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Aspects of the Morphology of Chonopeltis Thiele, 1900

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Aspects of the Morphology of Chonopeltis Thiele, 1900 ASPECTS OF THE MORPHOLOGY OF CHONOPELTIS THIELE,1900 (CRUSTACEA: BRANCHIURA) WITH SPECIAL REFERENCE TO THE REPRODUCTIVE SYSTEMS by Madeleina Jeanetta Grundlingh THESIS Submitted in partial fulfilment of the requirements for the degree DOCTOR OF PHILOSOPHY in ZOOLOGY in the FACULTY OF NATURAL SCIENCES at the ND AFRIKAANS UNIVERSITY Promotor : Prof. J.H. Swanepoel Co-promotor : Dr. A. Oldewage March 1996 This thesis is dedicated with love and appreciation to my parents, Hennie and Rita Blignaut. " The earth is the Lord's, and the fulness thereof; the world, and they that dwell therein. For he bath founded it upon the seas, and established it upon the floods. " - Psalm 24 :1-2 LED II WOULD LIKE TO EX SS MY G TITUDE TO My Maker, for granting me the mercy, ability and strength to complete this study. Prof. J.H. Swanepoel who gave me the opportunity to undertake this study, and to complete it under his competent guidance. Dr. A. Oldewage for her greatly appreciated assistance and constructive criticism during this study. The personnel at the Zoology Department of the RAU, especially Edie Lutsch for her much appreciated friendship, support and assistance in histochemical preparation procedures. Dr. Andrew Deacon, Gerhard Strydom and other Kruger National Park staff members for their assistance in the field work. Hester Roets, Karien Brink and Vernie Naidoo for their time and aid in preparing the bromides and micrographs. Dr. Louis Swanepoel and Rika Nel for their much appreciated assistence during the final preparation of the thesis. My Mother, Rita Blignaut, and Tienie van der Berg for the linguistic attention to the thesis. Ester Le Grange for the typing of the thesis and Hendre van der Berg for the typographic finishes of the thesis. My family, especially my husband, Martin, and daughter, Maritza, for their continuous patience, encouragement and support. The RAU, FRD and Sanlam for financial support during this study. ll Branchiuran crustaceans are fairly common parasites of freshwater fishes in Africa and are presently represented by not more than 200 described species belonging to four genera, namely: Argulus Miiller,1785; Dolops Audouin,1837; Chonopeltis Thiele,1900; and Dipteropeltis Calman,1912. Chonopeltis is, however, the only one of these genera which is endemic to Africa and species of this genus have been reported from a number of different freshwater fish hosts and localities throughout the Ethiopic region. Although 13 species of Chonopeltis have hitherto been described, lack of morphological detail and morphometric data in taxonomic and subsequent descriptions, has led to confusion and much controversy as to, for instance, the phylogenetic origin, identity and function of various structures. Consequently, the validity of the taxonomic identity or status and affinities of the various species are questionable. Lack of important morphological information has also caused a considerable number of gaps and deficiencies in our present understanding of various aspects of ecological and pathogenetic significance. In the present study, a comprehensive investigation on various aspects of the morphology, anatomy and histology of adult Chonopeltis specimens was conducted, using not only light microscopy, but also scanning electron microscopy (SEM), histological microtomizing and staining techniques and graphic reconstruction. This investigation is based on several specimens of Chonopeltis victori Avenant-Oldewage,1991 collected during surveys of freshwater fish parasites in 1990 and 1991 from four different fish hosts, namely: Labeo rosae S teindachner,1894; L. congoro Peters, 1852; L. ruddi Boulenger,1907 ; and Barbus marequensis A. Smith,1841, sampled at various localities in the Olifants River in the Kruger National Park, as well as several C. australis Boxshal1,1976 specimens found on L. capensis (A. Smith,1841) and L. umbratus (A. Smith,1841), sampled in Boskop Dam near Potchefstroom. ill By making use of the mentioned methods of investigation a more complete and reliable morphological description of the distinctive external structures, as well as the internal structures of the cephalic shield alae and especially the reproductive system of these parasites could be given. With the aid of semi-thin (2am) serial sections and graphic reconstruction the various components of the reproductive systems could clearly be identified and the mutual relations and histomorphology of these components be described, discussed and compared with those of other members of the Branchiura. In an attempt to determine the true taxonomic position and relationships of the recognised species of Chonopeltis, the original descriptions, morphometric data, geographical