Evaluating Biodiversity and Conserving Lake Malawi's Cichlid Fish Fauna

The cichlid fishes of the Great Lakes of Africa are extraordinary biological assemblages. Conservation strategies are urgently needed for these fish faunas, due to increased threats from over-fishing, habitat degradation, and species introductions. Here I investigate the patterns of biodiversity of the rock-dwelling cichlid fishes (known as mbuna) of Lake Malawi and present a series of recommendations to enhance their conservation. Specifically, I examine intralacustrine biogeograpbic distributions and diversity, and levels of endemism for both mbuna species and genera. The biogeographic patterns show a high degree of variation for both taxonomic designations. Significant relationships are found between diversity and distribution patterns. Speciose genera are more widespread and habitats close to other lands support higher biodiversity than in isolated islands. The results indicate that the genus level may be the appropriate taxonomic designation to consider for conservation purposes, and that areas of high generic endemism, such as the Muleri Islands, are significant for conservation. In order to account for geographic variation in biodiversity, consideration should be given to expanding Lake Malawi National Park boundaries to include areas in the northern and central parts of the lake, especially Likoma and Chisumulu Islands and Nkhata Bay. I discuss the establishment of an international park and the expansion of conservation efforts to include the non-mbuna cichlids. An ecosystem approach to biodiversity conservation and management is advocated.     

Evolutionary and Conservation Biology of Cichlid Fishes as Revealed by Faunal Remnants in Northern Lake Victoria

Lake Victoria until recently harbored the world's second richest lacustrine fish assemblage, but it is now experiencing a mass extinction. Here we report on the current status of the endemic cichlid fishes from a preliminary biotic inventory conducted as part of limnological reconnaissance in Kenyan and Ugandan waters during 1989–1992. Sixty-two haplochromine taxa were recovered, half of them new. Some are local endemics, while others are more widely distributed, based on review of earlier collections. Comparison with data from 1982–1984 and 1988 suggests that some shoreline assemblages have been locally stable over the past decade. In addition to littoral rock and vegetation, four kinds of refugia for indigenous fishes are recognized: (1) schools of Rastrineobola , (2) benthic microbial mats in deep water, (3) satellite lakes, (4) the oxycline. Reconnaissance by remote operated vehicle (ROV) documented fish kills due to deoxygenation of the water column, indicating the risk of using the oxycline as a refugium. Several species formerly abundant in Lake Victoria were found only in satellite lakes, revealing the importance of these lakes in the conservation of indigenous species. Recent events in Lake Victoria shed light on mechanisms of speciation and mass extinction, insights relevant to conservation planning for the valuable remnants of the lake's indigenous fauna.     

Conservation of the African Great Lakes: A Limnological Perspective

The limnology of Lakes Malawi Tanganyika, and Victoria is discussed with the objective of examining how the physical, chemical, and biological properties of the lakes will determine the response of these ancient great lake ecosystems to human activities. Of the physical properties discussed, large dilution capacities and long flushing times can make the detection and removal of chemical pollutants in these lakes difficult. The outflows of all three lakes are small because of high evaporation losses, and as a result lake levels are responsive to climate changes that would alter evaporation:precipitation ratios. Increased nutrient input to these lakes will likely result in a decrease in the volume of oxygenated water and available fish habitat. Plankton community composition will also change, probably toward dominance by cyanobacteria. While the effects of eutrophication on fish production are difficult to predict, changes in plankton composition would almost certainly be accompanied by changes in fish community structure. Recent studies of water chemistry and plankton productivity in Lake Victoria provide evidence of possible eutrophication. Because land use has a diffuse but potentially large impact on these lakes, conservation strategies must take into account human activities within entire drainage basins. This requires cooperation between riparian countries and technological and financial input from the international community. An immediate need is the establishment of monitoring programs to determine the impact, real or potential, of human activities around the lakes.     

