to the ovisacs (Fig. 10). Coelomic system In most piscicolid leeches, there is commonly a well developed coelomic (lacunar) system which is a modi- 24 _c B A 1 2 ~~ 3 FIG. 8. Model of annulation of complete somite: Aj-3, annulation of first degree, Bı.s, annulation of second degree; C;-12, annulation of third degree; Dj-24, annulation of fourth degree; Ts, testisac; G, ganglion; Ne, nerve cord. 16 | 1g E Epd es 8 9 C 10 Cotip AA. 12 13 7 14 Fee 15 a Ic+tLp 16 = di F En, 1% 2 Va PcctLp ete. R+Rb : 4 FIG. 9. Model of digestive system of piscicolid leeches: P, proboscis; Ep, esophagus; Epd, esophageal diverticula: C, crop; Cc+Lp, crop chamber with lateral processes; I, intestine; Ic+Lp, intestinal chamber with lateral processes: R+Rb, rectum with rectal dilation; A, anus; Pcc, posterior crop caeca; F, fenestrae; Pcc+Lp, lateral pro of posterior crop caeca; Nc, nerve cord; g, ganglion. 2c Pe BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 54/2007 On “J 10 11 12 3 Ne FIG. 10. Model of reproductive system of piscicolid leeches: Ts, testisacs; Ve, vasa efferentia; Vd, vasa deferentia; Sr, seminal reservoir; Ed, ejaculatory ducts; Tp, terminal parts of ejaculatory ducts (atrial cornu); Cp, common part of ejaculatory ducts (common atrium); Ag, accessory glands; Cb, copulatory bursa; Os, ovisacs; Od, oviducts; Ctm, conductive tissue mass; Ctc, conductive tissue cord; Nc, nerve cord; g, ganglion. 18 UTEVSKY, ANTARCTIC PISCICOLID LEE( FIG. 11. Model of coelomic system of piscicolid leeches: DI, dorsal lacuna; V1, ventral lacuna; LI, lateral lacuna; MI, marginal lacuna; Tl, testicular lacunae; Mc, main connection; Ac, accessory connection; Bw, body wall. FIG. 12. Coelomic system in ganglion region and main connections. A, absence of main connection; B, un- connected main connection with processes of the ventral lacuna; C, unconnected main connection with processes of the dorsal lacuna; D, connected main connection; DI, dorsal lacuna; V1, ventral lacuna; LI, lateral lacuna; Mc, main connection. 19 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 54/2007 fication of the coelom, and forms a regular system of channels (lacunae). It may consist of dorsal, ventral, lateral and marginal lacunae, pulsatile vesicles and lacunae of testisacs (Fig. 11). The dorsal lacuna may contain the dorsal blood vessel and the ventral lacu- nae contain the ventral blood vessel and the nerve cord. The marginal lacunae are located between the epithelium and muscle layer, and are usually muscu- larized. The arrangement of coelomic channels in a somite of the crop region which contains all main parts of this system is of great taxonomical signifi- cance. The structure of the coelomic system is very variable. The dorsal lacuna may be absent or present. This lacuna may be divided into two channels. The dorsal blood vessel may be located outside of the lacuna. The lateral, marginal and testicular lacunae may be absent or present. The lateral lacunae may be connected with various elements of the coelomic system: testicular lacunae, main and accessory connec- tions, marginal lacunae and pulsatile vesicles. Main connections (formed by ventral and dorsal lacunae) are located in the ganglion region, and vary in their development. They may be absent, with a small dilation of the ventral lacuna in the ganglion region (Fig. 12 A); unconnected, with longitudinal processes of the ventral lacuna which extend to the dorsal lacuna in the same region (Fig. 12 B); un- connected, with processes of the dorsal lacuna which extend to the ventral lacuna in the ganglion region (Fig. 12 C); connected by a junction of the ventral and dorsal lacunae in the ganglion region (Fig. 12 D). The accessory connection (formed by ventral, dor- sal and testicular lacunae) located in the testicular re- gion of the somite between two adjacent ganglia, may be developed to different degrees: it may be absent (Fig. 13 A); unconnected, with junction of testicular and ventral lacunae (Fig. 13 B); unconnected, with junction of testicular and dorsal lacunae (Fig. 13 C); connected by junction of dorsal, testicular and ventral lacunae (Fig. 13 D). FIG. 13. Coelomic system in testicular region and accessory connections: A, absence of accessory connec- tion; B, unconnected accessory connection with processes of the ventral lacuna; C, unconnected accessory connection with processes of the dorsal lacuna; D, connected accessory connection; DI, dorsal lacuna; VI, ventral lacuna; Ml, marginal lacuna; TI, testicular lacunae; Ac, accessory connection. 20 UTEVSKY, ANTARCTIC PISCICOLID LEECHI FIG. 14. Pulsatile vesicles: A, pulsatile vesicles of the subfam. Piscicolinae; B, pulsatile vesicles of the gen. Pontobdella (subfam. Pontobdellinae); C, pulsatile vesicles of the gen. Moorebdellina (subfam. Pontobdellinae); DI, dorsal lacuna; V1, ventral lacuna; LI, lateral lacuna; SII, sublateral lacuna; MI, marginal lacuna; Tl, testicular lacunae; Mc, main connection; Ac, accessory connection; Pv, pulsatile vesicles. Pulsatile vesicles may be absent (subfam. Platy- bdellinae) or present. It may be of different origin: pulsatile vesicles located in places of connections of lateral lacunae with the main connection (one pair of external vesicles per somite in leeches of the subfam. Piscicolinae) (Fig. 14 A); two pair of subepidermal vesicles per somite located in places of connections of lateral lacunae with the main and accessory connec- tions in the leeches of the subfam. Pontobdellinae (gen. Pontobdella) (Fig. 14 B); pulsatile vesicles as dilation of marginal lacunae (one or two pair of sub- epidermal vesicles per somite in leeches of the sub- fam. Pontobdellinae (gen. Moorebdellina) (Fig. 14 C). External pulsatile vesicles differ in size. BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 54/2007 4, BIOLOGY OF PISCICOLID EEECHES Piscicolid leeches (Hirudinea: Piscicolidae) inhabit fresh, brackish and salt waters all over the world. Marine leeches are benthic animals, which occur from the littoral to more than the 4000 m depth. Antarc- tic piscicolids may be classified into both stenobathic and eurybathic species, which are respectively steno- thermic and eurythermic (Table 1). Leeches are para- sitic on benthic and benthophagous fishes because they deposit their cocoons on bottom substrates such as stones, rhizoids, hard shelled invertebrates; young leeches infest demersal hosts (Meyer & Burreson 1990, Utevsky 2000). Life histories of the majority of Antarctic leeches are poorly known and their host-parasite relationships need further investigation. Unlike representatives of other families of Hirudinea, piscicolid leeches are closely associated with their hosts. A number of pis- cicolid leeches from the North Atlantic, the North Pacific and the Arctic lay their cocoons on decapod crustaceans and pycnogonids but feed on the blood of various fishes (Meyer & Barden 1955, Bakay et al. 1998). Similar cocoon-laying was found in the Ant- arctic leech Glyptonotobdella antarctica (Sawyer and White 1969). The leeches and their cocoons occur on the isopod Glyptonotus antarcticus (Janssen 1993). In addition, the sea urchin Sterechinus sp. and the benthic octopus Pareledone (charcoti?) serve as hosts. According to Janssen (1993), G. antarctica moves between Ant- arctic octopuses and their potential prey. There is no evidence of leeches feeding on these invertebrates. Piscicolid leeches may deposit their cocoons on eggs of their host fishes. Young leeches infest fish larvae (Khan & Paul 1995). There are no similar records for Antarctic leeches. However, it was found that the eggs of unidentified fishes and cocoons of unidentified leeches (Trulliobdella bacilliformis (Brinkmann 1948)?) stuck together in one of the lots of our collection. Many marine leeches deposit their cocoons on rocks (Khan & Paul 1995). Some Antarctic leeches frequently occur on rocks, rhizoids, etc., but there are TABLE 1. Depth of founding Antarctic piscicolid leeches. Species Depth (m) ie Cryobdella levigata Harding, 1922 10-40 2; C. antarctica Epstein, 197 11 3: C. hadovi Epstein & A.Utevsky, 1994 360-500 4. C. pallida A.Utevsky, 1997 - 5; Austrobdella translucens Badham, 1916 195 6. Glyptonotobdella antarctica Sawyer & White, 1969 0-665 7. Epsteinia alba (Epstein, 1970) = 8. Pleurobdella varituberculata (Moore, 1938) 128-546 9. P australis (Epstein, 1970) 100-400 FO, Pontobdella tasmanica Hickman, 1947 27-891 ine Moorebdellina rugosa (Moore, 1938) 15-300 12. M. biannulata (Moore, 1957) 137-580 15, M. uschakovi Epstein, 1974 58-410 14. M. meyeri A.Utevsky, 1997 140-300 15. Megaliobdella szidati Meyer & Burreson, 1990 549-556 16. Trulliobdella capitis Brinkmann, 1948 170-270 17. 7! bacilliformis (Brinkmann, 1948) 5-500 18. Trachelobdella bathyrajae Meyer & Burreson, 1990 - 19. Trachelobdellina glabra Moore, 1957 20-437 20. Nototheniobdella sawyeri A.Utevsky, 1993 50-200 21. Galatheabdella bruuni Richardson & Meyer, 1973 4052 i) i) UTEVSKY, ANTARCTIC PISCICOLID | TABLE 2. Hosts of the Antarctic piscicolid leeches. Leeches — — | S INT « x IS ax En SQ S m = > P N SII|S EN S | & Ri S S N Hosts .S Sy R ES Sn DE le (8 Sire] NS ae i SPSOESL SIRI STs IS Is |. ISIS] 87 «= Ss Ss IS: N ERS SS S Peery ~ m = 2 iS Na u Ra ES z x SISISISIS N SIRS|I|SIN| So S IS IS SS | SISIS SI 2 SISH.SISIOI SI SISISISISIIPISISISIRISISI SPR] S = SID. REIS ISIS | SI SIS|I|SISIC SIT JA FI TI FI s N = I S NS Sol ~ rR N ™ % a oe Bal N NIS = > Sl ee RB] SR ll eb ea ee S PS II sO 9 A : ; A . ie sie sie sic eS Ir - Ir. Ic Io - IS enn EEEEEE HNEBEBEEEEN Phyl. Arthropoda Cl) Crustacea Ord. Isopoda COOL Phyl. Chordata Cl. Elasmobranchii Ord. Rajiformes Fam. Rajidae [Fam,Rajidae ee II TTS T Fam. Rhinobatidae Ed) SS lll Cl. Actinopterygii Ord. Gadiformes Fam. zen i I TIIIIITI III GM Mr EEENENENEEEEE EEREEEE Fam. Macrouridae 7. Conphaenoidensp — || eee Ord. Ophidiiformes Fam. Ophidiidae 8. Bassozerus sp ZEEERAABEREREREAER Ord. Perciformes Fam. Nototheniidae i ee oe Ci III fit wnat PP PP Pe Stee OGRE i oe EBENE B = HH bo Oo BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 54/2007 TABLE 2. Continued. M. uschakovi M. meyeri Me P tasmanica M. biannulata M.rugosa 15. Trenatomus borchgrevinki HENEEEEEE EEEETSEM 117 dT LTT TT TT TTT | me TT TT TT ene eee 0 00 ee Fam. Bathydraconidae 5.Pocheenchhngogona | TTT 111111111111 Mil Fam. Chaenichthyidae 20. Chaenichthys rhinoceratus Leeches e. szidati if bacilliformis G. antarctica G. bruuni P australis C. levigata C. antarctica C. hadovi C. pallida Tr. glabra IR bathyrajae N. sawyeri P varituberculata I! capitis [a Nepagepicime | | I OO UT 22. Pseudochaenichthys georgianus II) | | BEE 23. Chaenocephatus aceras ||| 24. Cbowvetenis | TT | Be ee ae 26. Champsocephalus gunmari | | | | | | | tt | I | 27. Chionodraco katnleenae 28. C. hamatus 29. C. rastrospinosus Fam. Harpagiferidae HEEEEEEEEEEE EEE BEE [30.Chaenodracowilsoni_ | [III Po 000 2 no records on cocoon-laying on the bottom substrates and hosts of the leeches are unknown (Table 2). Young leeches hatch within from 30 to 300 days after cocoon deposition. There are annual and more long-term life cycles (Khan & Paul 1995). As for Antarctic piscicolids, it seems likely that small leeches are characterized by annual life cycles while large leeches live for a longer time. The list of host fishes and carrier invertebrates is given in Table 2. C. pallida, E. alba, G. anatarctica, and 7! bathyrajae may be classified as species-specific parasites. Broader host ranges are recorded for C. ha- dovi (2 host species of one genus), C. levigata (3 host 24 species of 2 related genera of one family), A. translu- cens (2 host species of related families of one super- family), C. antarctica (11 host species of 10 genera of 2 families of one superfamily). The lowest host spe- cificity is peculiar to N. sawyeri (7 host species of 6 genera of 3 related families) and 7. bacilliformis (10 host species of 8 genera of 3 related families). Hosts of the other leeches are unknown. Piscicolid leeches may be grouped according to the duration of stay on a host fish as temporary or per- manent parasites. The former usually leave their host when sated and reattach to different host when hun- ery. The latter do not detach from their hosts (Sawyer UTEVSKY, ANTARCTIC PISCICOLID LEECHI TABLE 3. Intensity of infestation of the hosts. Species Infestation Il Cryobdella levigata Harding, 1922 1-4 C. antarctica Epstein, 1970 1-36 3 C. hadovi Epstein & A.Utevsky, 1994 1-21 4 C. pallida A.Utevsky, 1996 1-6 ay Austrobdella translucens Badham, 1916 1-2 6. Glyptonotobdella antarctica Sawyer & White, 1969 1-132 7 Epsteinia alba (Epstein, 1970) ] 8. Pleurobdella varituberculata (Moore, 1938) | OR P australis (Epstein, 1970) ] 10. Pontobdella tasmanica Hickman, 1947 = Tole Moorebdellina rugosa (Moore, 1938) | 12: M. biannulata (Moore, 1957) = 13: M. uschakovi Epstein, 1974 = 14. M. meyeri.A.Utevsky, 1996 - 15: Megaliobdella szidati Meyer & Burreson, 1990 = 16. Trulliobdella capitis Brinkmann, 1948 1-36 17% T. bacilliformis (Brinkmann, 1948) 1-19 18. Trachelobdella bathyrajae Meyer & Burreson, 1990 3 1) Trachelobdellina glabra Moore, 1957 = 20. Nototheniobdella sawyeri A.Utevsky, 1993 1-29 2. Galatheabdella bruuni Richardson & Meyer, 1973 — 1986). Leeches locate on sites like fins, folds between of impossibility of storing blood. E. alba also lacks pectoral fins and body, head, mouth and gill cavities posterior crop caeca. However, it is unknown whether where the proboscis might easily penetrate through the leech is a permanent parasite because only one spe- the fish skin. cimen has been recorded. Leeches of the genus Cryobdella are examples of Leeches of the genera Pleurobdella, Moorebdelli- permanent parasites. They have a relatively large na, Megaliobdella, Galatheabdella, and Trachelobdel- posterior sucker for long-term attachment. An ex- — /ina found free living are apparently temporary para- treme case of such is the adaptation is C. Gadovi. The sites. Findings of some leeches of the above genera on unique shape of its posterior sucker is adapted to arthropods support this supposition. attachment to gill rakers. The lack of posterior crop Little is known about intensity and prevalence of caeca is another evidence of permanent parasitism. infestation by Antarctic piscicolids. Intensity varies The leech has to feed on its host all the time because within a broad range (Table 3). to J\ BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 54/2007 5. COLLECTION AND PRESERVATION Leeches may be collected from the fins, the head, the mouth and gill cavities of fishes. It should be em- phasized that these parasites often leave their hosts immediately after capture. Therefore it is necessary to examine a lot of fishes at once. Leeches may be found on crustaceans, among rhizoids of algae, and free- living on the bottom. They may be collected by dredge, Petersen grab, Otter trawl, Blake trawl, Men- zies trawl, and divers. Collected specimens should be narcotized in 10% ethanol, 7% MgCh, CdSO; or in chloral hydrate for relaxation. But eyes, eyes-like spots, segmental ocelli, ocelli on the posterior sucker and the coloration must be examined before narcotization and preservation 26 because leeches may be depigmented by these sub- stances. After relaxation the leeches should be fixed in 5% formalin, or Bouin’s fluid for histological preparations. The leeches may be stored in these fluids, large spe- cimens should be injected with fixative fluid for valid preservation of internal organs. Preserving and fixing solution based on formalin preserves pigmented struc- tures and coloration longer than that based on ethanol (70%). Moreover, ethanol macerate the body wall, in- ternal organs and worsens histological staining. Pre- served specimens must be stored in opaque vials. Leeches may be fixed in 96% ethanol and stored at low temperature conditions for DNA analysis. 