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FINAL VERSION JUNE 14 2010X Nematodes from the Bakwena Cave in Irene, South Africa Candice Jansen van Rensburg Academic year: 2009-2010 Thesis submitted in partial fulfillment of the requirements for the award of the degree Master of Science in Nematology in the Faculty of Sciences, Ghent University Promoter: Prof. Wilfrida Decraemer Co-Promoters: Dr. Wim Bert Dr. Antoinette Swart Nematodes from the Bakwena Cave in Irene, South Africa Candice JANSEN VAN RENSBURG 1*,2 1Nematology section, Department of Biology, Faculty of Sciences, Ghent University; K.L. Ledeganckstraat 35, 9000 Ghent, Belgium 2Dept. Zoology & Entomology, P.O. 339, University of the Free State, Bloemfontein, 9300, South Africa; [email protected] *Corresponding e-mail address: [email protected] 1 Summary A survey forming part of the Bakwena cave project was carried out from January 2009 to February 2010 at the Bakwena Cave South Africa. A total of 27 nematode genera belonging to 23 families were collected, 19 genera are reported for the first time from cave environments. Of the six localities sampled, the underground pool of the cave showed the highest species diversity with lowest diversity associated with fresh and dry guano deposits. Four of the sampling localities were dominated by bacterial feeders the remaining two localities being comprised of fungal feeders, obligate and facultative plant feeders and omnivores. Multidimensional scaling indicated six nematode assemblages corresponding with six localities, which might reflect substrate associated patterns. Three species are also described, two being new to science. Diploscapter coronatus is characterised by having a visibly annulated cuticle; a pharyngeal corpus clearly distinguishable from the isthmus, the vulva situated about mid-body and the stoma almost twice as long as the lip region width. Panagrolaimus n. sp. is characterised by an almost straight habitus for females and J-shaped habitus for males, lips amalgamated into three pairs, lateral field with two incisures, ovary with straight reflexed part and germinal zone extending till anal level or even beyond into the tail cavity, post uterine sac shorter than body width, vulva at 69% (mean value) with protruding lips, female tail elongate conoid to conoid, male tail conoid with short mucron, spicules robust with hemispheroid manubrium, a short calamus and lamina with dorsal hump and ventral wing, gubernaculum curved ventrad. Plectus n. sp. is characterised by its small body size, labial region not set off, cephalic setae almost as long as half labial diameter, amphidial fovea at posterior end of stoma, vulva at mid-body, each genital branch about two body widths long, tail ventrally arcuate and relatively short. In addition to the morphological diagnostic features described for Plectus n. sp. a polytomous key for females of the genus Plectus is also presented. Descriptions, measurements and illustrations including SEM micrographs are provided for the three species. Keywords: Ecology, karst system, Pretoria, morphology, new species, Diploscapter coronatus , SEM, taxonomy, key 2 The study of cave systems and particularly karst systems of South Africa are still in their initial stages (Durand, 2008). Karst is a special kind of landscape that is formed by the dissolution of soluble rocks such as dolomite. Sinkholes, caves and complex underground water flow networks are found associated with a karst system (Williams, 2008). Approximately 25% of the world’s population inhabit areas above karst regions and obtain water from its aquifers (Buchanan, 2008). Karst regions are therefore very important to be protected from any kind of pollutants moreover because they appear extremely sensitive to such disturbance (Buchanan, 2006). South Africa holds 12% of the world’s karst systems, with the Gauteng Province having one of the largest single repositories of karst in the world (Buchanan, 2008). In South Africa, Karst systems are facing immense pressures from unsustainable over-exploitation. This over- exploitation comes in the form of pollution of groundwater and surface habitats, urban development and roads, unauthorized removal of dolomites, fossils and cave formations by public and the over-use of natural resources in the areas, including lime operations and agriculture. Cave environments have been considered by many authors (Culver et al. , 1999, 2004; Culver & Sket, 2000; Hodda et al. , 2006) as extreme habitats which are inhabited by only a few specialised species (Abolafia & Peña-Santiago, 2006a). Cave fauna can include obligate (troglobites), edaphic (edaphobites) and aquatic (stygobites) cave dwellers while some species may be facultative (troglophiles), indifferent cave soil dwellers (trogloxenes) or even accidentals (Abolafia & Peña-Santiago, 