Abstract. Nile crocodile (Crocodilus niloticus) mass mortality events in the Olifants River between the Letaba River confluence in South Africa and Lake Massingir in Mozambique have been attributed to pansteatitis: a disease that affects fat depots of the animals. The disease is also found in sharptooth catfish (Clarias gariepinus) in the same area, and the cause of the disease is attributed to pollution. Although the Olifants River Valley is polluted, the impact of interventions such as dam construction on biodiversity receives little attention. We show that the onset of the pansteatitis epidemic in crocodiles and sharptooth catfish at the Olifants/Letaba confluence coincided with back-flooding of Lake Massingir that changed the Olifants River from a rock and sand substrate river to a clay substrate lake. Isotopic analysis shows that sharptooth catfish shifted from a predominantly vegetarian to a piscivorous diet that is highly correlated with pansteatitis prevalence, and crocodiles and tiger fish (Hydrocynus vittatus) show coincident trophic level increases. The evidence suggests that the ecosystem change altered the structure of the lotic foodweb and that an exotic or extralimital fish has invaded the confluence and is the vector of the pansteatitis epidemic. The invasive fish species is yet to be identified. The pansteatitis epidemic is an unintended ecological consequence of damming this river.

In late 2006 an unusual ulcerative condition in wild fish was reported for the first time in Africa from the Chobe and upper Zambezi Rivers in Botswana and Namibia. Concern increased with subsistence fishermen reporting large numbers of ulcerated fish in their catches. In April 2007 the condition was confirmed as an outbreak of epizootic ulcerative syndrome (EUS). The causative agent, Aphanomyces invadans, is a pathogenic water mould of fish that shows little host specificity. Ulcers follow infection of tissues by oomycete zoospores, resulting in a granulomatous inflammation associated with invading oomycete hyphae. Granulomatous tracts surrounding oomycete hyphae within the necrotic tissues characterise the diagnostic histological picture. The upper Zambezi floodplain at the confluence with the Chobe River spans the four countries of Botswana, Namibia, Zambia and Zimbabwe, making disease control a challenge. The floodplain ecosystem supports a high fish diversity of around 80 species, and is an important breeding and nursery ground. The annual cycle of flooding brings about changes in water quality that are thought to favour the infectivity of A. invadans, with diseased fish appearing soon after the plains become flooded. Since 2006 the disease has spread rapidly upstream along the upper Zambezi and its tributaries. By 2010 the disease was reported from the Okavango Delta in Botswana and in 2011 from the Western Cape Province of South Africa. EUS has the potential to disrupt floodplain ecosystems elsewhere in Africa where high fish diversity forms the basis of subsistence fisheries and local economies, and is a direct threat to freshwater fish culture.

Large numbers of adult Nile crocodiles

This study compares the aetiology of pansteatitis in Lake Loskop, relative to two other impoundments along the Olifants River. Macroscopic and microscopic pathology, age determination and analysis of stomach content, fatty acids and stable isotopes explain the high prevalence of pansteatitis in Oreochromis mossambicus (Peters) and several other species in Lake Loskop. All the dietary indicator comparisons between pansteatitis-affected and healthy fish fail to support a systemic cause. Pansteatitis in Lake Loskop was linked to size and weight of O. mossambicus, but not to ontogenic age. Fish in Lake Loskop showed abnormally high omega-3 to omega-6 fatty acid ratios normally only found in marine fish with no significant difference in degree of assimilation of these fatty acids between pansteatitis-affected and healthy fish. This explains the vulnerability to, but not the occurrence of, pansteatitis. As a cause for the pansteatitis, these results point towards sporadic vitamin E-depleting trigger events, known sporadic fish die-off occurrences that provide surviving fish with a rich source of rancid fats on which to scavenge. The mechanism ties pansteatitis to eutrophication and trophic cascade effects, the intrinsic drivers of the disease and suggests an adaptive management strategy that might be applied by relevant conservation authorities.