0SSYSTEMATIC PART Classification of the Family Piscicolidae Johnston, 1865. “Subfamily Platybdellinae” Epstein, 1970 Genus Platybdella Malm, 1863 No Antarctic representatives * Genus Cryobdella Harding, 1922 C. levigata Harding, 1922 C. antarctica Epstein, 1970 C. hadovi Epstein & A.Utevsky, 1994 C. pallida A.Utevsky, 1997 Genus Coitobdella S.Utevsky, 1997 No Antarctic representatives Genus Heptacyclus Vasilyev, 1939 No Antarctic representatives Genus Malmiana Strand, 1942 No Antarctic representatives Genus Oceanobdella Caballero, 1956 No Antarctic representatives Genus Sanguinotus de Silva & Burdon-Jones, 1961 No Antarctic representatives Genus Arctobdella de Silva & Kabata, 1961 No Antarctic representatives Genus Phyllobdellina Epstein & S.Utevsky, 1993 No Antarctic representatives * Genus Austrobdella Badham, 1916 A. translucens Badham, 1916 Genus Pterobdella Kaburaki, 1921 No Antarctic representatives Genus Phyllobdella Moore, 1939 No Antarctic representatives Genus Pterobdellina Bennike & Bruun, 1939 No Antarctic representatives Genus Notostomum Levinsen, 1882 No Antarctic representatives Genus Ostreobdella Oka, 1927 No Antarctic representatives * Genus Glyptonotobdella Sawyer & White, 1969 G. antarctica Sawyer & White, 1969 Genus Myzobdella Leidy, 1851 No Antarctic representatives Genus Piscicolaria Whitman, 1889 No Antarctic representatives Genus /llinobdella Meyer, 1940 No Antarctic representatives Genus Crangonobdella Selensky, 1914 No Antarctic representatives * Genera represented in the Antarctic region UTEVSKY, ANTARCTIC PISCICOLID LEECHES Genus Beringbdella Caballero, 1976 No Antarctic representatives * Genus Epsteinia A.Utevsky, 1993 E. alba (Epstein, 1970) Genus Mysidobdella Selensky, 1927 No Antarctic representatives Genus Marsipobdella Moore, 1952 No Antarctic representatives Genus Hemibdella van Beneden & Hesse, 1863 No Antarctic representatives Genus Aestabdella Burreson, 1976 No Antarctic representatives Genus Ringueletobdella Epstein, 1994 No Antarctic representatives Genus Makarabdella Richardson, 1959 No Antarctic representatives Genus Bathybdella Burreson, 1981 No Antarctic representatives * Genus Pleurobdella A.Utevsky, 1995 P varituberculata (Moore, 1938) P australis (Epstein, 1970) “Subfamily Pontobdellinae” Llewellyn, 1966 * Genus Pontobdella Leach, 1815 P tasmanica Hickman, 1947 Genus Oxytonostoma Malm, 1863 No Antarctic representatives * Genus Moorebdellina Epstein, 1972 M. rugosa (Moore, 1938) M. biannulata (Moore, 1957) M. uschakovi Epstein, 1974 M. meyeri A.Utevsky, 1997 * Genus Megaliobdella Meyer & Burreson, 1990 M. szidati Meyer & Burreson, 1990 “Subfamily Piscicolinae” Johnston, 1865 Genus Piscicola Blainville, 1818 No Antarctic representatives Genus Codonobdella Grube, 1873 No Antarctic representatives Genus Baicalobdella Dogiel, 1957 No Antarctic representatives Genus Zeylanicobdella de Silva, 1963 No Antarctic representatives Genus Caspiobdella Epstein, 1966 No Antarctic representatives Genus /talobdella Bielecki, 1993 No Antarctic representatives BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 54/2007 Genus Acipenserobdella Epstein, 1969 No Antarctic representatives Genus Limnotrachelobdella Epstein, 1968 No Antarctic representatives Genus Zaimenobdella Epstein, 1964 No Antarctic representatives * Genus Trulliobdella Brinkmann, 1948 7. capitis Brinkmann, 1948 T! bacilliformis (Brinkmann, 1948) * Genus Trachelobdella Diesing, 1850 T. bathyrajae Meyer & Burreson, 1990 * Genus Trachelobdellina Moore, 1957 T! glabra Moore, 1957 * Genus Nototheniobdella A.Utevsky, 1993 N. sawyeri A.Utevsky, 1993 Genus Bdellamaris Richardson, 1953 No Antarctic representatives Genus Branchellion Savigni, 1822 No Antarctic representatives Genus Orientobdella Epstein, 1962 No Antarctic representatives Genus Johanssonia Selensky, 1914 No Antarctic representatives Genus Cystobranchus Diesing, 1859 No Antarctic representatives Genus Calliobdella van Beneden & Hesse, 1863 No Antarctic representatives * Genus Galatheabdella Richardson & Meyer, 1973 G. bruuni Richardson & Meyer, 1973 Genus Brumptiana Knight-Jones & Llewellyn, 1984 No Antarctic representatives Key to the Subfamilies of the Family Piscicolidae The keys are based on characters peculiar to Ant- arctic leeches and are not intended for identifying any taxa beyond the Antarctic region. la Externally visible pulsatile vesicles absent ... 2 lb One pair of externally visible pulsatile vesicles per somite ...“Subfamily Piscicolinae” (p. 60) 2a Body with large tubercles; annuli distinctly div- ided; pulsatile vesicles subepidermal “Subfamily Pontobdellinae” (p. 28) 2b Body with small tubercles and papillae or smooth; annuli indistinctly divided; pulsatile vesicles absent * Genera represented in the Antarctic region 28 “Subfamily Pontobdellinae” Diagnosis. Leeches of varying sizes; body long, cylin- drical or subcylindrical. Externally visible pulsatile vesicles, segmental ocelli and ocelli on posterior sucker absent. Annuli of somites distinctly divided, with tubercles and papillae. Anterior sucker may bear tentacles, tubercles, papillae, and eye-like spots. Digestive system of varying structure, posterior crop caeca totally or nontotally fused (with fenestrae). Reproductive system of varying structure, copu- latory area absent. Coelomic system possesses dorsal, ventral, lateral, testicular lacunae; marginal lacunae absent or present. Usually with two subepidermal pulsatile vesicles on each side of an urosomal somite. References: Sawyer 1986, Epshtein er al. (1994). Key to the Antarctic Genera of the “Subfamily Pontobdellinae” la Number of annuli in complete somite less than 12, tubercles of varying size; not very large leeches 1b Complete somite 12-annulate with equal large conic tubercles; very large leeches Genus Megaliobdella (p. 39) 2a Large or medium-sized leeches, three peak eye- like spots on anterior sucker, posterior crop caeca totally fused, accessory glands absent Genus Pontobdella (p. 28) 2b Not large leeches, eye-like spots without peaks, posterior crop caeca nontotally fused with fe- nestrae, accessory glands present Genus Moorebdellina (p. 30) Genus Pontobdella Leach, 1815 (Syn.: Stibarobdella Leigh-Sharpe, 1925; Pentabdella Lewellyn, 1966) Type species: Pontobdella muricata Linnaeus, 1758. Medium-sized or large leeches. Body long, cylindrical or subcylindrical, indistinctly divided into trachelo- some and urosome. Complete somite sharply divided into small number of annuli with tubercles. External copulatory area on clitellum absent. Anterior sucker: small, sharply divided from tra- chelosome, with tentacles, tubercles and eye-like spots. Posterior sucker: small, indistinctly divided from urosome, facing directly posteriorly. Digestive system. Esophageal diverticula absent. Crop chambers lacking lateral processes. Posterior FIG. 15. Disribution of the Pontobdella tasmanica. crop caeca totally fused. Intestine possesses weakly developed chambers without lateral processes. Reproductive system. Seminal reservoirs long, bursa short, accessory glands absent. Ovisacs small, conductive tissue well developed. Coelomic system. Dorsal, ventral, lateral, and tes- ticular lacunae present. Two pairs of small subepi- dermal pulsatile vesicles per somite present. Host data: skates and sharks. Distribution: absent in polar waters of the North- ern Hemisphere, most abundant in tropical waters. Congenitors: P macrothela Schmarda, 1861 - equatorial world-wide P tarpobanensis de Silva, 1963 - Sri Lanca P planodiscus Baird, 1869 - Patagonia P vosmaeri Apathy, 1888 - north-eastern Atlantic, Me- diterranean P australiensis Goddard, 1909 - Australia P moorei Oka, 1910 - Japan UTEVSKY, ANTARCTIC PISCICOLID LEECHES P bimaculata Oka, 1910 - Japan P loricata Harding, 1924 - India P dispar Cordero, 1937 - Brazil P tasmanica Hickman, 1947 - Tasmania, Antarctica References: Lewellyn 1966, Sawyer 1986; Ep- shtein et al. (1994). Pontobdella tasmanica Hickman, 1947* (Syn. Stibarobdella tasmanica (Hickman, 1947) ) 1. External characters. Large leeches. Up to 80 mm in length and 10 mm in width. Body indistinctly divided into trachelosome and urosome, with tu- bercles. Anterior sucker small, distinctly divided from trachelosome, eccentrically attached, directed ventrad, cup-shaped, with 4 pairs of submarginal tentacles. One pair of three peak eye-like spots on anterior * Species described on literature data BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 54/2007 sucker. Mouth-pore centrally located. Posterior sucker small, but larger than anterior one, cup-shaped, cent- rally attached, facing directly posteriorly. Annulation. Clitellum 6-annulate, including 443, biannulate 5 and 6, 7A, which slightly constricted in diameter and lacking large tubercles, annulus 7A; very small. Male gonopore between annuli 5A; and 5A>, female gonopore between annuli 6A; and 6A». Copulatory area absent. Complete somite 3-annulate; annulus A) slightly wider than A; and A3. Annulus A> with 4 dorsal and 4 smaller ventral papillae; A; and A3 with 6 dorsal and 6 smaller ventral papillae. Anus on papilla, separated by 2 annuli from poste- rior sucker. Coloration. Body and suckers unpigmented, only clitellum with brownish tinge. 2. Digestive system. Structure unknown. 3. Reproductive system. Structure unknown. 4. Coelomic system. Structure unknown. 5. Host data: Trigonorhina fasciata (Rajiformes, Rhi- nobatidae) and unidentified skate. 6. Distribution (Fig. 15): Hobart, King Island, King- ston (Tasmania) - Pacific Ocean; Subantarctic region of Atlantic Ocean, S. Sandwich Is., S. Orkney Is., S. Georgia, S. Shetland Is., Elephant I., Graham Land - Atlantic sector, Ross Sea, Mawson Sea - Pacific sector. References: Hickman 1947, Llewellyn 1966, Meyer & Burreson 1990. Genus Moorebdellina Epstein, 1972 Type species: Pontobdella rugosa Moore, 1938 Not large leeches. Body long and cylindrical or short and subcylindrical, with large papillae. Complete somite strictly divided into 2-4 annuli (additional annulation may occur). Anterior sucker: medium-sized, eccentrically at- tached, with tubercles and papillae, and two eye-like spots, mouth-pore centrally located. FIG. 16. Distribution of the gen. Moorebdellina: 1— M. rugosa; 2 — M. biannulata, 3 — M. uschakovi; 4 — M. meyeri. 30 UTEVSKY, ANTARCTIC PISCICOLID LEECHES FIG. 17. Moorebdellina rugosa, external and internal structure: 1, common view; 2, anterior sucker (dorsal and ventral view); 3, posterior sucker; 4, clitellum (ventral view); 5, somite; 6, digestive system; 7, reproductive system (dorsal view); 8, reproductive system (lateral view). BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 54/2007 FIG. 18. Moorebdellina rugosa, lateral view. Posterior sucker: medium sized, centrally attached, facing directly posteriorly, ocelli absent. Digestive system. Esophageal diverticula present or absent. Crop chambers developed in varying de- grees, without lateral processes. Posterior crop caeca nontotally fused with 5 fenestrae without lateral pro- cesses. Intestinal chambers varying in their develop- ment, lateral processes present or absent. Rectum varying in its development. Reproductive system. Four to six pairs of testisacs. Seminal reservoirs long, loop-like. Common parts not great. Accessory glandular mass small, covering pos- terior surface of common part of ejaculatory ducts. Bursa short or long. Ovisacs not extending posterior to ganglion 8. Conductive tissue present. Copulatory area absent. Coelomic system. Dorsal, ventral, lateral, mar- ginal, and testicular lacunae present. Main and ac- cessory connections present. Small pulsatile vesicles (dilations of marginal lacunae) located at site of connection with lateral lacuna. Host data: only one specimen was collected from Pycnogonida. a2 Distribution (Fig. 16): Atlantic, Indian and Pa- cific sector of Antarctic region. Congenitors: M. biannulata Moore, 1957; M. uschakovi Epstein, 1974; M. meyeri A.Utevsky, 1997. Found only in Antarctic region. References: Moore 1938, Epshtein 1972, Sawyer 1986, Utevsky 2000. Key to the Species of the Genus Moorebdellina la Complete somite 2-annulatey eee nn... ELSE ee M. biannulata (p. 34) lb Complete somite more than 2-annulate ... 2 2a Complete somite 4-annulate; Al with 6 pa- pillae; A>, Bs, Be with 8:papillacsyeen...:-<.... Ree ree REN M. rugosa (p. 32) 2b Complete somite 3-annulate ee... 3 3a Annuli A; and A3 with many small tubercles, A with 4 papillae ......... M. uschakovi (p. 36) 3b Annulus A; with 6 papillae, Ao, and Az with 8 papillae nun M. meyeri (p. 38) Moorebdellina rugosa (Moore, 1938) (Syn. Pontobdella rugosa Moore, 1938) 1. External characters (Figs. 17, 18). Medium-sized leeches. Up to 40.5 mm in length and 2.5 mm in width. Body long, cylindrical, distinctly divided into trachelosome and urosome. Anterior sucker small, distinctly divided from trachelosome, eccentrically attached; with two eye-like spots and small tubercles and papillae; mouth-pore centrally located (Fig. 17: 2). Posterior sucker medium-sized, but larger than an- terior sucker, centrally attached, facing directly posteriorly, divided into 2 rings by furrows, without papillae (Fig. 17: 3). Annulation. Clitellum 6-annulate (Fig. 17: 4). Male gonopore located between annuli 1 and 2, fe- male gonopore between 4 and 5. External copulatory area absent. Complete somite 4-annulate (annulus A3 include annuli Bs, Be), and annulus A; with large papillae; Ao, Bs, Be with 8 large papillae 2 or 3 small papillae situated between large tubercles; additional annulation sometimes present (Fig. 17: 5). Anus on first annulus from posterior sucker (Fig. 17: 3). Coloration. Anterior sucker with transverse bar which unites a pair of eye-like spots. Radial brownish stripes on posterior sucker. Its dorsal surface light brownish with unpigmented spots around papillae; its ventral surface light-colored. Trachelosome with 5 pigmented bands. Each side of urosome with 13 oval UTEVSKY, ANTARCTIC PISCICOLID LEECHES fh E = FIG. 19. Moorebdellina biannulata, external and internal structure: 1, common view; 2, anterior sucker (dorsal and ventral view); 3, posterior sucker; 4, clitellum (ventral view); 5, somite (dorsal view); 6, digestive system; 7, reproductive system (dorsal view); 8, reproductive system (lateral view); 9, coelomic system (reconstruction). 33 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 54/2007 unpigmented spots on 1-3 annuli of somite sharply narrowing on more pigmented annulus 4. 2. Digestive system (Fig. 17: 6). Base of proboscis at ganglion 3 of ventral nerve cord. Esophageal di- verticula absent. Crop chambers weakly developed, lateral processes absent. Posterior crop caeca nonto- tally fused with 5 fenestrae, without lateral processes. 5 well developed intestinal chambers, without lateral processes. Rectal dilation weakly developed. 3. Reproductive system (Fig. 17: 7, 8). 