The Effects of Predation by Nile Perch, Lates niloticus L., on the Fish of Lake Nabugabo, with Suggestions for Conservation of Endangered Endemic Cichlids

As in Lake Victoria and Lake Kyoga, most of the native fish species of Lake Nabugabo have either been depleted or have disappeared since Lates niloticus , Nile perch, and Oreochromis niloticus , a tilapiine, were introduced. L. niloticus, O. niloticus, Schilbe intermedius, Brycinus jacksonii , and Rastrineobola argentea are the only species still abundant and widespread in the lake. Of the five haplochromine species endemic to Lake Nabugabo, at least one has vanished and the remaining ones are very rare. The prey of the Nile perch in Lake Nabugabo, with the exception of the prawn Caridina nilotica , which is absent from the lake, are similar to those the predator switched to in Lakes Victoria and Kyoga after haplochromines had been depleted. These consist of ephemeropterans—especially Povilla , the anisopteran nymphs R. argentea , tilapiines, and juvenile Nile perch. Three small lakes adjacent to Lake Nabugabo—Lake Kayanja, Lake Kayugi, and Lake Manywa—contain large numbers of Oreochromis esculentus and two haplochromine species previously known only in Lake Nabugabo. These three lakes should be designated conservation areas for the cichlids and other native species that were previously abundant in Lakes Victoria, Kyoga, and Nabugabo. The swamps around these lakes should be protected so that the Nile perch does not spread into them. Further research should be carried out to ensure the preservation of the endemic species.     

Effects of Introduced Salmonids on a Montane Population of Iberian Frogs

Abstract:  Amphibians are declining worldwide because of multiple factors, including human-mediated introduction of fishes into naturally fishless areas. Although several studies have focused on the effect of exotic fishes on native amphibians breeding in ponds or lakes, little is known about their effects on stream-breeding species. We studied the effects of introductions of native brown trout ( Salmo trutta ) and exotic brook trout ( Salvelinus fontinalis ) on the stream-breeding, endemic Iberian frog (  Rana iberica ) in a protected area in central Spain. We assessed occurrence patterns of tadpoles and salmonids and compared habitat use of the three species. We also determined experimentally whether chemical cues from salmonids elicited antipredator behavior in tadpoles. Finally, we assessed the relative influence of tadpole habitat preferences, differences in salmonid species, and invasion geography on tadpole occurrence. Despite widely overlapping habitat preferences, tadpoles and trout did not coexist, with the former restricted to fishless habitats. Tadpoles detected chemical cues from both trout species and reacted by decreasing their activity, although the response toward the native brown trout was stronger. The residual distribution of Iberian frogs in Peñalara is better explained by the geography of fish invasions than by the fish species involved. Measures such as fish extirpation from certain areas, aimed at recovering lost habitat and improving connectivity among remaining populations of Iberian frogs, seem critical for the species' long-term survival in central Spain.     

Patterns of the Remnant Cichlid Fauna in Southern Lake Victoria

During the years 1984–1987 Lake Victoria in East Africa experienced what is probably the largest mass extinction of contemporary vertebrates. Within a decade about 200 endemic species of haplochromine cichlids disappeared. The extinctions that occurred in the 1980s have been documented predominantly on species of offshore and sub-littoral waters in the Mwanza Gulf of southern Lake Victoria. Although the littoral fauna of southern Lake Victoria had not been examined in detail, their diversity seemed less affected by the changes in the ecosystem. We give results of the first comprehensive inventory of the littoral cichlid fauna in southern Lake Victoria and discuss its conservation status. We also report on new developments in the sub-littoral fauna after 1990. More than 50 littoral and 15 sub-littoral stations were sampled between the years 1991 to 1995. Of the littoral stations, 34 were sampled for the first time. One hundred sixty three species of haplochromines were collected. Of these, 102 species were previously unknown. About two thirds of them live in rocky areas that were sampled for the first time. Littoral rocky habitats harbored the highest diversity. Since 1990, however, 13 more species disappeared from established sampling stations in littoral habitats. Fishing practices, spreading of exotic fishes, water hyacinth, and eutrophication are considered important threats to the littoral fauna. Also in the upper sub-littoral the number of species declined further. On deeper sub-littoral mud bottoms individual and species numbers increased again, although they are nowhere close to those found before the Nile perch (Lates niloticus) upsurge. This fauna differs from the well studied pre-Nile perch fauna. At well-established sampling stations, the sub-littoral zone is dominated by previously unknown species, and some known species have performed dramatic habitat shifts.     