distribution, hosts and affinities are compared, summarised and discussed. The results of the present study reveal a horseshoe-shaped furrow dorsally that separates the alae from the cephalon. Ventrally the frontal ala contains two marginal chitinous grooves and four sclerotized ridges which support the frontal ala. Two pairs of dorso-ventrally orientated muscles account for the movements of the frontal ala, whilst the movements of each lateral ala are controlled by a pair of transverse muscles and some longitudinal muscles. The lateral alae appear to represent the fused pleurae of the cephalic segments. The male reproductive tract consists of paired testes, vasa efferentia, vasa deferentia, ejaculatory ducts and prostate complex as well as a single median seminal vesicle and a genital atrium. Several prominent secondary sexual structures occur on the four pairs of thoracopods. The mechanism of sperm transfer presumably involves a process during which the two female spermathecal spines, being inserted into the male genital atrium, penetrate the walls of the respective ejaculatory ducts whereafter semen is actively pumped into the spermathecal vesicles. The female reproductive tract consists of a single large ovary, two anteriorly fused oviducts of which only one is functional, a genital atrium, a single median genital aperture and a crescent-shaped fertilization chamber. The paired spermathecae, each consisting of a spermathecal vesicle, duct and spine, is located separately in the abdomen and are of considerable significance during sperm transfer from the male as well as during oviposition when stored semen is presumably injected into the yet uninseminated ova. Finally, aspects on the ecology of C. victori concerning the reproductive cycle, life cycle, and epidemiology are investigated, compared and discussed. iv ir- G Branchiuriese Crustacea is redelik algemene parasiete van varswatervisse in Afrika en word huidig verteenwoordig deur nie meer as 200 beskryfde spesies wat aan vier genera behoort, naamlik: Argulus Miiller,1785; Dolops Audouin,1837; Chonopeltis Thiele,1900; en Dipteropeltis Calman,1912. Chonopeltis is egter die enigste een van hierdie genera wat endemies aan Afrika is en spesies van die genus is al aangemeld vanaf heelwat verskillende varswatervisgashere en lokaliteite regdeur die Etiopiese wyk. Alhoewel 13 Chonopeltis spesies tot dusver beskryf is, het gebrek aan morfologiese detail en morfometriese data in taksonomiese en daaropvolgende beskrywings, aanleiding gegee tot verwarring en heelwat kontroversie oor, byvoorbeeld die filogenetiese oorsprong, identiteit en funksie van verskeie strukture. Gevolglik is die geldigheid van die taksonomiese identiteit of status en verwantskappe van die onderskeie spesies betwisbaar. Gebrek aan belangrike morfologiese inligting het ook heelwat leemtes en tekortkominge in ons huidige vertolking van verskeie aspekte van ekologiese en patogenetiese belang veroorsaak. In die huidige studie is 'n omvattende ondersoek na verskeie aspekte van die morfologie, anatomic en histologic van volwasse Chonopeltis eksemplare uitgevoer, deur gebruik to maak van nie net ligmikroskopie nie, maar ook skandeerelektronmikroskopie (SEM), histologiese mikrotomering- en kleuringstegnieke en grafiese rekonstruksie. Hierdie ondersoeke is gebaseer op verskeie eksemplare van Chonopeltis victori Avenant-Oldewage,1991 wat tydens opnames van varswatervisparasiete in 1990 en 1991 vanaf vier verskillende visgashere gevind is, naamlik: Labeo rosae Steindachner,1894; L. ruddi Boulenger, 1907; L.congoro Peters, 1852 en Barbus marequensis A. Smith,1841 wat versamel is by verskeie lokaliteite in die Olifantsrivier in die Nasionale Kruger Wildtuin, asook verskeie C. australis Boxshal1,1976 eksemplare gevind op L. capensis (A. Smith,1841) en L. umbratus (A. Smith,1841), versamel in Boskopdam naby Potchefstroom. Deur gebruik te maak van die genoemde ondersoekmetodes kon 'n meer volledige en betroubare morfologiese beskrywing van die kenmerkende uitwendige strukture, asook die inwendige strukture van die sefaliese skild-vleuels en veral die geslagstelsels gemaak word. Met behulp van semidun (2/2m) seriesnee en grafiese rekonstruksie kon die verskillende komponente van die geslagstelsels duidelik geidentifiseer word en die onderlinge verhoudings en histomorfologie van hierdie dele beskryf, bespreek en vergelyk word met die van ander verteenwoordigers van die Branchiura. In 'n poging om die ware taksonomiese posisie en verwantskappe van die erkende spesies van Chonopeltis te bepaal, is die oorspronklike beskrywings, morfometriese data, geografiese verspreiding, gashere en verwantskappe vergelyk, opgesom en bespreek. Die resultate van die huidige studie toon dat 'n perdeskoenvormige groef, die vleuels dorsaal van die sefalon skei. Ventraal bevat die vleuels twee marginale chitienagtige
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