3 or 4 pairs of testisacs. Long seminal reservoirs extended to gang- lion 10 of ventral nerve cord to coil. Ejaculatory ducts short, terminal parts wide. Accessory glandular mass covering posterior surface of common part of ejacu- latory ducts. Common part small, bursa short. Ovi- sacs long, loop-like, extending to ganglion 8. Con- ductive tissue in form of two cords connected with ovisacs and fused posterior to vagina. 4. Coelomic system. Structure unknown. 5. Host data: one specimen collected from Pyc- nogonida. 6. Distribution (Fig. 16): Antarctic shelf between 50°-110°E - Indian sector of Antarctic region, En- derby Land - Atlantic sector. References: Moore 1938; Epshtein 1972. Moorebdellina biannulata (Moore, 1957) (Syn. Pontobdella biannulata Moore, 1957) 1. External characters (Figs. 19, 20). Small or not large leeches. Up to 20 mm in length and 3.5 mm in width. Body long, cylindrical, distinctly divided in- to trachelosome and urosome (Fig. 19: 1). Anterior sucker medium-sized, distinctly divided from tra- chelosome, eccentrically attached, with two eye-like spots and small tubercles, mouth-pore centrally lo- cated (Fig. 19: 2). Posterior sucker small, smaller than anterior sucker, centrally attached, facing directly posteriorly, divided by furrows into 4 rings with pa- pillae (Fie; 19: 3). Annulation. Clitellum 4-annulate, with secondary annulation (Fig. 19: 4). Male gonopore located in pos- terior part of annulus 1, female gonopore in posterior part of annulus 3. External copulatory area absent. Complete somite 2-annulate. Annulus 1 with 6 large tubercles, annulus 2 with 8 tubercles, 2 or 3 papillae situated between tubercles; skin with longitudinal and transverse folds (Fig. 19: 5). Anus separated by 4 an- nuli from posterior sucker (Fig. 19: 3). Coloration. Anterior sucker with two brownish transverse bars separated by unpigmented area. Se- cond bar uniting a pair of eye-like spots. Radial light 34 brownish stripes on posterior sucker. Segmental bands of same color alternated with unpigmented areas on dorsum. 2. Digestive system (Fig. 19: 6). Base of proboscis at ganglion 3 of ventral nerve cord. Esophageal diverti- cula located between ganglia 4 and 5. Crop chambers weakly developed, lateral processes absent. Posterior crop caeca nontotally fused with 5 fenestrae, without lateral processes, extending to ganglion 21. Intestinal chambers weakly developed, without lateral proces- ses. Rectal dilation weakly developed. 3. Reproductive system (Fig. 19: 7, 8). Four or five pairs of testisacs. Seminal reservoirs long, loop-like, usually extending to ganglion 12 of ventral nerve cord, enlarged at ganglion 12. Initial parts of ejaculatory ducts short, loop-like; terminal parts wide, oval. Ac- cessory glandular mass covering posterior surface of terminal and common parts of ejaculatory ducts and bursa. Common part small, bursa short. Ovisacs short, voluminous, usually extending to ganglion 7. Conductive tissue in form of mass located adjacent to bursa and connected by two cords with ovisacs. 4. Coelomic system (Fig. 19: 9). Dorsal, ventral, lateral, testicular and marginal lacunae present. Main connections connecting dorsal and ventral lacunae at each ganglion. Accessory connections connecting each FIG. 20. Moorebdellina biannulata, lateral view. UTEVSKY, ANTARCTIC PISCICOLID LEECHES FIG. 21. Moorebdellina uschakovi, external and internal structure: 1, common view; 2, anterior sucker (dorsal and ventral view); 3, posterior sucker; 4, clitellum (ventral view); 5, somite (dorsal view); 6, digestive system; 7, reproductive system (dorsal view); 8, reproductive system (lateral view); 9, coelomic system (reconstruction). 3) BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 54/2007 testicular and dorsal lacunae. Lateral lacunae con- necting main connections with accessory connections. Two pairs of small pulsatile vesicles (dilations of mar- ginal lacunae) per mid-body somite. Pulsatile vesicles of one pair connected with main connection. Pulsa- tile vesicles of another pair connected with accessory connection. 5. Host data: all specimens were collected free- living. 6. Distribution (Fig. 16): Princess Martha Coast, Riiser-Larsen Sea - Atlantic sector of the Antarctic region; between 50°-70°E - Indian sector; Oates Coast and Cape Adare - Pacific sector. References: Moore 1957, Epshtein & Okun’ 1991. Moorebdellina uschakovi Epstein, 1974 1. External characters (Figs. 21, 22). Not large leeches. Up to 25 mm in length and 3.0 mm in width. Body short (may be stretching), cylindrical, distinctly divided into trachelosome and urosome (Figs. 21: 1; 22). Anterior sucker small, distinctly divided from trachelosome, eccentrically attached; with two eye-like spots, papillae and tubercles, mouth-pore centrally located (Fig. 21: 2). Posterior sucker small, but larger than anterior sucker, centrally attached, facing directly posteriorly, without papillae (Fig. 21: 3). Annulation. Clitellum 6-annulate (Fig. 21: 4). Male gonopore located between annuli 1 and 2, female gonopore between 4 and 5. External copulatory area absent. Complete somite 3-annulate, annuli A; and Az with many papillae, A2 with 4 tubercles (two lateral papillae larger than two dorsal), 2 or 3 papillae sit- uated between tubercles on dorsum of Az (Fig. 21: 5). Anus on third annulus from posterior sucker (Fig. 215). Coloration. Dorsal surface light brownish with tints of violet, ventral surface light. Annuli A; and A3 pigmented, A, without pigment. 2. Digestive system (Fig. 21: 6). Base of proboscis at ganglion 3 of ventral nerve cord. Esophageal diver- ticula absent. Five diamond-shaped crop chambers without lateral processes. Posterior crop caeca non- totally fused with 5 fenestrae, without lateral proces- ses, extending to ganglion 20. Three intestinal cham- bers with lateral processes and “folded organ” [Fo]. Rectal dilation present. 3. Reproductive system (Fig. 21: 7, 8). Five pairs of testisacs. Seminal reservoirs well developed, in form of a number of loops, extending posterior to gang- lion 12 of ventral nerve cord. Initial parts of ejacula- 36 FIG. 22. Moorebdellina uschakovi, lateral view. tory ducts short, terminal parts wide, kidney-shaped. Accessory glandular mass covering posterior surface of common parts of ejaculatory ducts and bursa. Common part short and wide, bursa long. Ovisacs long, in form of a number of loops located between ganglia 6 and 9. Conductive tissue in form of mass located posterior to female gonopore and connected by two cords with ovisacs. 4. Coelomic system (Fig. 21: 9). Dorsal, ventral, lateral, testicular, and marginal lacunae present. Main connections connecting dorsal and ventral lacunae at each ganglion. Accessory connections connecting each testicular and dorsal lacunae. Lateral lacunae con- necting main connections with accessory connections. One pair of pulsatile vesicles (dilations of marginal lacunae) per complete somite. Pulsatile vesicles connected with main connection. nennen UTEVSKY, ANTARCTIC PISCICOLID LEECHES Atrer I en. seta : APRs yyrac 10 mm FIG. 23. Moorebdellina meyeri, external and internal structure: 1, common view; 2, anterior sucker (dorsal and ventral view); 3, posterior sucker; 4, clitellum (ventral view); 5, somite (dorsal view); 6, digestive system; 7, reproductive system (dorsal view); 8, reproductive system (lateral view); 9, coelomic system (reconstruction). 37 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 54/2007 5. Host data: all specimens were collected free- living. 6. Distribution (Fig. 16): Crozet Is. - Indian sec- tor of the Antarctic region; S. Georgia I. - Atlantic sector. References: Epshtein 1974. Remark: this species has been included in the genus Moorebdellina by Epshtein (1974). However coelomic system differs from those in other members of Pontobdellinae by having only one pairs of pulsa- tile vesicles per somite. Thus, the subfamilia affiliati- on open to question. Moorebdellina meyeri A.Utevsky, 1997 1. External characters (Figs. 23, 24). Not large leeches. Up to 23.5 mm in length and 1.5 mm in width. Body long, cylindrical, distinctly divided into trachelosome and urosome (Figs. 23: 1; 24). Anterior sucker small, distinctly divided from trachelosome, eccentrically attached; with two eye-like spots and papillae; mouth- pore centrally located (Fig. 23: 2). Posterior sucker small, smaller than anterior sucker, centrally attached, facing directly posteriorly, without papillae (Fig. 23: 3). Annulation. Clitellum 6-annulate (Fig. 23: 4). Male gonopore located between annuli 1 and 2, female gonopore between 4 and 5. External copulatory area absent. Complete somite 3-annulate, annulus A; with 6 tubercles, Ay and Az with smaller tubercles (Fig. 23: 5). Anus on first annulus from posterior sucker (Fig. 23:3): Coloration. Anterior sucker with two pigmented area around eye-like spots. Body with light pigment bands (the examined specimens most likely are un- pigmented after alcohol fixation). 2. Digestive system (Fig. 23: 6). Base of proboscis located between ganglia 3 and 4 of ventral nerve cord. Esophageal diverticula absent. Crop chambers weakly developed, without lateral processes. Posterior crop caeca nontotally fused with 5 fenestrae, without la- teral processes, extending to posterior ganglion mass. Posterior portion of posterior crop caeca separated. 5 intestinal chambers well developed, without lateral processes. Rectal dilation present. 3. Reproductive system (Fig. 23: 7, 8). Six pairs of testisacs. Seminal reservoirs long, loop-like, usually located between ganglia 6 and 7 of ventral nerve cord, very coiled. Initial parts of ejaculatory ducts short, loop-like; terminal parts wide. Accessory glandular mass small, covering posterior surface of common part. Common part small, bursa short. Ovisacs short, 38 usually located between ganglia 6 and 7. Conductive tissue in form of mass located between ganglia 6 and 7, and two cords extending to vagina and connected with ovisacs. 4. Coelomic system (Fig. 23: 9). Dorsal, ventral, lateral, testicular, and marginal lacunae present. Main connections connecting dorsal lacuna with ventral lacuna near each ganglion. Accessory connections of 1-2 pair of testisacs connecting testicular lacunae with ventral lacuna. Accessory connection of 3-6 pairs of testisacs connecting testicular lacunae with dorsal lacuna. Lateral lacunae connecting main connections with accessory connections. Iwo pairs of small pulsa- tile vesicles (dilations of marginal lacunae) per com- plete somite. Pulsatile vesicles of one pair connected with main connection. Pulsatile vesicles of another pair connected with accessory connection. The num- ber of vesicles per somite may be inconstant. 5. Host data: all specimens were collected free- living. FIG. 24. Moorebdellina meyeri, dorso-lateral view. 6. Distribution (Fig. 16): S. Orkney Is.- Atlantic sector of the Antarctic region. References: Utevsky 1997. Genus Megaliobdella Meyer & Burreson, 1990 Type species: Megaliobdella szidati Meyer & Burreson, 1990 Very large leeches. Body long, cylindrical, tapering to suckers, with large, cone-like tubercles. Complete somite sharply divided into annuli. External copula- tory area on clitellum absent. Anterior sucker: small, eccentrically attached, sharply separated by neck from trachelosome; with one pair of eyes; with transverse furrows on dorsal sur- face. Posterior sucker: small, indistinctly divided from urosome; without ocelli, with papillae on dorsal sur- face. Structure of Digestive, Reproductive and Coelo- mic systems poorly known. FIG. 25. Distribution of the Megaliobdella szidati. UTEVSKY, ANTARCTIC PISCICOLID LEECHES No host data. Monotypic genus. Distribution (Fig. 25): Pacific sector of the Ant- arctic region. Reference: Meyer & Burreson 1990. Remark: Megaliobdella szidati was placed only provisionally in the subfam. Pontobdellinae (Meyer & Burreson 1990). The description of this species and genus was based on external characters. Thus, the subfamiliar affiliation of M. szidati is open to ques- tion. Megaliobdella szidati Meyer & Burreson, 1990* 1. External characters (Fig. 26). Very large leeches. Up to 340 mm in length and 13 mm in width. Body long, cylindrical, indistinctly divided into trachelo- some and urosome, enlarged from anterior to poste- rior sucker, with many truncated conic tubercles on trachelosome and urosome. Anterior sucker small, sharply separated by neck from trachelosome, eccen- 93 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 54/2007 FIG. 26. Megaliobdella szidati, external and internal structure: 1, common view; 2, clitellum (ventral view); 3, somite; 4, elements of digestive system (dorsal view) 5, anterior sucker (lateral view); 6, posterior sucker (ventral view). trically attached, with 9 transverse furrows on dorsal surface; one pair of eyes present (Fig. 26: 4). Posterior sucker small, but larger than anterior sucker, terminal, indistinctly divided from urosome, eccentrically at- tached, with many tubercles similar to those on body, margin of sucker with smaller tubercles, ocelli absent (Fig. 26:5). Annulation. Preclitellar region 60-72-annulate, clitellum 12-17-annulate, male gonopore on annulus 1, female gonopore between annuli 10 and 11 (copu- latory area absent) (Fig. 26: 2). Complete somite 12- annulate with 18-25 large tubercles on each annulus and some small tubercles (papillae) (Fig. 26: 3). Coloration. Dorsal surface with two brownish stripes. Ventrum lighter. Segmental bands absent. Body of preserved specimens light brown. 2. Digestive system. Structure poorly known. Last chamber of crop with 3 pairs of short lateral proces- ses. Posterior crop caeca nontotally fused, with 3 40 chambers and 3 fenestrae, without lateral processes. Intestine wide, without lateral processes and distinct chambers. Rectal dilation absent. 3. Reproductive system. Structure unknown. 4. Coelomic system. Structure unknown. 5. No host data. 6. Distribution (Fig. 25): collected at 77°42.0’S, 167°22.0’W (Ross Sea) - Pacific sector of the Antarctic region. Reference: Meyer & Burreson 1990. “Subfamily Platybdellinae” Diagnosis. Leeches of varying sizes; body long or short, cylindrical, subcylindrical or flattened. Exter- nally visible pulsatile vesicles absent. Tubercles, pa- pillae, marginal flanges, eyes, eye-like spots, segmen- tal ocelli, ocelli on posterior sucker absent or present. Digestive system of varying structure, posterior crop caeca of varying degrees of fusion (from totally free to totally fused) or absent. Reproductive system of varying structure. Coelomic system of varying structure: from sim- ple coelomic system consisting of ventral lacunae on- ly to well developed system with connections, lateral and marginal lacunae. Pulsatile vesicles absent. References: Sawyer 1986; Epshtein er al. 1994. Key to the Antarctic Genera of the “Subfamily Platybdellinae” la Body sharply divided into trachelosome and urosome, wide and flattened 1b Body not sharply divided into trachelosome and urosome, not wide and not flattened ... 