Helpers in fish

Summary Field data show that in the cichlid fish Lamprologus brichardi conspecifics other than the reproducing pair help in brood care and territory maintenance. The expected degree of relatedness between helpers and the eggs or larvae they tend lies between 0.25 and 0.5, decreasing with the helper's age. This decrease might influence the point of time at which helpers depart. Five other endemic Lake Tanganyika cichlids showing rather similar helping behaviour are described.     

Benthic-slope fishes found at 900m depth along a transect in the western N. Atlantic Ocean

The species composition of the benthic-slope fish fauna of the Middle Atlantic Bight is described from trawl catches taken from a depth of about 900 m in August, 1969. Two species associations and an intermediate area are apparent. The difference between the two associations is partially attributable to differences in species composition, but primarily to a difference in species dominance. The small number of samples precludes geographic and bathymetric delineation of these associations, as well as conclusions about their temporal stability. Diversity values appear to be relatively high for the benthic fishes of the upper slope, but comparison with values for temperate estuaries and coastal fishes indicates a narrow range for all of these habitats, and considerable overlap in values from the different environments. Environmental stability may be of less importance in determining the diversity of fish faunas than of benthic invertebrate faunas because of the mobility of fishes and ontogenetic changes in their habitat preference.     

A Regional Perspective on the Diversity and Conservation of Tropical Andean Fishes

Abstract: The tropical Andes harbor an extraordinarily varied concentration of species in a landscape under increasing pressure from human activities. Conservation of the region's native plants and animals has received considerable international attention, but the focus has been on terrestrial biota. The conservation of freshwater fauna, particularly the conservation of fishes, has not been emphasized. Tropical Andean fishes are among the most understudied vertebrates in the world. We estimate that between 400 and 600 fish species inhabit the diverse aquatic environments in the region. Nearly 40% of these species are endemic. Tropical Andean fishes are vulnerable to ongoing environmental changes related to deforestation, water withdrawals, water pollution, species introductions, and hydropower development. Additionally, their distributions and population dynamics may be affected by hydrologic alterations and warmer water temperatures associated with projected climate change. Presently, at least three species are considered extinct, some populations are endangered, and some species are likely to decline or disappear. The long‐term persistence of tropical Andean fishes will depend on greater consideration of freshwater systems in regional conservation initiatives.     

Conservation of Stream Fishes: Patterns of Diversity, Rarity, and Risk

Abstract: North America has a rich fauna of freshwater fishes which attains greatest diversity in the central and southeastern United States. Many stream fishes have limited ranges and are locally rare and patchily distributed. Local diversity increases downstream and total diversity follows typical species-area relationships. Drainages linked with the large Mississippi system support more species than those of comparable area flowing directly to the sea and rivers isolated by falls have notably few species. The between-drainage component of diversity is large. Threats to this fauna, which are not addressed by management focused on threatened species, include fragmentation of drainage networks by impoundments and homogenization of faunas by interbasin connections and introductions. Conservation efforts require a biogeographic perspective; they should focus on streams of intermediate size (orders 4–6) plus the upstream portion of each drainage and should attempt to maintain total diversity and, inclusively, populations of rare or threatened species.     

Environmental issue of exotic catfish culture in Uttar Pradesh

A survey was conducted in Uttar Pradesh to enlist the available exotic fish species in culture. The culture and feeding details of unofficially introduced Clarias gariepinus was collected from 419 ponds existing in rural areas of 23 districts of Uttar Pradesh. The data indicated that 32% ponds were culturing exotic fishes of which 79% cases are of monoculture of exotic African catfish. Feeding analysis of these exotic African catfish revealed that 7% cases subsist on natural pond feed while 29% cases are fed on slaughter house waste. In 32% cases of the exotic magur monoculture, chicken waste was given whereas 11% cases were recorded with fish waste feeding. The possible threats and environmental risks of such unauthorised and indiscreminate culture of exotic magur has been discussed in this paper.     