2006a). A cave may be classified into three main zones according to light intensity: the twilight zone, middle zone and deep zone (Durand, 2008). The most diverse fauna in a cave occurs in the twilight zone near the entrance. The middle zone is in complete darkness but has variable temperature and supports several species, some of which may commute to the surface (Poulson & White, 1969). The deep zone is characterised by complete darkness and constant temperatures. The different fauna associated with these three zones show varying degrees of adaptivity towards troglomorphism, with many cave-dwelling organisms becoming obligatory troglobites, which reproduce, feed and spend their whole life underground (Poulson & White, 1969). Other organisms such as certain bat species need caves for roosting, hibernation and 3 reproduction, but hunt outside caves. The guano deposits of these bats serve as food for decomposers such as bacteria and fungi (Durand, 2008). The most widely studied group of organisms from cave environments are arthropods (Welbourn, 1999) with other taxa such as nematodes only being mentioned as present without specification (Abolafia & Peña-Santiago, 2006a). Little is known on the nematofauna of cavernous ecosystems especially of those forming part of a karst system. Altogether 28 nematode genera have been reported from cave habitats worldwide (Hodda et al. , 2006). The nematode species found in this unusual type of environment seem to consist of a few endemic species on the one hand and accidental occupants that can tolerate a wide variety of environmental conditions on the other hand (Hodda et al. , 2006). Muschiol & Traunspurger (2007) showed that bacterial mats in caves are dominated by five nematode genera: Chronogaster Cobb, 1913, Monhystrella Cobb, 1918, Panagrolaimus Fuchs, 1930, Poikilolaimus Fuchs, 1930 and Udonchus Cobb, 1913. Chronogaster troglodytes Poinar & Sarbu, 1994 can be considered as a truly cavernicolous nematode which is regularly found associated with bacterial mats relying on autochthonous primary production from thermal springs (Poinar & Sarbu, 1994). Members of the genera Plectus Bastian, 1865 and Panagrolaimus have previously been reported from cave environments (Cayrol, 1973; Poinar & Sarbu, 1994) from the south of France, and Romania respectively. Diploscapter Cobb, 1913 on the other hand is not listed from this extreme habitat (Hodda et al. , 2006). A checklist of free living nematodes recorded from freshwater habitats in southern Africa which catalogued about 37 families, 66 genera and 140 nematode species (Heyns, 2002), did not report on any nematodes from cave habitats. The available information of species from the families Panagrolaimidae Thorne, 1937, Diploscapteridae Micoletzky, 1922 and Plectidae Örley, 1880 in South Africa is limited with only a few records being known (Heyns, 1971; Dassonville, 1981; Botha & Heyns, 1993; De Bruin & Heyns, 1993). Since the passing of Juan Heyns no recent taxonomic information is available on free living nematodes from South Africa. 4 The current study is part of the Bakwena Cave Project initiated by the South African Karst Ecology Study Group of the University of Johannesburg and the Biosystematics division of the Plant Protection Research Institute (PPRI) of the Agricultural Research Council (ARC), Pretoria. The objectives of the study undertaken herein were: 1) to identify all nematodes from the Bakwena cave to genera level; 2) to taxonomically describe some key species or new species from the different habitats within the cave; 3) to determine possible distribution patterns of the nematode genera and 4) to determine if the cave shows any signs of an ecological disturbance Overall the above mentioned objectives aim to contribute to the Bakwena cave biodiversity study, as well as to increase our current knowledge on free living nematodes from South Africa. Material and methods STUDY AREA : BAKWENA CAVE The Bakwena Cave located south of Pretoria in the Gauteng Province, was chosen as study area. The cave appears as a sinkhole in dolomite with its entrance obscured by a thicket and a few Celtis africana Burm.f. trees along the periphery . The perimeter of the cave is about 10m in diameter and tapers down to an almost vertical shaft of about 9m in diameter and 21m deep, sloping in a north-easterly direction (Fig. 1B). At the bottom of the shaft there is a talus slope consisting of scree (about 13m along its east-west axis) forming the entrance to the main chamber which is approximately 15m wide along its north-south axis, and 21m long along its east-west axis. 5 Fig. 1: Diagram showing simplified map of A) a vertical and B) a horizontal section through the Bakwena Cave in Irene South Africa (courtesy of Francois Durand, Dept. Zoology, University
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