3 2a Eye, eye-like spots, ocelli absent; tegument opa- que; posterior crop caeca absent Genus Epsteinia (p. 41) 2b 2 eye-like spots on anterior sucker; tegument translucent with pigment cells; posterior crop caeca nontotally fused with 4 fenestrae Genus Austrobdella (p. 42) 3a Body with marginal flanges Genus Pleurobdella (p. 46) 3b Body without marginal flanges 4a 10-12 pairs of tubercles on urosome; anterior sucker with eye-like spots; posterior sucker small, facing directly posteriorly Genus Glyptonotobdella (p. 49) 4b Tubercles absent; anterior sucker without eye- like spots; posterior sucker medium-sized or large, Genus Cryobdella (p. 52) facing ventraly Genus Epsteinia A. Utevsky, 1994 Type species: Trulliobdella alba Epstein, 1970 Not large leeches. Body distinctly divided into tra- chelosome and urosome. Segmental ocelli absent. Complete somite 3-annulate with additional annu- lation. Anterior sucker: small, distinctly divided from trachelosome, surface smooth, eyes or eye-like spots absent. Posterior sucker: small, distinctly divided from urosome, surface smooth, ocelli absent. Digestive system. Esophageal diverticula present. Crop chambers diamond-shaped with small lateral processes. Posterior crop caeca absent. Intestine with lateral processes. Rectal dilation present. Reproductive system. Five pairs of testisacs. Se- minal reservoirs long, ejaculatory ducts short, terminal parts wide, common part small, bursa short, acces- FIG. 27. Distribution of the Epsteinia alba. UTEVSKY, ANTARCTIC PISCICOLID LEECHES sory glands absent. Conductive tissue absent, exter- nal copulatory area present. Coelomic system. Structure unknown. Host data: fam. Nototheniidae (Ord. Percifor- mes). Distribution (Fig. 27): Enderby Land - Atlantic sector of the Antarctic region. Monotypic genus. References: Epshtein 1970b, Utevsky 1994. Epsteinia alba (Epstein, 1970) (Syn. Trulliobdella alba Epstein, 1970) 1. External characters (Figs. 28, 29). Not large leeches. Up to 30 mm in length and 8 mm in width. Body short and wide, distinctly divided into subcylindrical trachelosome and flattened urosome. Anterior sucker small, distinctly divided from trachelosome, centrally attached, surface smooth, mouth-pore centrally lo- 4] BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 54/2007 FIG. 28. Epsteinia alba, dorsal and ventral view (Photo. Epshtein V.M.). cated (Figs. 28, 29: 2). Posterior sucker small, but larger than anterior one, excentricaly attached, sur- face smooth (Figs. 28, 29: 3). Annulation. Clitellum 8-annulate (Fig. 29: 4). Male gonopore located in depression between annuli 3 and 4, female gonopore in depression between annuli 6 and 7. 3rd annulus enlarged in central part, 4th bent posteriorly and covering annuli 5 and 6, annulus 7 bent posteriorly also. Copulatory area in depression around female gonopore. Complete somite 3-annu- late with additional annulation of 2nd degree. Di- mensions of annuli variable, without tubercles or papillae, wrinkled. Coloration. Body light colored but dorsum more dark than venter. No numerous large brown pigment cells visible in skin. 2. Digestive system (Fig. 29: 5). Base of proboscis located between ganglia 3 and 4 of ventral nerve cord. Large esophageal diverticula between ganglia 4 and 5. Five diamond-shaped crop chambers with small lateral processes. Posterior crop caeca absent. Five well 42 developed intestinal chambers with voluminous lateral processes. Rectal dilation weakly developed. 3. Reproductive system (Fig. 29: 6, 7). Five pairs of testisacs. Seminal reservoirs well developed in form of some loops between ganglia 6 and 7 of ventral nerve cord. Initial parts of ejaculatory ducts slightly coiled, sharply divided from more wide terminal parts. Accessory glands absent. Common part small, bursa short. Ovisacs voluminous extending behind ganglion 8. Conductive tissue absent. 4. Coelomic system. Structure unknown. 5. Host data: on caudal fin of Trematomus borch- grevinki. 6. Distribution (Fig. 27): Enderby Land - Atlantic sector of the Antarctic region. References: Epshtein 1970b, Utevsky 1994. Genus Austrobdella Badham, 1916 Type species: Austrobdella translucens Badham, 1916. Not large or small leeches. Body flattened, distinctly divided into trachelosome and urosome, surface UTEVSKY, ANTARCTIC PISCICOLID LEECHES eed Tye “ ALIEN coy OR > 6% Kae ey Ig ihe FIG. 29. Epsteinia alba, external and internal structure: 1, dorsal view; 2, anterior sucker (ventral view); 3, poosterior sucker (ventral view); 4, clitellum (ventral view); 5, digestive system (dorsal view); 6, reproductive system (dorsal view); 7, reproductive system (lateral view). 43 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 54/2007 smooth, without ornamentation. Complete somite 6-annulate, without segmental ocelli and pigment bands. Anterior sucker: small, distinctly divided from tra- chelosome, surface smooth, eye-like spots present or absent. Posterior sucker small, distinctly divided from urosome, surface smooth, ocelli absent. Digestive system. Esophageal diverticula present or absent. Crop chambers well developed, lateral pro- cesses present or absent. Posterior crop caeca nonto- tally fused, with fenestrae. Intestine with chambers, lateral processes present or absent. Reproductive system. Bursa long, accessory glands present. Conductive tissue and copulatory area absent, vagina long. Coelomic system. Dorsal, ventral, marginal and testicular lacunae present. Main connections absent, accessory connections present or absent. FIG. 30. Distribution of the Austrobdella translucens. a) Host data: Sillago ciliata (A. translucens), Rhom- bosolea tapirina (A. bilobata); Hippoglossina stomata, Pleuronichthis verticalis, P decurrens, Scorpaena guttata (A. californiana) Distribution (Fig. 30): coastal waters of Green- land, California, Tasmania, Australia and East Ant- arctica. Congenitors: A. anoculata Moore, 1940 - Greenland A. bilobata Ingram, 1957 - Tasmania, Pacific Ocean A. californiana Burreson, 1977 - California, Pacific Ocean References: Badham 1916, Sawyer 1986, Epsh- tein et al. (1994). Austrobdella translucens Badham, 1916* 1. External characters. Small leeches. Up to 13 mm in length and 3.25 mm in width. Body short and wide, distinctly divided into subcylindrical trache- UTEVSKY, ANTARCTIC PISCICOLID LEECHES S III SI S FIG. 31. Austrobdella translucens, external and internal structure (reproduced and modified from Badham 1916). losome and flattened urosome (Fig. 31: 1). Anterior sucker small, distinctly divided from trachelosome, surface smooth with pair of eye-like spots, mouth- pore centrally located ( Fig. 31: 2). Posterior sucker small, but larger than anterior one, distinctly divided from urosome, surface smooth without ocelli. Annulation. Complete somite 3-annulate with additional annulation of 2nd degree. D) BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 54/2007 Coloration. Skin semitransparent with brownish- red, yellow and purple individual pigment cells. 2. Digestive system (Fig. 31: 3). Base of proboscis near ganglion 3 of ventral nerve cord. Esophageal diverticula between ganglia 4 and 5. Six small crop chambers with long lateral processes which may be bifurcated at their endings. Posterior crop caeca non- totally fused with 4 fenestrae and 6 lateral processes which may be bifurcated to end. Intestine with 3 large chambers. Rectal dilation absent. 3. Reproductive system (Fig. 31: 3, 4). Five pairs of testisacs. Seminal reservoirs short, coiled at ganglion 6 of ventral nerve cord. Initial parts of ejaculatory ducts short, loop-like, terminal parts wide, common part voluminous, bursa long. Accessory glandular mass covering terminal and common parts of male reproductive system. Ovisacs long, extending to gang- lion 12, vagina long and glandular. Ovisacs with pro- jections in place of connections with vagina. Con- ductive tissue and copulatory area absent. 4. Coelomic system (Fig. 31: 5). Dorsal, ventral, marginal lacunae present. Main connection comple- tely closed: connecting dorsal and ventral lacunae. Marginal lacunae connected with main connections and hypodermal lacunae. 5. Host data: Collected from Sillago ciliata - Pa- cific Ocean, Notothenia sp. and Chaenocephalus sp. in the Antarctic region. 6. Distribution (Fig. 30): Southern Coast of Aus- tralia, Kerguelen Is. - Indian sector of the Antarctic region. References: Badham 1916, Moore 1957. Genus Pleurobdella A.Utevsky, 1995 Type species: Oxytonostoma varituberculata Moore, 1938 Not large leeches. Body long, cylindrical or sub- cylindrical, with tubercles, papillae, and/or marginal flanges from anterior to posterior sucker. Complete somite 6-annulate, with additional annulation. Seg- mental pigmentation present. FIG. 32. Distribution of the gen. Pleurobdella. 1 — P varituberculata; 2 — P australis. 46 Anterior sucker: various size, eccentrically at- tached, with pair of eye-like spots, mouth-pore cen- trally located. Posterior sucker: small, indistinctly divided from urosome, facing directly posteriorly. Digestive system. Small esophageal diverticula pre- sent. Crop chambers with small double lateral pro- cesses. Posterior crop caeca nontotally fused, with 5 fenestrae. Intestine chambers with small lateral pro- cesses. Rectal dilation present. Reproductive system. Five or six pairs of testisacs. Seminal reservoirs long, loop-like. Common part small. Accessory glands absent. Bursa long, with in- ternal copulatory area. Ovisacs long. Conductive tis- sue well developed, begining prior to terminal parts of male reproductive system, rounding terminal parts, common part, and copulatory bursa and extending to posterior parts of ovisacs. External copulatory area absent or present. Coelomic system. Dorsal, ventral, marginal and testicular lacunae present. Main and accessory con- nections absent. Host data: Pycnogonida. Distribution (Fig. 32): Atlantic and Indian parts of the Antarctic region. Congenitors: P australis (Epstein, 1970). References: Moore 1938, Utevsky 1995, Utevsky 2000. Key to the Species of the Genus Pleurobdella la Anterior sucker small, body with small tuber- P varituberculata (p. 47) 1b Anterior sucker large, body with small papil- PR australis (p. 49) Pleurobdella varituberculata (Moore, 1938) (Syn. Oxytonostoma varituberculata Moore, 1938) 1. External characters (Figs. 33, 34). Not large leeches. Up to 27 mm in length and 2.4 mm in width. Body long, subcylindrical, sharply narrowing to anterior sucker, indistinctly divided into trachelosome and urosome. Body surface with small tubercles and pa- pillae. Two marginal flanges extending from anterior to posterior sucker (Fig. 33: 1). Anterior sucker small, distinctly divided from trachelosome, eccentrically attached; papillae and eyes-like spots on dorsal sur- face, edge rough, mouth-pore centrally located (Fig. 33: 2). Posterior sucker small, but larger than anterior UTEVSKY, ANTARCTIC PISCICOLID LEECHES sucker, indistinctly divided from urosome, centrally attached, facing directly posteriorly, divided by 2 fur- rows into 3 rings with papillae (Fig. 33: 3). Annulation. Clitellum 5-annulate with additional annulation of 2nd degree, annulus 5 more shorter than other (Fig. 33: 4). Male gonopore at posterior part of annulus 2 on tubercle; female gonopore in centre of annulus 5, external copulatory area absent. Complete somite 6-annulate with obscure ad- ditional annulation. Each annulus may be divided into 3 annuli, annulus 2 with small tubercles. Anus between annuli 1 and 2 from posterior sucker (Fig. 33,9): Coloration. Margin of anterior sucker brownish. Posterior sucker with brownish stripe on dorsum which narrowing to margin. Body with bands of same colors, dorsum more colored than venter. Three bands on trachelosome and 15 on urosome. On urosome, annuli B1-B3 bearing segmental bands. This annuli more pigmented than B4-B6 which unpigmented on their lateral surface. Annuli B4-B6 longer than B1- Bo: 2. Digestive system (Fig. 33: 6). Base of proboscis located anterior to ganglion 3 of ventral nerve cord. Esophageal diverticula posterior to ganglion 3. Five voluminous crop chambers with small double lateral processes. Posterior crop caeca nontotally fused, with 5 fenestrae, separated anterior to ganglion 21 and ex- tending to posterior ganglion mass, lateral processes absent. Five intestinal chambers with small lateral pro- cesses. Rectal dilation weakly developed. 3. Reproductive system (Fig. 33: 7, 8). Five pairs of testisacs. Seminal reservoirs long, forming 2 loops between ganglia 8 and 9. Initial parts of ejaculatory ducts loop-like, terminal parts wide. Accessory glands absent. Common part medium-sized, bursa long. Ovisacs medium-sized, wide, extending almost to ganglion 8. Hollow conductive tissue mass beginning at ganglion 4 and deviding into 2 cords rounding ter- minal, common parts and bursa, fusing at ganglion 6 to form mass located between ganglia 7 and 8. In- ternal copulatory area formed by conductive tissue cords adjoining to bursa. 4. Coelomic system (Fig. 33: 9). Dorsal, ventral, testicular and marginal lacunae present. Main connec- tions absent: ventral lacuna enlarged at ganglia, not extending to dorsal lacuna. Accessory connections absent: testicular lacunae not joining with dorsal and ventral lacunae. Marginal lacunae not traced for their entire length. 47 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 54/2007 Sat oe Do" LAO 29 D0 > } 7 —s PE aT 722.930 ar 4 Eee f FIG. 33. Pleurobdella varituberculata, external and internal structure: 1, common view; 2, anterior sucker (dorsal and ventral view); 3, posterior sucker; 4, clitellum (ventral view); 5, somite (dorsal view); 6, digestive system; 7, reproductive system (dorsal view); 8, reproductive system (lateral view); 9, coelomic system (reconstruction). 48 FIG. 34. Pleurobdella varituberculata, dorsal view. 5. Host data: only one specimen collected from Pycnogonida. 6. Distribution (Fig. 32): S. Orkney Is., Enderby Land - Atlantic sector of the Antarctic region; Wil- kes Land - Indian sector. References: Moore 1938, Utevsky 1995. Pleurobdella australis (Epstein, 1970) (Syn. Pterobdellina australis Epstein, 1970) 1. External characters (Figs. 35, 36). Not large leeches. Up to 27 mm in length and 2.4 mm in width. Body long, subcylindrical, sharply narrowing to anterior sucker, indistinctly divided into trachelosome and urosome. Body surface with small papillae. Two mar- ginal flanges extending from anterior to posterior sucker (Fig. 35: 1). Anterior sucker large, distinctly divided from trachelosome, eccentrically attached; surface smooth with oval eye-like spots, mouth-pore centrally located (Fig. 35: 2). Posterior sucker small, not larger than anterior sucker, indistinctly divided from urosome, eccentrically attached, facing directly posteriorly, surface smooth (Fig. 35: 3). Annulation. Clitellum 8-annulate with additional annulation of 2nd-3rd degree. Male gonopore on tu- bercle between annuli 3 and 4 (2 folds formed ex- ternal copulatory area), female gonopore between UTEVSKY, ANTARCTIC PISCICOLID LEE( annuli 6 and 7 (Fig. 35: 4). Complete somite 6-annu- late with additional annulation, surface smooth. Anus on annulus 3 from posterior sucker (iis. 552-9), Coloration. Margin of anterior sucker brownish. Radial brownish stripes on posterior sucker. Body pig- mented by brownish pattern against gray background which may form some black bands on trachelosome. 2. Digestive system (Fig. 35: 6). Base of proboscis located between ganglia 3 and 4 of ventral nerve cord. Esophageal diverticula between ganglia 4 and 5. Six voluminous crop chambers with small double lateral processes. Posterior crop caeca nontotally fused, with 5 fenestrae, separated anterior to ganglion 21 and ex- tending to posterior ganglion mass, lateral processes absent. Five intestinal chambers with small lateral pro- cesses. Rectal dilation weakly developed. 3. Reproductive system (Fig. 35: 7, 8). 