Effects of Widespread Fish Introductions on Paedomorphic Newts in Europe

Abstract:  As a result of factors such as global warming, habitat destruction, and species introduction, amphibians are declining worldwide. No one, however, has analyzed the status of polymorphic amphibian species at a national or continental scale, although some local reports exist. Our aim was to report on the loss of intraspecific heterochrony as a loss to diversity in determining the consequences of fish stocking on European populations of paedomorphic newts. Paedomorphosis is a polymorphism in which larval traits are retained in the adult stage. We surveyed 39 paedomorphic populations of the alpine (  Triturus alpestris ) and palmate ( T. helveticus ) newts, all but one of which initially occupied fishless ponds and lakes in France, Italy, Slovenia, Bosnia, Montenegro, and Greece. Exotic fishes were found in 44% of the studied aquatic habitats, with a 100% presence in Montenegro. At all sites paedomorphs disappeared and metamorphs declined. Only fish explained these population changes because alternative factors such as drying were not significant. More catastrophically, fish introductions occurred in habitats known to support the largest populations of newts and even some endemic subspecies. If management and legislative measures are not taken to stop fish stocking, protect paedomorphs as conservation units at national and international levels, and restore natural habitats, all the largest paedomorphic populations may disappear in the near future. Their disappearance would represent a loss of one of the rare, fascinating examples of intraspecific heterochrony.     

Sex and status in a cooperative breeding fish: behavior and androgens

Among taxonomically widespread cooperatively breeding vertebrates, those with non-breeding helpers-at-the-nest provide an excellent opportunity to understand the proximate mechanisms underlying care and allocare. In this study, we examined androgen levels in relation to care behavior in a cooperatively breeding cichlid fish, Neolamprologus pulcher, from Lake Tanganyika. We concentrated on androgens, as these hormones have been linked to the defense behavior, and the defense of young is a common form of parental care in fishes. N. pulcher dominant female breeders performed the most care and also displayed the highest levels of plasma testosterone (T) compared with other individuals within the social group. We also found that dominant male breeders provided a similar amount of care as did the subordinate helpers, but breeding males had the highest levels of 11-ketotestosterone (11KT), an important androgen in fish. Breeders had higher levels of both androgens (T and 11KT) compared to helpers. There was a weak but significant positive correlation between T levels and the frequency of care regardless of sex and status. Our results suggest that androgens may promote defense of young and are in contrast to the commonly reported trade-off between androgen and parental care.     

Littoral Fish Communities in Lake Tanganyika: Irreplaceable Diversity Supported by Intricate Interactions among Species

The rocky littoral areas of Lake Tanganyika harbor diverse fish communities, mainly composed of cichlids. Their stability, structure, and organizing mechanism were examined at three locations by census, behavioral observations, and dietary analyses. These fish communities were stable, characterized by their persistence during a 10-year period and resilience after a perturbation. Partitioning of spawning sites among substrate spawners was evident, which might be a factor in their ability to coexist. The fish communities were consistently composed of 12 food-habit groups, but composition of species in each group differed from location to location. Among species of different food-habit groups, facultative commensalisms were prevalent. Although aggressive interactions are common among fishes of the same food-habit group, each predatory fish obtained an advantage in feeding efficiency from the different feeding behaviors of other species of the same group. This mutualism and the facultative commensalism should both increase species richness in different parts of the communities. These intricate interactions among species may be a base of stability of the fish communities. We assert that maintenance of ecological networks among species are most important for conservation of biodiversity in tropical regions.     

