(Aspidocotylea) with Comments on Postlarval Development
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Proc. Helminthol. Soc. Wash. 55(1), 1988, pp. 62-67 Revision of the Multicalycidae (Aspidocotylea) with Comments on Postlarval Development D. A. THONEY AND E. M. BURRESON Virginia Institute of Marine Science, School of Marine Science, College of William and Mary, Gloucester Point, Virginia 23062 ABSTRACT: Based upon examination of a large number of specimens from both the Pacific and Atlantic oceans, Taeniocotyle Stunkard, 1962, and Trigonostoma Szidat, 1966, are considered junior synonyms of Multicalyx Faust and Tang, 1936. Trigonostoma callorhynchi Szidat, 1966, is considered a synonym of Multicalyx elegans (Olsson, 1869). The family Multicalycidae consists of a single genus, Multicalyx, and 2 species, M. elegans from holocephalans, and M. cristata from elasmobranchs. Multicalyx elegans matures at a much smaller size than M. cristata. KEY WORDS: Multicalyx, Taeniocotyle, Trigonostoma, Platyhelminthes, Multicalycidae, Aspidocotylea, tax- onomy, gall bladder, elasmobranch, holocephalan, postlarval development, synonyms. The study of aspidogastrids in the gall bladder probably belonged to the same genus, and Gib- of chondrichthian fishes has a long, confused son and Chinabut (1984) based a new family on history (Brinkmann, 1957; Stunkard, 1962; Bray, Multicalyx because the validity of Taeniocotyle 1984) that has resulted in the establishment of and Trigonostoma was open to question. 3 monotypic genera. Olsson (1869) established Our review of the literature in conjunction with the genus Macraspis for M. elegans, a gall blad- our previous studies (Thoney and Burreson, 1986, der parasite of the holocephalan Chimaera mon- 1987) indicated that an examination of a large strosa from the North Sea. Faust and Tang (1936) number of specimens was needed to determine described Stichocotyle cristata from a single spec- if the 3 existing genera, Multicalyx, Taeniocotyle, imen collected from the spiral valve of the cow- and Trigonostoma, were valid. Consequently, a nose ray, Rhinopterus bonasus, in Biloxi Bay, careful review of pertinent literature, and an ex- Mississippi. In order to accommodate their new amination of available deposited specimens and species into Stichocotyle Cunningham, 1884, recently collected material was conducted. The Faust and Tang (1936) emended the generic de- review clearly established the need for a revision scription and subdivided the genus into 2 sub- of the family Multicalycidae. genera; Stichocotyle, to include S. (Stichocotyle) nephropis Cunningham, 1884, and Multicalyx, Materials and Methods to include S. (Multicalyx) cristata Faust and Tang, Most of the individuals examined were stained and 1936. Manter (1954) transferred S. cristata to mounted museum specimens, but some specimens of Macraspis Olsson, 1869. Stunkard (1962), noting M. cristata from our collections were studied in tem- porary glycerin mounts. In all, 47 mature specimens, that Macraspis Olsson, 1869, was preoccupied 16 immature specimens, and the relevant literature by Macraspis MacLeay, 1819, proposed the new form the basis of this revision. Important morpholog- genus Taeniocotyle to replace Macraspis, and el- ical characters were photographed and measurements evated Multicalyx to generic rank for M. cristata, were made with the aid of an ocular micrometer. Spec- which he considered genetically different. Szidat imens examined are listed below. (1966) established a new genus and species, Tri- Multicalyx cristata gonostoma callorhynchi, for worms from Cal- Two mature specimens (1981.11.23.40-45, lorhynchus callorhynchus collected off Argentina. 1981.11.23.46-48) from the British Museum (Natural He separated his new genus from Taeniocotyle History) deposited by R. van der Elst and G. J. B. Ross, respectively; 2 mature (USNM 74467) and 2 immature elegans (although Szidat did not reference Stun- specimens (USNM 74468, 74469) in the U.S. National kard [1962] and called it Macraspis elegans) based Museum Helminth Collection deposited by S. S. Hen- on the presence of a triangular cephalic shield at drix; 2 immature specimens (USNM 51694) deposited the anterior end of the worms. The validity of by H. W. Manter; 1 immature specimen (HWML 22977) in the Harold W. Manter Laboratory, Univer- some of the existing genera has been questioned sity of Nebraska State Museum deposited by H. W. in subsequent papers. Hendrix and Overstreet Manter; 3 mature specimens (USNM 79412, 79413) (1977) stated that T. elegans and M. cristata deposited by D. A. Thoney; 13 mature specimens from 62 Copyright © 2011, The Helminthological Society of Washington 63 Table 1. Meristics and morphometrics of Multicalyx elegans (mean with range in parentheses, in millimeters). Collection BM (NH)* MACN HWML, USNM USNM ZMUB Location NE Atlantic SW Atlantic SW Pacific NE Pacific NE Atlantic Host Chimaera Callorhynchus Callorhynchus