6 pairs of testisacs. Seminal reservoirs long, forming loops poste- rior to ganglion 8. Initial parts of ejaculatory ducts loop-like, terminal parts wide. Accessory glands ab- sent. Common part short and wide, bursa long. Ovi- sacs long, wide, extending posterior to ganglion 8. Hollow conductive tissue mass beginning at ganglion 5 and dividing into 2 cords rounding terminal, com- mon parts and bursa, fusing at ganglion 6 to form mass extending posterior to ganglion 8. Internal co- pulatory area formed by conductive tissue cords ad- joining to bursa. 4. Coelomic system (Fig. 35: 9). Dorsal, ventral, testicular and marginal lacunae present. Main connec- tions absent: ventral lacuna forming small lateral pro- cesses at ganglia, not extending to dorsal lacunae. Accessory connections absent: testicular lacunae not joining with dorsal and ventral lacunae. 5. Host data: only one specimen collected from Pycnogonida. 6. Distribution (Fig. 32): Atlantic Ocean near Argentina, S. Orkney Is. - Atlantic sector of the Ant- arctic region. References: Epshtein 1970a, Utevsky 1995. Genus Glyptonotobdella Sawyer & White, 1969 Type species: Glyptonotobdella antarctica Sawyer & White, 1969 Not large leeches. Body long, cylindrical, indistinct- ly divided into urosome and trachelosome. Complete somite 12-annulate, with pair of lateral tubercles, seg- mental ocelli absent. Anterior sucker: small, distinctly divided from tra- chelosome, eccentrically attached, with one pair of eye-like spots. 49 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 54/2007 FIG. 35. Pleurobdella australis, external and internal structure: 1, common view; 2, anterior sucker (dorsal and ventral view); 3, posterior sucker; 4, clitellum (ventral view); 5, somite (dorsal view); 6, digestive system; 7, reproductive system (dorsal view); 8, reproductive system (lateral view); 9, coelomic system (reconstruction). 50 FIG. 36. Pleurobdella australis, dorsal view. Posterior sucker: small, centrally attached, cup- shaped, facing directly posteriorly, ocelli absent. Digestive system. Esophageal diverticula absent. Crop chambers without lateral processes. Posterior crop caeca nontotally fused, with fenestrae. Intesti- nal chambers and rectal dilation weakly developed. Reproductive system. Six pairs of testisacs. Bursa long, accessory glands absent. Ovisacs short, con- ductive tissue and copulatory area absent. Coelomic system. Dorsal lacuna absent. Main and accessory connections absent. Host data: Glyptonotus antarcticus (Ord. Isopoda), fam. Chaenichthyidae (Ord. Perciformes). Distribution (Fig. 37): Atlantic and Indian parts of the Antarctic region. Monotypic genus. References: Sawyer & White 1969, Sawyer 1976, Sawyer 1986, Janssen 1993. UTEVSKY, ANTARCTIC PISCICOLID LEECHI Glyptonotobdella antarctica Sawyer & White, 1969 (Syn. Notobdella streptocheles Yang, 1987) 1. External characters. Not large leeches. Up to 26 mm in length and 1.0 mm or more in width. Body long, cylindrical, indistinctly divided into trachelo- some and urosome, with 10-12 pairs of lateral tu- bercles on urosome (Fig. 38: 1). Anterior sucker small, distinctly divided from trachelosome, eccentrically attached; edge rough, surface smooth with pair of eye- like spots on pigment area, mouth-pore centrally located (Fig. 38: 3). Posterior sucker small, not larger than anterior, distinctly divided from urosome, cen- trally attached, facing directly posteriorly, divided by furrows into 4-5 rings, without papillae (Fig. 38: 4). Annulation. Clitellum 7(14)-annulate, male go- nopore between annuli 7 and 8, female gonopore be- tween annuli 12 and 13, external copulatory area absent (Fig. 38: 5). Complete somite 12-annulate (may be 12-18 annulate), annulus A; wider than A; and Ao, lateral tubercles on annulus A; (Fig. 38: 6). Anus on annulus 3 from posterior sucker (Fig. 38: 4). Coloration. [rachelosome with 5 red-brown pig- ment bands, urosome with 14 bands of same color, which include annulus Az. Anterior sucker with shoe- like pigment bar which unites eye-like spots, poste- rior sucker with some pigment radial stripes. Colo- ration varying from translucent ochre to light brow- nish-red. 2. Digestive system (Fig. 38: 2). Base of proboscis at ganglion 3 of ventral nerve cord. Esophageal di- verticula absent. Six crop chambers large, without la- teral processes. Posterior crop caeca nontotally fused, with 5 fenestrae, without lateral processes. Intestine chambers weakly developed, without lateral proces- ses. Rectal dilation weakly developed. 3. Reproductive system (Fig. 38: 2). Six pairs of testisacs. Seminal reservoirs narrow. Initial parts of ejaculatory ducts voluminous, loop-like, extending anteriorly to ganglion 5. Terminal parts deflected anteriorly. Common part large, wide. Accessory glands absent. Bursa long (?). Ovisacs short, located between ganglia 6 and 7. Conductive tissue absent. 4. Coelomic system. Dorsal and lateral lacunae absent. Ventral lacuna forming wing-like projections in ganglion region. Main and accessory connections absent. 5. Host data: most specimens collected from G/yp- tonotus antarcticus (Eights, 1852), other specimens Dil BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 54/2007 collected from Chionodraco sp. and Chaenocephalus aceratus. 6. Distribution (Fig. 37): S. Orkney Is., S. Sand- wich Is., Scottia Sea, Princess Martha Coast - Atlan- tic sector of the Antarctic region; McMurdo Sd. (Ross Sea) - Pacific sector; Marion I. - Indian sector. References: Sawyer & White 1969; Yang 1987. Genus Cryobdella Harding, 1922 Type species: Cryobdella levigata Harding, 1922. Not large or small leeches. Body cylindrical or sub- cylindrical. Annulation nondeterminated, surface smooth, without tubercles and segmental ocelli. Pig- mentation variable. Anterior sucker: varying in size, eccentrically at- tached, mouth may be prior to centre, without eyes or eye-like spots. Posterior sucker: large, strictly divided from uro- some, centrally or eccentrically attached, without ocelli. Digestive system. Esophageal diverticula absent or present. Crop chambers varying in their development, without lateral processes. Posterior crop caeca non- totally fused, with fenestrae, without lateral proces- ses. Intestine chambers developed in varying degrees. Rectum varying in their development. Reproductive system. [wo to six pairs of testisacs. Seminal reservoirs short, common part of ejaculatory ducts small, accessory glands absent, bursa short. Ovi- sacs short, conductive tissue and copulatory area ab- sent. Coelomic system. Dorsal, ventral, testicular la- cunae present. Lateral lacunae present or absent. Main and accessory connections developed in varying de- grees. FIG. 37. Distribution of the Glyptonotobdella antarctica. 52 UTEVSKY, ANTARCTIC PISCICOLID LEECHES Host data: fam. Nototheniidae (ord. Perciformes), C. ljadovi Epstein & A.Utevsky, 1994 Muraenolepidae (ord. Gadiformes). C. pallida A.Utevsky, 1997 Distribution (Fig. 39): Atlantic, Indian and Paci- References: Harding 1922, Sawyer 1986, Epsh- fic sectors of the Antarctic region. tein et al. (1994), Utevsky 2000. Congenitors: C. antarctica Epstein, 1970 | jt i / | oer ne FIG. 38. Glyptonotobdella antarctica, external and internal structure (reproduced and modified from Sawyer and White 1969, by permission of R.T. Sawyer, BIOPHARM (Uk) Ltd). N Oo BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 54/2007 Key to the Species of the Genus Cryobdella la Posterior sucker in complicated form, large, possesses two parts which just one to ones C. hadovi (p. 56) 1b Posterior sucker round form, medium-sized or 2a Proportion of posterior sucker C5/D>3>22 ... C. pallıda (p. 58) 2b Proportion of posterior sucker Cy/D> =1 ... 3 3a Posterior sucker centrally attached C. levigata (p. 54) 3b Posterior sucker eccentrically attached C. antarctica (p. 56) Cryobdella levigata Harding, 1922 (Syn. Platybdella levigata (Harding, 1922)) 1. External characters (Fig. 40). Not large leeches. Up to 29 mm in length and 3.5 mm in width. Body long, subcylindrical, distinctly divided into trachelo- some and urosome. Anterior sucker small, distinctly divided from trachelosome, eccentrically attached; surface smooth, edge rough, mouth-pore prior to centre. Posterior sucker large, larger than anterior sucker, centrally attached, facing directly posteriorly surface smooth (Fig. 40: 1). Annulation. Clitellum 8-annulate. Male gonopore located between annuli 3 and 4, female gonopore be- tween annuli 5 and 6. External copulatory area ab- sent. Complete somite 3-annulate, additional annu- lation present, pattern of additional annulation va- riable. Anus between annuli 2 and 3 from posterior sucker. Coloration. After alcohol fixation brownish-grey or fully unpigmented. 2. Digestive system (Fig. 40: 2). Proboscis ex- tending prior to 2nd ganglion of ventral nerve cord. Esophageal diverticula present. Crop chambers 1 and 2 weakly developed without lateral processes, other FIG. 39. Distribution of the gen. Cryobdella. 1 — C. levigata; 2 — C. antarctica; 3 — C. adovi; 4 — C. pallida. 54 UTEVSKY, ANTARCTIC PISCICOLID LEECHES FIG. 40. Cryobdella levigata, external and internal structure: 1, common view (reproduced from Moore 1938); 2, internal structure (reproduced and modified from Harding 1922). four chambers with two pairs of lateral processes. Posterior crop caeca nontotally fused with 5 fenestrae, without lateral processes. Three chambers of intesti- ne well developed, without lateral processes. Rectal dilation well developed. 3. Reproductive system (Fig. 40: 2). Four pairs of testisacs. Seminal reservoirs very short, exending to ganglion 6 of ventral nerve cord. Initial parts of eja- culatory ducts short and thin, terminal parts wide, oval. Accessory glandular mass absent, common part BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 54/2007 very small, bursa very short. Ovisacs long and wide, extending to ganglion 8. Conductive tissue absent. 4. Coelomic system. Structure unknown. 5. Host data: collected from Trematomus hansoni, T. bernacchit. 6. Distribution (Fig. 39): Ross Sea (Victoria Land) - Pacific sector of the Antarctic region; Davis Sea, Mawson Sea, Kerguelen Is. - Indian sector. References: Harding 1922, Moore 1938, Meyer & Burreson 1990. Cryobdella antarctica Epstein, 1970 (Syn. Glyptonotobdella epshteini Sawyer, 1986) 1. External characters (Figs. 41, 42). Not large leeches. Up io 22 mm in length and 1 mm in width. Body long, cylindrical, indistinctly divided into trachelo- some and urosome (Fig. 42). Anterior sucker me- dium-sized, distinctly divided from trachelosome, eccentrically attached, the first one third with many small papillae; the other two thirds without papillae, divided by 6 transverse furrows, mouth-pore centrally located (Fig. 41: 2). Posterior sucker medium-sized, but larger than anterior one, distinctly divided from urosome, eccentrically attached; the first one third adjacent to urosome divided by 2 transverse furrows, other two thirds divided by 5-6 longitudinal furrows, with small papillae (Fig.41: 3). Annulation. Clitellum 4-annulate with additional annulation of second degree (Fig. 41: 4). Male gono- pore located between annuli 2 and 3 of 2nd degree, female gonopore between annuli 5 and 6. External copulatory area absent. Annulus 3 with row of small papillae on dorsum, small papillae on other annuli disorderly located. Complete somite 3-annulate with additional annulation of 2nd degree (Fig. 41: 5). An- nulus Az shorter than others. Annuli with small pa- pillae. Anus between annuli 3 and 4 from posterior sucker (Fig. 41: 3). Coloration. Segmental brownish bands alternated with unpigmented areas. Unpigmented after alcohol fixation. 2. Digestive system (Fig. 41: 6). Base of proboscis posterior to 2nd ganglion of ventral nerve cord. Eso- phageal diverticula absent. Crop chambers weakly developed, without lateral processes. Posterior crop caeca nontotally fused with 5 fenestrae, without la- teral processes. Two chambers of intestine developed without lateral processes. Rectal dilation absent. 3. Reproductive system (Fig. 41: 7, 8). Two to six pairs of testisacs (3-6 pairs may be reduced). Seminal reservoirs well developed, in form of a number of 56 loops, extending to ganglion 6 of ventral nerve cord. Initial parts of ejaculatory ducts short and wide, loop- like, sharply divided from wide terminal parts. Ac- cessory glands absent. Common part small, bursa very short. Ovisacs long and wide, extending to ganglion 8 (may be willed between ganglia 6 and 7), asym- metrical. Conductive tissue absent. 4. Coelomic system (Fig. 41: 9). Dorsal, ventral, lateral and testicular lacunae present. Lateral lacunae with muscular layer. Main connection connecting dorsal lacuna with ventral lacuna near each ganglion. Accessory connections connecting dorsal and testi- cular lacunae. Main and accessory connections connected through lateral lacunae. 5. Host data: one specimen collected from body of Trematomus bernacchit. 6. Distribution (Fig. 39): Davis Sea - Indian sec- tor of the Antarctic region; Scottia Sea - Atlantic sec- tor. References: Epshtein 1970b, Sawyer 1986. Cryobdella hadovi Epstein & A.Utevsky, 1994 1. External characters (Figs. 43, 44). Small leeches. Up to 13.4 mm in length and 2.2 mm in width. Body short, cylindrical, distinctly divided into tracheloso- me and urosome, sharply narrowing to posterior sucker, surface smooth (Fig. 44). Anterior sucker small, indistinctly divided from trachelosome, eccentrically attached; surface smooth, edge rough, mouth-pore prior to centre (Fig. 43: 2). Posterior sucker large, lar- ger than anterior sucker, sharply divided from uro- some, centrally attached, facing directly posteriorly, surface smooth (Fig. 43: 3). It divided into right and left halves. Each half divided into two parts. Left half juts into right one. Annulation. Clitellum 8-annulate. Male gonopore between annuli 3 and 5, female gonopore between annuli 6 and 7, external copulatory area absent (Fig. 43: 4). Annulation of complete somite and location of anus obscure. Coloration. All specimens unpigmented after al- cohol fixation, skin semitransparent. 2. Digestive system (Fig. 43: 5). Base of proboscis located immediately prior to ganglion 2 of ventral nerve cord. Esophageal diverticula located between ganglia 3 and 4. Four crop chambers small, diamond- shaped, without lateral processes. Posterior crop caeca absent. Four intestinal chambers spherical, without lateral processes. Rectal dilation present. 3. Reproductive system (Fig. 43: 6,7). Five pairs of large testisacs. Seminal reservoirs in form of several UTEVSKY, ANTARCTIC PISCICOLID LEECHES FIG. 41. Cryobdella antarctica, external and internal structure: 1, common view; 2, anterior sucker (dorsal and ventral view); 3, posterior sucker; 4, clitellum (ventral view); 5, somite (dorsal view); 6, digestive system; 7, reproductive system (dorsal view); 8, reproductive system (lateral view); 9, coelomic system (reconstruction). N BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 54/2007 FIG. 42. Cryobdella antarctica, dorsal and ventral view. loops berween ganglia 5 and 6. Initial parts of ejacu- latory ducts short, coiled in the form of several loops, indistinctly divided from wide terminal parts. Acces- sory glands absent. Common part small, bursa very short. Ovisacs of intermediate length extending to ganglion 8. Ovisacs coiled between ganglia 6 and 7. Conductive tissue absent. 