Deep-sea bottom-living fishes at two repeat stations at 2 200 and 2 900 m in the Rockall Trough,northeastern Atlantic Ocean

Repeated sampling of deep-sea bottom-living fishes was conducted at two stations in the Rockall Trough at depths of 2 200 and 2 900 m. The 2 200 m station (M) was sampled 16 times by an Agassiz trawl between April 1978 and April 1985 and yielded 473 fish belonging to 17 species. The 2 900 m permanent station (PS) yielded 781 fish belonging to 11 species from 9 Agassiz trawls, 2 small box otter trawls, 4 semi-balloon otter trawls and 7 epibenthic sledges between March 1975 and April 1985. Macrourid fishes were numerically dominant at both stations with Coryphaenoides guentheri and Coryphaenoides (Nematonurus) armatus being the most abundant species at M and PS, respectively. In terms of biomass the morid, Antimora rostrata, was dominant at M but C. (N.) armatus was dominant at PS. The only species that showed clear evidence of a seasonal reproductive cycle was C. guentheri. Many species showed no indication of reproducing in the Rockall Trough. The diets of almost all the species are described and considered in relation to the food resources exploited. Investigations on the pelagic and benthic invertebrate faunas at these stations should reveal the extent to which the bottom-living fishes exploit the available resources.     

Human Impacts on Drainages of the Mesa Central, Mexico, and Its Genetic Effects on an Endangered Fish, Zoogoneticus quitzeoensis

Abstract:  The Mesa Central of Mexico is of special conservation interest due to its high richness of freshwater fish species, of which the goodeines are one of the most representative groups. Through an integrated approach, we determined conservation priorities for goodeine populations. We based our recommendations on the genetic diversity (variation in five microsatellite DNA loci) in 10 populations of Zoogoneticus quitzeoensis and on an analysis of ecological (e.g., presence of exotic species), social (e.g., political situation), and environmental (e.g., pollution) information for 52 historical occurrence points for species in the genus Zoogoneticus . Patterns of genetic erosion and genetic diversity indices were closely associated with human impact. Recent bottleneck events were most evident in the populations from remnants of the lakes drained at the beginning of the twentieth century. We identified seven operational conservation units (OCUs), all of which should be conserved because they contain unique portions of the total variation of the species. Special attention needs to be given to increase genetic variability, recover population sizes, and reestablish contact among populations within OCUs. It is imperative to create an integrative and effective approach for the recovery and conservation of the freshwater fish diversity of Central Mexico that is based on social and natural sciences.     

Introduced trout sever trophic connections in watersheds: consequences for a declining amphibian

Trophic linkages between terrestrial and aquatic ecosystems are increasingly recognized as important yet poorly known features of food webs. Here we describe research to understand the dynamics of lake food webs in relation to a native riparian amphibian and its interaction with introduced trout. The mountain yellow-legged frog Rana muscosa is endemic to alpine watersheds of the Sierra Nevada Mountains and the Transverse Ranges of California, but it has declined to a small fraction of its historical distribution and abundance. Although remaining frogs and introduced trout feed in different habitats of alpine lakes, our stable-isotope analyses clearly show that the same resource base of benthic invertebrates sustains their growth. During one period, insect emergence from naturally fishless lakes was nearly 20-fold higher compared to adjacent lakes with trout, showing that fish reduce availability of aquatic prey to amphibious and terrestrial consumers. Although trout cannot prey on adult frogs due to gape limitation, foraging post-metamorphic frogs are 10 times more abundant in the absence of trout, suggesting an important role for competition for prey by trout in highly unproductive alpine watersheds. Most Sierran lakes contain fish, and those that do not are usually small isolated ponds; in our study, these two lake types supported the lowest densities of post-metamorphic frogs, and these frogs were less reliant on local, benthic sources of productivity. Since Rana muscosa was formerly the most abundant vertebrate in the Sierra Nevada, the reduction in energy flow from lake benthos to this consumer due to fish introductions may have had negative consequences for its numerous terrestrial predators, many of which have also declined. We suggest that disruptions of trophic connections between aquatic and terrestrial food webs are an important but poorly understood consequence of fish introduction to many thousands of montane lakes and streams worldwide and may contribute to declines of native consumers in riparian habitats.     