Hydrolagus colliei Chimaera monstrosa callorhynchus rnilii monstrosa Site Gall bladder Gall bladder Gall bladder Gall bladder Gall bladder No. examined 1 4 8 6 17 Alveoli anterior 17 12(10-14) 13(12-13) 18(15-22) 14(13-18) to ovary Alveoli anterior 91 15(11-26) 40(31-54) 84 (77-96) 41 (11-77) to testis Total alveoli 99 21 (11-37) 55 (27-75) 101(89-113) 49(11-100) Total length 15.1 3.6 (2.3-6.5) 11.7(5.3-15.5) 18.3(14.8-21.3) 9.0 (2.5-22.6) % length anter- 21 52 (46-58) 47 (32-63) 32 (23-37) 48(24-81) ior to ovary % length anter- 94 78(71-85) 81 (65-92) 89 (80-92) 85 (72-90) ior to testis Testis L x W 0.70 x 0.62 0.39(0.14-0.72) x 0.80(0.31-1.40) x 0.76(0.40-1.25) x 0.64(0.08-1.28) x 0.25(0.14-0.40) 0.54(0.18-0.95) 0.58(0.40-1.00) 0.56(0.10-1.35) Ovary L x W 0.55 x 0.39 0.30 x 0.15 0.40(0.17-0.63) x 0.61 (0.20-0.80) x 0.32(0.10-0.67) x 0.45(0.15-0.90) 0.40(0.15-0.60) 0.23 (0.08-0.50) Egg length 0.12(0.11-0.13) 0.14(0.13-0.16) 0.12(0.10-0.15) 0.12(0.09-0.14) 0.12(0.10-0.13) Author Szidat, 1966 Manter, 1954 Henrix and Brinkmann, 1957 Overstreet, 1977 * BM (NH) = British Museum (Natural History), MACN = Museo Argentine de Ciencias Naturales, HWML = H. W. Manter Laboratory, USNM = U.S. National Museum, ZMUB = Zoological Museum, University of Bergen. our own collection; and 2 mature specimens (35 mm slides) loaned by R. A. Campbell, Southeastern Mas- Family Multicalycidae sachusetts University. Gibson and Chinabut, 1984 Taeniocotyle elegans Multicalyx Faust and Tang, 1936 One mature specimen (1976.4.8.155-156) from the =Macraspis Olsson, 1869; Taeniocotyle Stun- British Museum (Natural History), deposited by G. kard, 1962; Trigonostoma Szidat, 1966. Rees; 6 mature specimens (USNM 70861, 70862, 70863) in the U.S. National Museum Helminth Col- DIAGNOSIS: Body greatly elongated, cylindri- lection, deposited by J. E. Lynch; 1 mature specimen cal. Ventral haptor extending nearly entire length (USNM 49154) deposited by H. W. Manter; 1 im- of body, consisting of a well denned single, con- mature and 6 mature specimens (HWML 22978) in the Harold W. Manter Laboratory, University of Ne- tinuous row of alveoli, subdivided by transverse braska State Museum, deposited by H. Manter; and 7 septa. Marginal organs present laterally on each immature and 10 mature specimens (43700, 43701, transverse septum. Mouth terminal, funnel- 43702, 43703, 43704, 43708, 44230) in the Zoological shaped; pharynx muscular; intestine simple, ter- Museum, University of Bergen, Norway, deposited by A. Brinkmann, Jr. minating blindly at posterior end. Testis single, situated at posterior extremity of uterus. Vas def- Trigonostoma callorhynchi erens extending anteriorly ventral or lateral to One mature and 3 immature specimens (27.949b, intestine, forming seminal vesicle distally. Cirrus 27.950) in the Museo Argentino de Ciencias Naturales pouch claviform, enclosing seminal vesicle, "Bernardino Rivadavia" e Institute Nacional de In- vestigacion de las Ciencias Naturales, deposited by L. prostate complex, and ejaculatory duct; protru- Szidat. sible cirrus thick walled, muscular. Genital pore ventral, anterior to haptor. Ovary anterior to tes- Results tis; uterus descending from ovary posteriorly to A revision of the family Multicalycidae is pre- level of testis, then turning and ascending ante- sented below. Comparative meristics and mor- riorly to genital pore. Eggs operculate, oval in phometrics of M. elegans from various parts of shape, usually filling uterus in mature specimens. its geographic range are listed in Table 1. Vitellaria consisting of 2 laterally situated bands Copyright © 2011, The Helminthological Society of Washington 64 PROCEEDINGS OF THE HELMINTHOLOGICAL SOCIETY of follicles extending from near posterior end posterior end of body. This pattern can be seen anteriorly to level between ovary and genital pore. by examination of Table 1 and Figures 6-8. Two lateral excretory ducts descending poste- Number of alveoli anterior to testis is greater in riorly from level of pharynx to end of vitellaria larger, and presumably older, individuals. Spec- where ducts fuse to form excretory vesicle and imens from Hydrolagus collet, which were all exit through dorso-terminal pore. Parasitic, usu- mature, have many more alveoli anterior to testis ally in gall bladder of Chondrichthyes. than those from Callorhynchus callorhynchus, TYPE SPECIES: Multicalyx cristata Faust and which were mostly juveniles. The same trend is Tang, 1936. seen in percentage of length anterior to ovary, where higher percentages are found in juvenile Multicalyx cristata Faust and Tang, 1936 worms. Worms become mature during the time (Figs. 1-3) required for development of 27 to 37 alveoli. =Stichocotyle (Multicalyx) cristata Faust and Tang, 1936; Macraspis cristata (Faust and Discussion Tang, 1936) Manter, 1954; Macraspis sp. of Taxonomy Manter (1954); Multicalyx multicristata Pa- Our examination of the specimens of Multi- rukhin and Tkachuk, 1980.