4. Coelomic system (Fig. 43: 8). Dorsal, ventral and testicular lacunae present. Main connections absent: ventral lacuna forming a small lateral processes at ganglion region. Accessory connections unclosed: connecting dorsal and testicular lacunae. 5. Host data: collected from gill rakers of Mura- enolepis marmoratus and Muraenolepis microps. 6. Distribution (Fig. 39): Kerguelen coastal waters - Indian sector of the Antarctic region; S. Orkney Is. - Atlantic sector. Reference: Epshtein & Utevsky 1994. Cryobdella pallida A.Utevsky, 1997. 1. External characters (Figs. 45, 46). Small leeches. Up to 9 mm in length and 0.7 mm in width. Body 58 long, cylindrical, indistinctly divided into trachelo- some and urosome (Fig. 46). Anterior sucker large, distinctly divided from trachelosome, eccentrically attached, surface smooth, edge rough, mouth-pore prior to centre (Fig. 45: 2). Posterior sucker large, larger than anterior sucker, eccentrically attached, sur- face smooth (Fig. 45: 3). Annulation. Clitellum 5-annulate (may be an- nulation of 2nd degree). Male gonopore located on annulus 2, female gonopore on annulus 3. External copulatory area absent (Fig. 45: 4). Annulation of complete somite obscure, basically 3-annulate with annulation of 2nd-3rd degree (Fig. 45: 5). Surface of annuli smooth. Anus on annulus 2 from posterior sucker (Fig. 45: 3). Coloration. Skin semitransparent (all examined specimens unpigmented after alcohol fixation). 2. Digestive system (Fig. 45: 6). Proboscis ex- tending to Ist ganglion of ventral nerve cord. Eso- phageal diverticula located between ganglia 4 and 5. Five crop chambers well developed, diamond-shape without lateral processes. Posterior crop caeca non- UTEVSKY, ANTARCTIC PISCICOLID LEECHES FIG. 43. Cryobdella hadovi, external and internal structure: 1, common view; 2, anterior sucker (dorsal and ventral view); 3, posterior sucker; 4, clitellum (ventral view); 5, digestive system; 6, reproductive system (dorsal view); 7, reproductive system (lateral view); 8, coelomic system (reconstruction). 59 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 54/2007 FIG. 44. Cryobdella ladovi, ventral view. totally fused with 5 fenestrae, without lateral proces- ses, extending to posterior ganglion mass. Intestinal chambers weakly developed, without lateral processes. Rectal dilation weakly developed. 3. Reproductive system (Fig. 45: 7, 8). Five pairs of testisacs. Seminal reservoirs short and wide, ex- tending posterior to ganglion 6 of ventral nerve cord. Initial parts of ejaculatory ducts short and wide, sharply divided from roundish-wide terminal parts. Accessory glands absent. Common part small, bursa very small. Ovisacs long and voluminous, asymme- trical, extending posterior to ganglion 7. Conductive tissue absent. 4. Coelomic system (Fig. 45: 9). Dorsal, ventral, lateral, and testicular lacunae present. Main connec- tion nontotally close, ventral lacuna enlarges at gang- lion region, dorsal lacuna forms two long projections extended to dilation of ventral lacuna. Accessory connection absent. Testicular lacunae connected by lateral lacunae on each side. 60 5. Host data: all specimens collected from gills of Notothenia squamifrons. 6. Distribution (Fig. 39): Crozet I.- Indian sector of the Antarctic region. References: Utevsky 1997. “Subfamily Piscicolinae” Diagnosis. Leeches of varying sizes; body long or short, cylindrical, subcylindrical or flattened. Tuber- cles, papillae, eyes, eye-like spots, segmental ocelii, ocelli on posterior sucker present or absent. Digestive system of varying structure, posterior crop caeca of varying degrees of fusion or absent. Reproductive system of varying structure. Coelomic system possesses dorsal, ventral, lateral, testicular lacunae, main and accessory connections of varying degrees developed, one pair of externally visible subepidermal pulsatile vesicles formed by mo- dified lateral lacunae per urosomal somite. References: Sawyer 1986; Epshtein er al. 1994. Key to the Antarctic Genera of the “Subfamily Piscicolinae” la Body wide and flattened lb Body long, cylindrical or subcylindrical .... 3 2a Pulsatile vesicles well developed; Li=L»; surface of anterior sucker smooth, 1 pair of eyes, pos- terior sucker without ocelli Genus Trachelobadellina (p. 60) Pulsatile vesicles small; Lı<; anterior sucker with papillae and 2 groups of ocelli; posterior sucker with ocelli; 2 groups of ocelli on first oO annulus of trachelosome Genus Trulliobdella (p. 63) Posterior crop caeca absent Genus Galatheabdella (p. 69) Posterior crop caeca present Anterior and posterior suckers small; facing directly posteriorly; prepuce present Genus Trachelobdella (p.72 Anterior and posterior suckers large, facing ventrally; prepuce absent Genus Nototheniobdella (p. 75) Genus Trachelobdellina Moore, 1957 Type species: Trachelobdellina glabra Moore, 1957 Not large leeches. Trachelosome cylindrical, up to one half of the entire body length; urosome wider and UTEVSKY, ANTARCTIC PISCICOLID LEECHES - — BUD Ade bi POST Ann ( MTT TTT rc raya a Tenner, : ray Wah 9) 5 3 u Sera eee aT UT FIG. 45. Cryobdella pallida, external and internal structure: 1, common view; 2, anterior sucker (dorsal and ventral view); 3, posterior sucker; 4, clitellum (ventral view); 5, somite (dorsal view); 6, digestive system; 7, reproductive system (dorsal view); 8, reproductive system (lateral view); 9, coelomic system (reconstruction). 61 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 54/2007 more flattened than trachelosome. There are ocelli on the dorsum and venter of the body. Complete somite 6-annulate with pair of pulsate vesicles. Segmental pigmentation absent. Anterior sucker: small, cardiform, with pair of eyes. Posterior sucker: small, without ocelli. Digestive system. Esophageal diverticula present, posterior crop caeca nontotally fused, number of lateral processes of intestine reduced. Reproductive system. Five pairs of testisacs, bur- sa short, accessory glands absent. Coelomic system. Dorsal, ventral, pair of lateral and testicular lacunae present. Main and accessory connections present. Host data: hosts poorly known, one specimen collected from Glyptonotus antarcticus (Isopoda). Distribution (Fig. 47): Indian and Antlantic parts of the Antarctic region. Monotypic genus. Reference: Moore 1957. FIG. 47. Distribution of the Trachelobdellina glabra. 62 FIG. 46. Cryobdella pallida, lateral view. FIG. 48. Trachelobdellina glabra, common view (re- produced from Moore 1957). Trachelobdellina glabra Moore, 1957* 1. External characters (Fig. 48). Not large leeches. Up to 30 mm in length and 7 mm in width. Body distinctly divided into long, narrow trachelosome and wide, subcylindrical urosome. They are approximately equal in length. Body smooth, with 12 pairs of large pulsatile vesicles on urosome, five pairs of ventral and dorsal ocelli: three pairs on clitellar segments, one pair on last trachelosomal segment and one pair on first urosomal segment. Anterior sucker small, cardiform, with a pair of anterior rounded lobes separated by a median cleft, distinctly divided from trachelosome, eccentrically attached. One pair of eyes. Mouth-pore centrally located. Posterior sucker small, but larger than anterior one. Annulation. Clitellum 9-annulate; somites of tra- chelosome 3-annulate, without additional annulation. Male gonopore between annuli 3 and 4, female gono- pore between annuli 8 and 9 (according to Moore 1957, male gonopore between annulus A3 of 10th somite and 11 somite, female gonopore between an- nuli A» and Az of 12th somite). Complete somite 6(8)-annulate with additional annulation. UTEVSKY, ANTARCTIC PISCICOLID LEECHES 2. Digestive system. Proboscis extend to ganglion 6 of ventral nerve cord, very coiled at ganglion 3. Eso- phageal diverticula based between ganglia 5 and 6, ex- tending cephalad between ganglia 4 and 5. Crop tubular, without distinct chambers and lateral pro- cesses. Posterior crop caeca nontotally fused, with fenestrae. Intestine tubular, with 1 pair of lateral pro- cesses at ganglion 13. 3. Reproductive system. Five pair of testisacs. Ter- minal parts of male reproductive system typical (?). Bursa short. Accessory glands absent. 4. Coelomic system. Dorsal, ventral, lateral and testicular lacunae present. Lateral lacunae paired (thin- and thick-walled) on each side of the body. Main connections closed, dorsal and ventral lacunae connected through thin-walled ventro-lateral lacu- nae in ganglion region. Accessory connection closed, dorsal and ventral lacunae connected through testi- cular lacunae. Thin-walled ventro-lateral lacunae con- nected with testicular lacunae. Anterior parts of pulsa- tile vesicles connected with thin-walled ventro-late- ral lacunae at ganglion region. Posterior part of pulsa- tile vesicles connected with thick-walled lateral lacu- nae. 5. Host data: 1 specimen collected from Glypto- notus antarcticus (Isopoda). 6. Distribution (Fig. 47): Mawson Coast - Indian sector of the Antarctic region, S. Shetland Is., Arthur Harbor (Palmer Station) - Atlantic sector. References: Moore 1957, Meyer & Burreson 1990. Genus 7rulliobdella Brinkmann, 1948 (Syn.: Cryobdellina Brinkmann, 1948; Antarctobdella Dollfus, 1964; Ophthalmobdella Szidat, 1965) Type species: Trulliobdella capitis Brinkmann, 1948 Medium-sized leeches. Body variable in form, di- stinctly divided into trachelosome and urosome. Ira- chelosome with 2 groups of 3-5 ocelli in each on annuli 1-3. Complete somite 3-annulate with addi- tional annulation, pulsatile vesicles on annulus A3. Segmental ocelli in young individuals specimens on dorsum and venter of annulus Az or absent. Seg- mental pigmentation absent. Anterior sucker: small, eccentrically attached, with 4-6 tentacles and two 2 groups of 5 ocelli in each. Posterior sucker: small, surface smooth, with 7—15 ocelli. Digestive system. Large esophageal diverticula present. Crop chambers with long and voluminous double lateral processes. Posterior crop caeca nonto- tally fused, with 4 large fenestrae and lateral proces- ses similar to processes of crop chambers. Intestine 63 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 54/2007 with 4 small chambers and long lateral processes. Rec- tal dilation present. Reproductive system. Five pairs of testisacs. Se- minal reservoirs short. Initial parts of ejaculatory ducts thin. Posterior parts wide. Common part voluminous. Bursa short. Accessory glands absent. Oviducts and conductive tissue cords connects with voluminous seminal receptacle, which form internal copulatory area. External copulatory area around gonopores present. Coelomic system. Dorsal, ventral, lateral and testicular lacunae present. Dorsal lacuna divided into two coelomic channels. Main and accessory connec- tions unclosed. Lateral lacunae form typical for Pis- cicolinae pulsatile vesicles. Host data: fam. Nototheniidae, Bathydraco- nidae, Chaenichthyidae (ord. Perciformes). Distribution (Fig. 49): Atlantic, Pacific and Indian parts of the Antarctic region. Congenitors: 7. bacilliformis (Brinkmann, 1948) References: Brinkmann 1948, Sawyer 1986, Mey- er & Burreson 1990, Utevsky 2000. Remark: Meyer & Burreson (1990) synonymized species of the genera Cryobdellina, Trulliobdella, Ant- arctobdella, Ophthalmobdella with Notobdella noto- theniae Benham, 1909. We cannot hold this syno- nymy as the description of latter species is based on some external characters and lacks any illustrations. Benham did not record tentacles on the anterior sucker. The shape, arrangement and the number of the eyes do not agree with the characters of the other “tentacled” leeches. Key to the Species of the Genus Trulliobdella la Body yellowish gray, posterior sucker with 7 ocelli I! capitis (p. 65) 1b Body without pigment, semitransparent; pos- terior sucker with 15 ocelli T! bacilliformis (p. 66) FIG. 49. Distribution of the gen. Trulliobdella: 1 - T! capitis; 2 - 7! bacilliformis. 64 UTEVSKY, ANTARCTIC PISCICOLID LEECHES Trulliobdella capitis Brinkmann, 1948 able from leaf to cylindrical) sharply narrowing to 1. External characters (Figs. 50, 51). Medium-sized suckers, distinctly divided into subcylindrical trache- leeches. Up to 60 mm in length and 12 mm in width. losome and lancet-shaped (more flattened) urosome. Body short and wide, leaf-shaped (body shape vari- Ten to twelve pairs of very small pulsatile vesicles, seg- FIG. 50. Trulliobdella capitis, external and internal structure: 1, common view; 2, anterior sucker (dorsal and ventral view); 3, posterior sucker; 4, clitellum (ventral view); 5, somite (dorsal view); 6, digestive system; 7, reproductive system (dorsal view); 8, reproductive system (lateral view); 9, coelomic system (reconstruction). 65 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 54/2007 mental ocelli absent on urosome (Fig. 50: 1). Anterior sucker small, distinctly divided from trachelosome, eccentrically attached; with 4 tentacles and 2 fields with 3-5 ocelli in each near the base of sucker (Fig. 50: 2). Ocelli present in 2 fields of 3 first annuli of trachelosome, each field consisting of 5 ocelli; mouth- pore prior to centre. Posterior sucker small, but larger than anterior one, nearly centrally attached; surface smooth, with a maximum 7 ocelli on its marginal dorsal posterior part (Fig. 50: 3). Annulation. Clitellum 4-annulate (may be addi- tional annulation of 2-3 degree). Male gonopore on annulus 1, female gonopore on annulus 4. Gonopores situated on swelling of ventral surface of clitellum. This swelling is the external copulatory area (Fig. 50: 4). Complete somite 3-annulate (may be additional annulation of 2nd-3rd degree), surface smooth, with- out segmental ocelli (Fig. 50: 5). Annulus Az consists of a pair of pulsatile vesicles. Anus on 2nd annulus from posterior sucker (Fig. 50: 3). Coloration. Body and suckers yellowish gray, ocelli may be depigmented after fixation. 2. Digestive system (Fig. 50: 6). Base of proboscis between ganglia 3 and 4 of ventral nerve cord. Eso- phageal diverticula between ganglia 4 and 5, long and voluminous. Five small crop chambers with double lateral processes which may be bifurcated in turn. Pos- terior crop caeca nontotally fused with 4 large fene- strae and 5 pairs of lateral processes similar to crop chamber processes, do not extend to posterior gang- lion mass. Four intestinal chambers small with long processes laterally directed, Sth chamber in form of “folded organ”. Rectal dilation small. 3. Reproductive system (Fig. 50: 7, 8). Five pairs of testisacs. Seminal reservoirs short, in form of small loops between ganglia 6 and 7 of ventral nerve cord. Initial parts of ejaculatory ducts short and thin, sharply divided from spherical wide terminal parts. Accessory glands absent. Common part extensive, bursa short. Ovisacs long, coiled between ganglia 7 and 8, flowing extensive seminal receptacle which formed internal copulatory area (depression of ven- tral body wall). Conductive tissue in form of short cords between ganglia 6 and 7 which connect semi- nal receptacle and ovisacs. 4. Coelomic system (Fig. 50: 9). Dorsal, ventral, lateral and testicular lacunae present. Dorsal lacuna consisting of two channels. Main connections un- closed, projections of ventral lacuna extending to dorsal lacuna in ganglion region. Accessory connec- 66 FIG. 51. Trulliobdella capitis, lateral and ventral view. tions unclosed, connecting ventral and testicular lacunae. Lateral lacunae connected with main connec- tions through pulsatile vesicles in ganglion regions. 