Pansteatitis has been identified in wild populations of sharptooth catfish, Clarias gariepinus (Burchell), and Nile crocodiles, Crocodylus niloticus Laurenti, inhabiting the same waters in the Olifants River Gorge in the Kruger National Park, South Africa. Mesenteric and pectoral fat tissue was investigated microscopically and by fatty acid analysis in healthy and pansteatitis-affected catfish from both captive and wild populations. Variation in fatty acid composition between pectoral and mesenteric fat was noted. Composition of mesenteric fat differed between fish from various localities as a result of differences in diet. Pansteatitis in the captive population, resulting from ingestion of high amounts of dietary oxidized fat, reflected higher levels of unsaturated fatty acids within the mesenteric fat. Mesenteric fat of pansteatitisaffected wild catfish was characterized by an increase in moisture content, a decrease in fat content and a decrease in stearic and linoleic acids. The n-3 to n-6 fatty acid ratio of mesenteric fat was higher in pansteatitis-affected wild catfish than in healthy catfish from the same locality, reflecting higher polyunsaturated fat intake by pansteatitisaffected fish. The possible role of alien, invasive, phytoplankton-feeding silver carp, Hypophthalmichthys molitrix (Valenciennes), in the aetiology of pansteatitis in both catfish and crocodiles in the Olifants Gorge is discussed.

Pansteatitis was confirmed in sharptooth catfish, Clarias gariepinus (Burchell), from three main locations within the Kruger National Park (KNP); the Olifants River Gorge, Engelhard Dam on the Letaba River and from the Sabie River in the Sabiepoort. An increasing prevalence of pansteatitis was observed in catfish during repeated samplings from the Olifants Gorge from 2009 to 2011 and co-existence of old and recent lesions indicated on-going incitement of pansteatitis. Only a low prevalence of pansteatitis was observed in catfish sampled from the Olifants River upstream of the Gorge in the KNP and no pansteatitis was observed in catfish sampled from a rain-filled dam not connected to the Olifants River. Common to both the Olifants Gorge and the Sabiepoort is the damming of the rivers in Mozambique to form lakes Massingir and Corumana respectively. Anthropogenic activities resulting in potential pollution of the rivers differ greatly between these two catchments, providing argument against a primary pollution-related aetiology of the pansteatitis found at these two sites. Compared with other sites, analysis of stomach contents of catfish from the Olifants Gorge and the Sabiepoort strongly suggested that consumption of a predominantly fish diet was associated with the development of pansteatitis in these fish. In a farmed population of catfish used as positive control, development of pansteatitis could be ascribed to consumption of rancid fish waste from a trout slaughterhouse. In the Olifants Gorge, alien invasive silver carp, Hypophthalmychthys molitrix (Valenciennes), seasonally migrate upstream out of Lake Massingir to spawn. This schooling species is an obligate phytoplankton feeder with consequent high levels of adipose tissue n-3 polyunsaturated fatty acids. In the Olifants Gorge, at least, this may explain seasonal exposure to levels of polyunsaturated fats in the diets of catfish and crocodiles to which these animals are not adapted. The possible roles of diet, membrane lipid composition and metabolic rate of fish, sediment pollution and seasonal drop in environmental temperature in the pathogenesis of pansteatitis in the catfish are discussed. Further studies are needed to verify some of these speculations.

In late 2006, diseased fish of a variety of species began to appear in the Chobe and upper Zambezi rivers in southern Africa. In April 2007, investigations showed that the levels of pesticides and heavy metals in the tissues of the fish were very low, discounting pollution as an underlying cause for the disease. However, histological evidence showed that the disease closely resembled the epizootic ulcerative syndrome caused by the oomycete Aphanomyces invadans, a serious aquatic pathogen that has been isolated from freshwater and estuarine fish in Japan, south-east Asia, Australia and the usa since the 1970s, but not previously recorded in Africa.

A field outbreak of pox virus infection in juvenile Nile crocodiles (Crocodylus niloticus), in which high morbidity and negligible mortality occurred, is described. Histopathological examination of the skin lesions revealed numerous large intracytoplasmic inclusions in the dermis and a very mild dermal inflammatory reaction. Scanning electron microscopical examination of the skin revealed the presence of large numbers of virus particles in the inclusions. Skin lesions persisted for 5 to 6 months.