Assessing Changes in Amphibian Population Dynamics Following Experimental Manipulations of Introduced Fish

Abstract: Sport‐fish introductions are now recognized as an important cause of amphibian decline, but few researchers have quantified the demographic responses of amphibians to current options in fisheries management designed to minimize effects on sensitive amphibians. Demographic analyses with mark–recapture data allow researchers to assess the relative importance of survival, local recruitment, and migration to changes in population densities. I conducted a 4‐year, replicated whole‐lake experiment in the Klamath Mountains of northern California (U.S.A.) to quantify changes in population density, survival, population growth rate, and recruitment of the Cascades frog (Rana cascadae) in response to manipulations of non‐native fish populations. I compared responses of the frogs in lakes where fish were removed, in lakes in their naturally fish‐free state, and in lakes where fish remained that were either stocked annually or no longer being stocked. Within 3 years of fish removals from 3 lakes, frog densities increased by a factor of 13.6. The survival of young adult frogs increased from 59% to 94%, and realized population growth and recruitment rates at the fish‐removal lakes were more than twice as high as the rates for fish‐free reference lakes and lakes that contained fish. Population growth in the fish‐removal lakes was likely due to better on‐site recruitment of frogs to later life stages rather than increased immigration. The effects on R. cascadae of suspending stocking were ambiguous and suggested no direct benefit to amphibians. With amphibians declining worldwide, these results show that active restoration can slow or reverse the decline of species affected by fish stocking within a short time frame.     

An Evaluation of Restoration Efforts in Fishless Lakes Stocked with Exotic Trout

Abstract: Detrimental effects of introduced fishes on native amphibian populations have prompted removal of introduced cutthroat ( Oncorhynchus clarki ), rainbow ( Oncorhynchus mykiss ), and brook trout (  Salvelinus fontinalis ) from naturally fishless lakes at Mt. Rainier National Park, Washington ( U.S.A.). Using paleolimnological indicators (diatoms, invertebrates, and sediment characteristics) in eight 480-year-old sediment cores from eight lakes, we (1) derived estimates of baseline environmental conditions and natural variation, (2) assessed the effects of stocking naturally fishless lakes, and (3) determined whether lakes returned to predisturbance conditions after fish removal (restoration). Diatom floras were relatively stable between 315 and 90 years before present in all lakes; we used this time period to define lake-specific "baseline" conditions. Dissimilarity analyses of diatoms revealed sustained, dramatic changes in diatom floras that occurred approximately 80 years ago (when fish were introduced) in four of five stocked lakes, whereas the diatom floras in two unstocked lakes had not changed significantly in the last 315 years. Diatoms were not preserved in an eighth lake. State changes also occurred in two lakes over 200 years before European settlement of the Pacific Northwest. Preserved invertebrate densities fluctuated dramatically over time in all cores, providing a poor reference for assessing the effects of fishes. Nevertheless, fish-invertebrate interactions have been demonstrated in other paleolimnological studies and may be useful for lower-elevation or more productive lakes. Because diatom communities have not returned to predisturbance assemblages in restored lakes, even 20–30 years after fish removal, we conclude that Mt. Rainier lakes were not successfully restored by the removal of fishes.     

Female-biased immigration and male peace-keeping in groups of the shell-dwelling cichlid fish Neolamprologus multifasciatus

The small cichlid fish Neolamprologus multifasciatus is endemic to Lake Tanganyika and is unique among fish because it lives in complex social groups with several reproductive males and females, all of which participate in defending their territory against neighbors and intruders. Individuals use empty snail shells for breeding and shelter. Previous parentage analysis using microsatellites suggested occasional exchange of individuals between groups. In field experiments, we found that females showed a higher tendency than males to migrate into territories already occupied by a resident pair. The phenomenon and causes of female-biased immigration were further investigated in aquarium experiments: Nine of 15 females, but only 2 of 15 equally sized males, settled in territories of established pairs. Territorial males exhibited more aggression toward strange males (potential reproductive competitors) than toward strange females (potential additional mates); their females were more aggressive toward strange females (probably competitors for shells) than toward strange males. Apparently, a conflict exists between the sexes regarding the immigration of additional females. This conflict seems to be the selective basis for observed active male interference in aggressive disputes between females in the territory. Interfemale tolerance in a group was greater in the male’s presence than in his absence. Received: 1 February 2000 / Received in revised form: 2 May 2000 / Accepted: 5 May 2000