5. Host data: Notothenia rossi, Parachaenichthys georgianus, Neopagetopsis ionah, Pseudochaenichthys georgianus, Chaenichthys rhinoceratus, Chaenocephalus bouvetensis, Cryodraco antarcticus, Champsocephalus gunnari, Chionodraco kathleenae, C. hamatus, C. rastro- spinosus, Chaenodraco wilsoni. 6. Distribution (Fig. 49): Bouvet I., S. Shetland Is., S. Orkney Is., Weddell Sea, Argentine Is. - Atlantic sector of Antarctic region; Wilkes Land - Indian sec- tor; Ross Sea - Pacific sector. References: Brinkmann 1948, Meyer & Burreson 1983, Meyer & Burreson 1990, Sawyer 1986. Trulliobdella bacilliformis (Brinkmann, 1948) (Syn.: Cryobdellina bacilliformis Brinkmann, 1948; Antarctobdella tcherniai Dollfus, 1964; Antarctobdella crozetensis Sawyer, 1972; Ophthalmobdella bellisioi Szidat, 1965) 1. External characters (Figs. 52, 53). Medium-sized leeches. Up to 45 mm in length and 9 mm in width. Body short and wide, leaf-shaped (body shape vari- able) sharply narrowing to suckers, distinctly divided on subcylindrical trachelosome and lancet-shaped (more flattened) urosome (Fig. 52: 1). Urosome pos- sessing 10-12 pairs of very small pulsatile vesicles and segmental ocelli on venter and dorsum. Anterior sucker small, distinctly divided from trachelosome, eccentrically attached; with 6 tentacles and 2 fields UTEVSKY, ANTARCTIC PISCICOLID LEECHES wog: Ueda FIG. 52. Trulliobdella bacilliformis, external and internal structure: 1, common view; 2, anterior sucker (dorsal and ventral view); 3, posterior sucker; 4, clitellum (ventral view); 5, somite (dorsal view); 6, digestive system; 7, reproductive system (dorsal view); 8, reproductive system (lateral view); 9, coelomic system (reconstruction). 67 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 54/2007 FIG. 53. Trulliobdella bacilliformis and other “tentacled” leeches: A, Trullibdella bacilliformi (Brinkmann, 1948); B, Antarctobdella tcherniai Dollfus, 1964; C, Antarctobdella crozetensis Sawyer, 1972 (reproduced from Sawyer 1972); D, Cryobdellina bacilliformis Brinkmann, 1948 (reproduced from Brinkmann 1948); E, Ophthalmobdella bellisioi Szidat, 1965 (reproduced from Szidat 1965). near the base with 3-5 ocelli per each (Fig. 52: 2). Ocelli present in 2 fields of 2 first annuli of trache- losome, each field consisting of 5 ocelli; mouth-pore prior to centre. Posterior sucker small, but larger than anterior one, eccentrically attached; surface smooth, with a maximum 15 ocelli on its marginal dorsal posterior part (Fig. 52: 3). Annulation. Clitellum 5-annulate (may be addi- tional annulation of 2nd-3rd degree). Male gonopore between annuli 2 and 3, female gonopore between annuli 4 and 5. Gonopores situated on swelling of ventral surface of clitellum which evidently formed external copulatory area (Fig. 52: 4). Complete somite 3-annulate (may be additional annulation of 2nd-3rd degree). Annulus Ap consists a pair of pulsatile vesicles and two pair of segmental ocelli on dorsum and venter (Fig. 52: 5). Anus on 2nd annulus from posterior sucker (Fig. 52: 3). Coloration. Body without pigments, ocelli may be absent. 2. Digestive system (Fig. 52: 6). Base of proboscis between ganglia 2 and 3 of ventral nerve cord. Eso- phageal diverticula based after ganglion 3 and ex- tending to ganglion 5, long and voluminous. Six small crop chambers with double lateral processes which may be bifurcate also. Posterior crop caeca nontotally fused with 4 large fenestrae, possessing 5 pairs of lateral processes similar to processes of crop chambers, do not extend to posterior ganglion mass. Four in- testinal chambers small with long lateral “V”-shaped processes at their base. Fifth chamber in form of “folded organ”. Rectal dilation small. 68 3. Reproductive system (Fig. 52: 7, 8). Five pairs of testisacs. Seminal reservoirs (short) in form of smail loops near ganglion 6 of ventral nerve cord. Initial part of ejaculatory ducts short and thin, sharply divided from trapezium-shaped wide terminal parts. Acces- sory glands absent. Common part extensive, bursa short, opening in pocket (depression of ventral body wall). Ovisacs long, coiled at ganglion 7, flowing into extensive seminal receptacle which forms inter- nal copulatory area (depression of ventral body wall) also. Conductive tissue in form of very short cords near ganglion 6 which connects seminal receptacle and ovisacs. 4. Coelomic system (Fig. 52: 9). Dorsal, ventral, lateral and testicular lacunae present. Dorsal lacuna consisting of two channels. Main connections un- closed, projections of ventral lacuna extending to dorsal lacuna in ganglion region. Accessory connec- tion unclosed, connecting ventral and testicular la- cunae. Lateral lacunae connected with main connec- tions through pulsatile vesicles in ganglion regions. 5. Host data: Notothenia coriiceps, N. neglecta, No- totheniops larsent; Parachaenichthys georgianus;, Pseu- dochaenichthys georgianus, Chaenichthys rhinoceratus, Chaenocephalus aceratus, C. bouvetensis, Champsoce- phalus gunnari, C. rastrospinosus. 6. Distribution (Fig. 49): Bouvet I., S. Shetland Is., S. Orkney Is., S. Georgia I.- Atlantic sector of the Antarctic region; Prince Edward Is., Marion I., Cro- zet Is., Kerguelen Is.- Indian sector. References: Brinkmann 1948, Benham 1909, Dollfus 1964, Sawyer 1972, Szidat 1965. Genus Galatheabdella Richardson & Meyer, 1973 Type species: Galatheabdella bruuni Richardson & Meyer, 1973 Large leeches. Body long, cylindrical, distinctly di- vided into trachelosome and urosome; tapering to- wards suckers; with 11 pairs of pulsatile vesicles, with- out ornamentation and segmental ocelli. Complete somite 14-annulate. Anterior sucker: medium-sized, distinctly divided from trachelosome, eccentrically attached, surface smooth, eyes or eye-like spots absent, mouth-pore prior to centre. Posterior sucker: large, distinctly divided from urosome, eccentrically attached, surface smooth, with- out ocelli. Digestive system. Esophageal diverticula absent. Crop tubular, without distinct chambers and lateral processes. Posterior crop caeca absent. Intestine with chambers. Rectal dilation present. ’S00 FIG. 54. Distribution of the Galatheabdella bruuni. UTEVSKY, ANTARCTIC PISCICOLID LEECHES Reproductive system. Six pairs of testisacs. Semi- nal reservoirs long, ejaculatory ducts long, common part small, bursa long. Accessory glands present. Con- ductive tissue mass present, copulatory area absent. Coelomic system. Structure unknown. Host data: Coryphaenoides sp, Bassozetus sp. Distribution (Fig. 54): Tasman Sea; Atlantic sec- tor of the Antarctic region. Monotypic genus. Reference: Richardson & Meyer 1973. Galatheabdella bruuni Richardson & Meyer, 1973* 1. External characters (Figs. 55, 56). Large leeches. Up to 90 mm in length, 5 mm in width. Body long, cylindrical, distinctly divided into trachelosome and urosome; surface smooth, semitransparent, without ornamentation and ocelli, 11 pairs of pulsatile vesicles on urosome (Fig. 56). Trachelosome tapering cephalad. Urosome tapering caudad. Anterior sucker medium- 69 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 54/2007 sized, distinctly divided from trachelosome, eccentri- urosome, eccentrically attached, surface smooth, ocelli cally attached, eyes or eye-like spots absent, mouth- _ absent (Fig. 56). pore prior to centre (Fig. 55: 2). Posterior sucker large, Annulation. Clitellum 7-annulate, annulation larger than anterior sucker, distinctly divided from — weakly defined. Male gonopore located between an- FIG. 55. Galatheabdella bruuni, external and internal structure: 1, common view; 2, anterior sucker with tra- chelosome and clitellum; 3, somite; 4, digestive system; 5, reproductive system (dorsal view); 6, reproductive system (lateral view) (reproduced and modified from Richardson & Meyer 1973). 70 FIG. 56. Galatheabdella bruuni, common view (re- produced from Richardson & Meyer 1973). nuli 1 and 2, female gonopore between 4 and 5. Ex- ternal copulatory area absent (Fig. 55: 2). Complete somite 14-annulate (with additional annulation), an- nulation weakly defined (Fig. 55: 3). Length of annuli variable. Pulsatile vesicles situated on annuli 4—6 and crossed by two interannular grooves, first pair of pulsa- tile vesicles on first annuli of urosome. Anus on an- nulus 2 from posterior sucker. Coloration. No dark pigment and pattern, tes- tisacs and ganglia of ventral nerve cord not visible, UTEVSKY, ANTARCTIC PISCICOLID LEECHES pinkish individual large cells visible trough the body wall, other body wall accepting this area dusky white. 2. Digestive system (Fig. 55: 4). Base of proboscis located before ganglion 3 of ventral nerve cord. Eso- phageal diverticula absent. Crop tubular, without distinct chambers and lateral processes. Posterior crop caeca absent. Four intestinal chambers small, without lateral processes. Rectal dilation present, tubular, re- ducing in diameter before ending at the anus. 3. Reproductive system (Fig. 55: 5, 6). Six pairs of elongate ovoid testisacs covered by clitellar gland cells and difficult to detect. Vasa deferentia joining smoothly seminal reservoir at ganglion 6 of ventral nerve cord. Seminal reservoir extending caudad to ganglion 7 to coil severely (may be asymmetrical), continue cephalad to ganglion 5, joining initial por- tions of curved, loop-shaped in vertical plane ejacu- latory ducts. Ejaculatory ducts sharply divided from FIG. 57. Distribution of the Trachelobdella bathyrajae. 7a BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 54/2007 wide terminal part. Common part small, obtusely co- nical; bursa long, evertable by muscular strands which connected with lateral papillae on each side of lower region of common part of mail reproductive system. Accessory glandular mass covering posterior portions of loops of ejaculatory ducts and its common part. Ovisacs extending caudad to ganglion 8 of ventral nerve cord, consisting of some lobes, asymmetrical. Conductive tissue in form of vertical, obtusely cy- lindrical fibrous mass between gonopores, which is connected by two fibrous cords with bursa and by two similar cords with ovisacs which extend caudad to each lobe. 4. Coelomic system. Structure unknown. 5. Host data: collected from Coryphaenoides sp., Bassozetus sp. in Tasman Sea; in Antarctic no host data. 6. Distribution (Fig. 54): Bransfield Str. - Atlan- tic sector of the Antarctic region; Tasman Sea - Paci- fic Ocean. References: Richardson & Meyer 1973, Meyer & Burreson 1990. Genus Trachelobdella Diesing, 1850 Type species: Trachelobdella muelleri Diesing, 1850 Not large or small leeches. Body short (may be stretching), distinctly divided into cylindrical trache- losome and subcylindrical or flattened urosome with 12-13 pairs of pulsatile vesicles. Complete somite 3- annulate, with additional annulation. Anterior sucker: small, eyes present or absent, mouth-pore centrally located. Posterior sucker: small, ocelli absent. Digestive system. Esophageal diverticula absent or present. Crop chambers various in shape. Posterior crop caeca nontotally fused, with fenestrae. Lateral processes of intestine chambers weakly developed or absent. Rectal dilation present. Reproductive system. Five to six pairs of testisacs. Seminal reservoirs very coiled. Initial parts of ejacu- latory ducts short. Terminal parts wide, with accessory glands. Bursa long. Ovisacs short, with long muscular vagina. Conductive tissue and copulatory area absent. Coelomic system. Dorsal, ventral, lateral and testi- cular lacunae present. Main and accessory connections in different structure. Lateral lacunae forming pulsa- tile vesicles typical for Piscicolinae. Distribution (Fig. 57): Atlantic, Pacific, Indian Ocean, Antarctic Seas. 22 FIG. 58. Trachelobdella bathyrajae, external and in- ternal structure: 1, common view; 2, anterior sucker; 3, posterior sucker; 4, complete somite; 5, coelomic system (reconstruction) (© 1990 American Geo- physical Union. Reproduced /modified by permission of American Geophysical Union and E.M. Burreson, Virginia Institute of Marine Science. M.C. Meyer and E.M. Burreson Some leeches (Hirudinea: Piscicolidae) of the Southern oceans. Biology of the Antarctic Seas XXI, Antarctic Research Series, V. 52, p 231). Congenitors: 7. lubrica (Grube, 1840) - equatorial world-wide T. maculata Moore, 1898 I! rugosa Moore, 1898 T. vividus Moore, 1898 1. australis Blanchard, 1900 - South America 7. luederitzi Augener, 1936 - south-west Africa T! leptocephali Ingram, 1957 — Tasman Sea 7. oregonensis Burreson, 1976 — North Pacific I. bathyrajae Meyer & Burreson, 1990 — Antarctic Seas References: Epshtein 1968, Epshtein er al. (1994), Sawyer 1986, Meyer & Burreson 1990. Trachelobdella bathyrajae Meyer & Burreson, 1990* 1. External characters (Fig. 58). Small leeches. Up to 13 mm in length and 2 mm in width. Body long, sub- cylindrical, tapering to suckers, distinctly divided in- to trachelosome and urosome by prepuce, without tubercles. Urosome with 11 pairs of pulsatile vesicles (Fig. 58: 1). Anterior sucker small, distinctly divided from trachelosome, eccentrically attached, without eyes, eye-like spots and tubercles, mouth-pore in centre (Fig. 58: 2). Posterior sucker small, but large than anterior sucker, centrally attached, indistinctly divided from urosome, cylindrical, facing directly posteriorly; without ocelli and tubercles (Fig. 58: 3). Annulation. Male and female gonopores separated by 3 annuli. Complete somite 6-13 annulate (addi- tional annulation), pulsatile vesicles on annulus B;, segmental ocelli absent (Fig. 58: 4). Coloration. 5 yellowish bands on trachelosome and 13 band of same color on urosome, annuli By and Bs unpigmented. Suckers unpigmented. 2. Digestive system. Base of proboscis located near ganglion 3 of ventral nerve cord. Esophageal diverti- cula based between ganglia 3 and 4. Crop enlarged FIG. 59. Distribution of the Nototheniobdella sawyeri. UTEVSKY, ANTARCTIC PISCICOLID LEECHES between testisacs. Posterior crop caeca nontotally fused with 4 fenestrae between ganglia 13 and 16, without lateral processes. Five intestinal lateral pro- cesses directed anteriorly, decreasing in size from Ist to 5th pair. Rectum tubular, tapering to anus. 3. Reproductive system. Six pairs of testisacs. Se- minal vesicles small, tubular, located near ganglion 7. Ejaculatory ducts long, loop-like, extending to gang- lion 4. Terminal and common parts tubular, bursa long. Accessory glands present, covering terminal parts of male reproductive system. Ovisacs short, tapering to gonopore. Copulatory area and distinct conduc- tive tissue mass absent. 4. Coelomic system (Fig. 58: 5). Dorsal, ventral lateral and testicular lacunae present. Main connec- tions unclosed, projections of ventral lacuna connec- ted with pulsatile vesicles. Accessory connections un- closed, connecting dorsal and testicular lacunae. Lateral lacunae connected with main connection through pulsatile vesicles in ganglion region. 73 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 54/2007 FIG. 60. Nototheniobdella sawyeri, external and internal structure: 1, common view; 2, anterior sucker (dorsal and ventral view); 3, posterior sucker; 4, clitellum (ventral view); 5, somite (dorsal view); 6, digestive system; 7, reproductive system (dorsal view); 8, reproductive system (lateral view); 9, coelomic system (reconstruction). 74 FIG. 61. Nototheniobdella sawyeri, lateral view. 5. Host data: collected from Bathyraja maccaini. 6. Distribution (Fig. 57): S. Shetland Is. - Atlan- tic sector of the Antarctic region. Reference: Meyer & Burreson 1990. Genus Nototheniobdella A. Utevsky, 1993 Type species: Nototheniobdella sawyeri A. Utevsky, 1993 Medium-sized leeches. Body long, cylindrical, with- out papillae, tubercles, segmental ocelli and segmen- tal pigment bands. Complete somite with well defined annulation. Anterior sucker: large, strictly divided from tra- chelosome, eccentrically attached, with one pair of eyes, mouth-pore centrally located. Posterior sucker: large, strictly divided from uro- some, centrally attached, without ocelli. Digestive system. Esophageal diverticula absent. Crop chambers without lateral processes. Posterior crop caeca nontotally fused with 5 fenestrae, without lateral processes. Reproductive system. Five to six pairs of testisacs. Accessory glands present, bursa short. Ovisacs long. Conductive tissue, internal and external copulatory area absent. UTEVSKY, ANTARCTIC PISCICOLID LEECH] Coelomic system. Structure of main and accessory connections variable in different somites. Host data: fam. Nototheniidae, Bathydraconidae, Chaenichthyidae (ord. Perciformes). Distribution (Fig. 59): Atlantic, Indian and Paci- fic parts of the Antarctic region. Monotypic genus, 1 species in Antarctic region. References: Utevsky 1993, Utevsky 2000. Nototheniobdella sawyeri A.Utevsky, 1993 1. External characters (Figs. 60, 61). Medium- sized leeches. Up to 60 mm in length and 5 mm in width. Body long, cylindrical, sharply narrowing to anterior sucker, distinctly divided into trachelosome and urosome. Trachelosome narrower and thinner than urosome, clitellum somewhat wider than prec- litellum. Urosome with 12-13 pairs of pulsatile vesicles (Fig. 61). Anterior sucker small, distinctly divided from trachelosome, eccentrically attached, with one pair of eyes, surface smooth (may be trans- verse furrows and rough edge) mouth-pore centrally located (Fig. 60: 2). Posterior sucker large, larger than anterior sucker, centrally attached, surface smooth (Fis. 6023). Annulation. Clitellum 11-annulate (may be ad- ditional annulation). Male gonopore located on an- nulus 5, female gonopore between annuli 9 and 10, external copulatory area absent (Fig. 60: 4). Complete somite 6-annulate with additional annulation of 2nd- 4th degree (Fig. 60: 5). Annulus B; bearing a pair of pulsatile vesicles. Anus on 2nd annulus from poste- rior sucker (Fig. 60: 3). Coloration. Segmental brownish bands on trahe- losome and urosome present. 10 bands on trahelo- some and 12-13 ones (which include the annuli with pulsatile vesicles) on urosome. Body and suckers un- pigmented after alcohol or formalin fixation. 2. Digestive system (Fig. 60: 6). Base of proboscis located before ganglion 3 of ventral nerve cord. Eso- phageal diverticula absent. Five diamond-shaped crop chambers without lateral processes. Posterior crop caeca nontotally fused, with 5 fenestrae which are filled by connective tissue. Five intestinal chambers well developed, without lateral processes. Rectal di- lation present. 3. Reproductive system (Fig. 60: 7, 8). Six pairs of testisacs, testisacs of first pair may be reduced. Late- rally located vasa deferentia joining smoothly semi- nal reservoir at ganglion 6 of ventral nerve cord. Me- dial seminal reservoir extending caudad to ganglion 7 to coil severely, continueing cephalad, joining greatly NN BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 54/2007 coiled initial portions of curved, loop-shaped ejacu- latory ducts. Ejaculatory ducts sharply divided from spherical wide terminal parts. Common part small, bursa short. Accessory glandular mass covering pos- terior parts of terminal, common parts and bursa. Ovisacs long, forming numerous loops and extending posterior to ganglion 8. Conductive tissue absent. 4. Coelomic system (Fig. 60: 9). Dorsal, ventral, la- teral and testicular lacunae present. Main connections at ganglia 8 and 9 completely closed: connecting dor- sal and ventral lacunae; other main connections un- closed: long projections of ventral lacuna extending to dorsal lacuna in ganglion region. Accessory con- nections of testisacs Ist and 2nd pairs connecting dor- 76 sal, ventral and testicular lacunae. Lateral lacunae connecting testicular lacunae and forming pulsatile ve- sicles connected with projections of ventral lacunae in ganglion region. 5. Host data: collected from body, mouth and gill cavity of Neopagetopsis ionah, Cryodraco antarcticus, Chaenodraco wilsoni, Chionodraco kathleenae, C. ha- matus, Parachaenichthys georgianus, Harpagiferidae (?). 6. Distribution (Fig. 59): Palmer Coast, S. Shet- land Is., Filchner Ice Shelf (Weddell Sea) - Atlantic sector of the Antarctic region; Clarie Coast - Indian sector; Scott Coast, Franklin I. (Ross Sea) - Pacific sector. Reference: Utevsky 1993. 7. REFERENCES Badham, C. 1916. On an ichthyobdellid parasitic on the Australian Sand whiting (Sillago ciliata). Quart. Journ. Micr. Sci. 62: 1-41. Bakay, Yu.I., Kuzmin, S.A., & S.Y. Utevsky. 1998. Ecolo- gical and parasitological investigations on the Barents Sea Red Crab Paralithodes camtshatica (the first results). In: International Council for the Exploration of the Sea (Session on Stock Components in Managment, AA 4, 195-202. Benham, W.B. 1909. Preliminary report on two Hirudinea from the subantarctic islands of New Zealand. Pp. 372-376 in Chilton, C. (ed.). The Subantarctic Islands of New Zeland. Philosophical Institute of Canterbury, Wellington. Brinkmann, A. 1948. Some new and remarkable leeches from the Antarctic seas. Sci. Results Norw. Antarct. Ex- ped. 1927-1928. 29: 1-12. Dolifus, R. 1964/1965. Sangsue tentaculifere de la peau d’un teleosteen du genre Chaenichthys J. Richardson, 1844. Bull. Mus. D’Hist. Naturelle 2-e Serie, 36: 831-843. Epshtein, V.M. 1967. Regularities of the geographical dis- tribution of the marine fish leeches (Hirudinea, Pisci- colidae). Zool. Zhurn. XLVI: 680-691 (in Russian). Epshtein, V.M. 1968. Zoological analysis of the fish leeches in the Antarctic and a revision of the genus Trachelob- della Diesing, 1850. Pp. 137-138 in Fifth All-Union Conference of the Diseases and Parasites of Fishes and Marine Invertebrates. Leningrad (in Russian). Epshtein, V.M. 1970a. Bipolar distribution of marine fish leeches (Hirudinea, Piscicolidae). Pp. 143-146 in Vo- lyanitsky, V.A. (ed.). First All-Union Symposium Di- seases and Parasites of Marine Animals. Sevastopol, Kiev (in Russian). Epshtein, V.M. 1970b. Fish leeches (Hirudinea, Piscicoli- dae) in the Antarctic seas from the collection of the Zoological Institute of the Academy of Sciences of the USSR. Pp. 146-149 im Volyanitsky, V.A. (ed.). First All-Union Symposium Diseases and Parasites of Marine Animals. Sevastopol, Kiev (in Russian). Epshtein, V.M. 1972. Systematic status of the Antarctic leech Pontobdella rugosa Moore (Piscicolidae). Zool. Zhurn. LI: 1142-1146 (in Russian). Epshtein, V.M. 1974. New data on tropical origin elements in Antarctic Piscicolid Leech fauna. Pp. 301-305 in Six All-Union Symposium Diseases and Parasites of Mari- ne Anımals. Moscow (in Russian). Epshtein, V.M., & S.V. Okun’. 1991. On the structure, systematic status and distribution of the antarctic leech Pontobdella biannulata Moore, 1957. Informatsionnyi bjulleten’ Sovetskoi Antarcticheskoi Ekspedicii 116: 99-103 (in Russian). UTEVSKY, ANTARCTIC PISCICOLID LEECHES Epshtein, V.M., & A.Y. Utevsky. 1994. A new marine leech Cryobdella hadovi sp.n. from Antarctic Seas. Zoosyste- matica Rossica 3: 86-88. Epshtein, V.M., Utevsky, A.Y., & S.Yu. Utevsky. 1994. The system of fish leeches. Genus 5: 401-409. Harding, W.A. 1922. Hirudinea. Nat. Hist. Rep. Br. Ant- arct. Terra Nova Exped. 1910. Zool. X: 257-261. Hickman, V.V. 1947. Pontobdella tasmanica nom.nov. (Hi- rudinea). Pap. Proc. R. Soc. Tasmania 27. Janssen, H.H. 1993. Morphology, egg cocoons, and trans- mission paths of the Antarctic leech Glyptonotobdella antarctica Sawyer & White, 1969 (Hirudinea: Rhyncho- bdelliformes: Piscicolidae). Polar Biol. 13: 347-354. Khan, R.A., & A.J. Paul. 1995. Life cycle studies on arcto- boreal leeches (Hirudinea). J. Helminthol. Soc. Wash. 62: 105-110. Llewellyn, L.C. 1966. Pontobdellinae (Hirudinea: Piscico- lidae) in the British Museum (Natural History) with a revue of the Subfamily. Bull. of the British Museum (Natural History). Zoology 14: 341-439. Meyer, M.C., & A.A. Barden. 1955. Leeches symbiotic on Arthtropoda, especially Decapoda Crustacea. Wassmann Journal of Biology 13: 297-311. Meyer, M.C., & E.M. Burreson. 1983. Redescription of the piscicolid leech Trulliobdella capitis Brinkmann. Proc. Helminthol. Soc. Wash. 50: 138-142. Meyer, M.C., & E.M. Burreson. 1990. Some leeches (Hi- rudinea: Piscicolidae) of the southern oceans. Biology of the Antarctic Seas. Antarctic research series. 52: 219-226. Moore, J.P. 1938. Leeches (Hirudinea). Australasian Ant- arctic Expedition 1911-1914. Sci. Rep.(C) 10: 5-16. Moore, J.P. 1957. Hirudinea. B.A.N.Z. Antarctic Research Expedition. Rep. Ser. B (Zoology and Botany) 6: 102-105. Richardson, L.R., & M.C. Meyer. 1973. Deep-sea fish leeches (Rhynchobdellae: Piscicolidae). Galathea report: Scientific Results of the Danich Deep-Sea Expedition Round the World 1950-52. Pp. 113-126. Sawyer, R.T. 1972. A new species of “tentacled” marine fish leech parasitic on Notothenia from subantarctic Marion and Crozet Islands. Hydrobiologia 40: 345-354. Sawyer, R.T. 1976. Notes on two marine leeches (Anneli- da, Hirudinea) from Subantarctic Marion Islands in- cluding a new record. Hydrobiologia 48: 267-268 Sawyer, R.T. 1986. Leech biology and behaviour. Feeding biology, ecology, and systematics. Oxford. Sawyer, R.T., & M.G. White. 1969. A new genus and species of marine leech, Glyptonotobdella antarctica, from the antarctic isopod. Brit. Antarct. Surv. Bull. 22: 1-14. 1. BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 54/2007 Szidat, L. 1965. Los parasitos de Notothenia neglecta Nybe- lin. Estudios sobre la fauna de parasitos de peces Ant- arcticos. Republica Argentina, Secretaria de Marina, Ser- vicio de hidrografia naval. 910: 1-84. Utevsky, A.Y. 1993. A new marine leech Nototheniobdella sawyeri gen.nov., sp.nov. (Hirudinea, Piscicolidae) from Antarctic seas. Zoosystematica Rossica 2: 237-240. Utevsky, A.Y. 1994. A new Genus of Antarctic Piscicolid leeches (Hirudinea, Rhynchobdellida). Vestnik Zoologii 4-5: 73-76 (in Russian). Utevsky, A.Y. 1995. A New Antarctic leech genus (Hiru- dinea, Piscicolidae). Vestnik Zoologii 5-6: 3-12 (in Russian). Utevsky, A.Y. 1997. A new species of piscicolid leeches (Hirudinea, Piscicolidae) from Antarctic seas. Vestnik Zoologii 31: 17-24 (in Russian). 78 Utevsky, A.Y. 2000. Fauna of Antarctic fish leeches (Hiru- dinea, Piscicolidae). Visnyk Kharkivskogo Universyte- tu 456: 104-107 (in Russian). Utevsky, A.Y. 2005. An identification key to Antarctic fish leeches (Hirudinea:: Piscicolidae). Ukrains’kij Antark- tychnij Zhurnal 3: 135-144. Utevsky, S.Y., Utevsky, A.Y., Schiaparelli, S., & P. Trontelj. 2007. Molecular phylogeny of pontobdelline leeches and their place in the descent of fish leeches (Hirudinea, Pis- cicolidae). Zoologica Scripta 36: 271-280. Williams, J.I., & E.M. Burreson. 2006. Phylogeny of the fish leeches (Oligochaeta, Hirudinida, Piscicolidae) based on nuclear and mitochondrial genes and morphology. Zoologica Scripta 35: 627-639. Yang, T. 1980. Two marine leeches from Antarctic fish of the genus Notothenia. Acta Zool. Sinica 33: 373-377. 8. INDEX Accessory connection 7, 19-21 Accessory glands 7, 15, 18 Antarctobdella crozetensis 10, 66, 68 Antarctobdella tcherniai 10, 66, 68 Anus 7, 17 Atrium 7, 15 Austrobdella 27, 40, 42 Austrobdella translucens 10, 22-25, 27, 42, 44, 45 Body 7, 12 E@litellum 7, 12, 13, 15 Coelomic (lacunar) system 16, 19-21 Common part (ejaculatory ducts) 7, 15, 18 Conductive tissue 7, 15, 18 Conductive tissue cord 7, 18 Conductive tissue mass 7, 18 Copulatory area 7, 15 Copulatory bursa 7, 15, 18 @rop7,15,.17 Crop chambers 7, 15, 17 Crop region 7, 12, 13 Cryobdella 25, 27, 40, 52 Cryobdella antarctica 10, 22, 23-25, 27, 53, 54, 56-58 Cryobdella levigata 10, 22-25, 27, 54, 55 Cryobdella badovi 11, 22-25, 27, 53, 54, 56, 59, 60 Cryobdella pallida 11, 22-25, 27, 53, 54, 58, 61, 62 Cryobdellina bacilliformis 10, 22-25, 64, 66, 68 Digestive system 15, 17 Dorsal lacuna 7, 19-21 Dorsal vessel 7, 20 Ejaculatory ducts 7, 15, 18 Epididymis 7 Epsteinia 27, 40, 41 Epsteinia alba 22-25, 27, 41-43 Esophageal diverticula 7, 15, 17 Esophagus 7, 15, 17 Eye-like spots 7, 12, 14 Eyes 7, 12, 14 Female reproductive system 15, 18 Galatheabdella 11, 28, 60, 69 Galatheabdella bruuni 69-71 Ganglion region 7, 19 Glyptonotobdella 10, 27, 40, 49 UTEVSKY, ANTARCTIC PISCICOLID LEECHES Glyptonotobdella antarctica 10, 22-25, 27, 49, 51-53 Glyptonotobdella epshteini 56 Flosts 23,24 Initial part (ejaculatory ducts) 7, 15 Intestinal region 12, 13 Intestine 15, 17 Lateral lacunae 7, 19-21 Lateral processes 7, 17 Male reproductive system 15, 18 Main connection 7, 19, 20 Marginal flanges 7, 16, 48, 50 Marginal lacunae 7, 19-21 Megaliobdella 11, 27, 28, 39 Megaliobdella szidati 11, 22-25, 39, 40 Moorebdellina 10, 27, 28, 30, 32 Moorebdellina biannulata 22-25, 27, 30, 32-34 Moorebdellina meyeri 11, 22-25, 27, 30, 32, 37, 38 Moorebdellina rugosa 11, 22-25, 27, 30-32 Moorebdellina uschakovi 10, 11, 22-25, 27, 30, 32, 5,36 Mouth-pore 7, 15 Nerve cord 15, 17, 18 Notobdella nototheniae 64 Notobdella streptocheles 51 Nototheniobdella 11, 28, 60, 75 Nototheniobdella sawyeri 11, 22-25, 28, 73-75 Ocelli 7, 12, 14 Ophthalmobdella bellisioi 10, 11, 66, 68 Oviduets 7,.15; 18 Ovisacs 7, 15, 18 Oxytonostoma varituberculata 10, 46, 47 Papillae 7, 14, 16 Piscicolinae 21, 27, 28, 60 Platybdella levigata 10, 53 Platybdellinae 21, 27, 28, 40 Pleurobdella 27, 46, 47 Pleurobdella australis 22-25, 27, 46, 47, 49-51 Pleurobdella varituberculata 22-25, 27, 46-49 Pontobdella 10, 11, 27, 28 Pontobdella biannulata 10, 34 Pontobdella rugosa 10, 30, 32 79 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 54/2007 Pontobdella tasmanica 11, 22-25, 27-29 Pontobdellinae 21, 27, 28 Posterior crop caeca 7, 15, 17 Preclitellum 7, 12, 13 Preservation 26 Proboscis 7, 15, 17 Pterobdellina australis 49 Pulsatile vesicles 7, 20, 21 Rectal dilation 7, 15, 17 Rectum 7,15; 17 Seminal receptacle 66, 68 Seminal reservoirs 7, 15, 18 Sucker 7, 12-14 Stibarobdella tasmanica 28, 29 Tentacles 7, 14 Terminal part (Ejaculatory ducts) 7, 15, 18 Testicular lacuna 8, 19-21 Authors address Andrei Y. Utevsky, Ph.D. Dept. of Zoology and Animal Ecology V.N. Karazin Kharkiv National University 61077, Kharkiv, Ukraine Email: autevsk@yandex.ru 80 Testicular region 7, 20 Testisacs 15, 16, 18 Trachelobdella 11, 28, 60, 72 Trachelobdella bathyrajae \1, 22-25, 28, 71, 72 Trachelobdellina 10, 11, 28, 60 Trachelobdellina glabra 10, 22-25, 28, 62, 63 Trachelosome 8, 12, 13 Trulliobdella 10, 11, 28, 60, 63 Trulliobdella bacilliformis 22-25, 28, 64, 66-68 Trulliobdella capitis 10, 11, 22-25, 28, 63-66 Trullioobdella alba 10, 41 Tubercles 8, 16 Urosome 8, 12, 13 Vagina 8, 16 Vasa deferetia 8, 18 Vasa efferentia 8, 18 Ventral lacuna 8, 19-21 Ventral vessel 8, 20 aie TITUTION LIBRARIES HL 1388 410 wii BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 54, 2007 Editor: Karl-L. Schuchmann Zoologisches Forschungsmuseum Alexander Koenig (ZFMK) Ornithologie Adenauerallee 160, D-53113 Bonn, Germany Druck: JF. CARTHAUS, Bonn ISBN-13: 978-3-925382-58-1 ISSN 0302-671X