Process and pattern in cichlid radiations - inferences for understanding unusually high rates of evolutionary diversification

The cichlid fish radiations in the African Great Lakes differ from all other known cases of rapid speciation in vertebrates by their spectacular trophic diversity and richness of sympatric species, comparable to the most rapid angiosperm radiations. I review factors that may have facilitated these radiations and compare these with insights from recent work on plant radiations. Work to date suggests that it was a coincidence of ecological opportunity, intrinsic ecological versatility and genomic flexibility, rapidly evolving behavioral mate choice and large amounts of standing genetic variation that permitted these spectacular fish radiations. I propose that spatially orthogonal gradients in the fit of phenotypes to the environment facilitate speciation because they allow colonization of alternative fitness peaks during clinal speciation despite local disruptive selection. Such gradients are manifold in lakes because of the interaction of water depth as an omnipresent third spatial dimension with other fitness-relevant variables. I introduce a conceptual model of adaptive radiation that integrates these elements and discuss its applicability to, and predictions for, plant radiations.

Nematodes and meiofauna in sediments obtained from RRS Charles Darwin cruise CD179, Portuguese continental margin

Samples collected at two different depths (ca. 3200 and ca. 4200 m) in the Setúbal and Cascais canyons off the Portuguese coast, during the HERMES RRS Charles Darwin cruise CD179, were analysed for (1) sediment biogeochemistry (TOC, TN) and (2) composition, and structural and trophic diversity of nematode communities. Multivariate PERMANOVA analysis on the nematode community data revealed differences between sediment layers that were greater than differences between canyons, water depths, and stations. This suggests that biogeochemical gradients along the vertical sediment profile are crucial in determining nematode community structure. The interaction between canyon conditions and the nematode community is illustrated by biogeochemical patterns in the sediment and the prevalence of nematode genera that are able to persist in disturbed sediments. Trophic analysis of the nematode community indicated that non-selective deposit feeders are dominant, presumably because of their non-selective feeding behaviour compared to other feeding types, which gives them a competitive advantage in exploiting lower-quality food resources. This study presents a preliminary conceptual scheme for interactions between canyon conditions and the resident fauna.

Results from field observations and in situ 13C feeding experiments in the Nazaré Canyon conducted during RRS James Cook 10 cruise Leg 2

Submarine canyon systems provide a heterogeneous habitat for deep-sea benthos in terms of topography, hydrography, and the quality and quantity of organic matter present. Enhanced meiofauna densities as found in organically enriched canyon sediments suggest that nematodes, as the dominant metazoan meiobenthic taxon, may play an important role in the benthic food web of these sediments. Very little is known about the natural diets and trophic biology of deep-sea nematodes, but enrichment experiments can shed light on nematode feeding selectivity and trophic position. An in-situ pulse-chase experiment (Feedex) was performed in the Nazaré Canyon on the Portuguese margin in summer 2007 to study nematode feeding behaviour. 13C-labelled diatoms and bacteria were added to sediment cores which were then sampled over a 14-day period. There was differential uptake by the nematode community of the food sources provided, indicating selective feeding processes. 13C isotope results revealed that selective feeding was less pronounced at the surface, compared to the sediment subsurface. This was supported by a higher trophic diversity in surface sediments compared to the subsurface, implying that more food items may be used by the nematode community at the sediment surface. Predatory and scavenging nematodes contributed relatively more to biomass than other feeding types and can be seen as key contributors to the nematode food web at the canyon site. Non-selective deposit feeding nematodes were the dominant trophic group in terms of abundance and contributed substantially to total nematode biomass. The high levels of 'fresh' (bioavailable) organic matter input and moderate hydrodynamic disturbance of the canyon environment lead to a more complex trophic structure in canyon nematode communities than that found on the open continental slope, and favours predator/scavengers and non-selective deposit feeders.

Effects of elevated CO2 and temperature on an intertidal meiobenthic community

In the near future, the marine environment is likely to be subjected to simultaneous increases in temperature and decreased pH. The potential effects of these changes on intertidal, meiofaunal assemblages were investigated using a mesocosm experiment. Artificial Substrate Units containing meiofauna from the extreme low intertidal zone were exposed for 60 days to eight experimental treatments (four replicates for each treatment) comprising four pH levels: 8.0 (ambient control), 7.7 & 7.3 (predicted changes associated with ocean acidification), and 6.7 (CO2 point-source leakage from geological storage), crossed with two temperatures: 12 °C (ambient control) and 16 °C (predicted). Community structure, measured using major meiofauna taxa was significantly affected by pH and temperature. Copepods and copepodites showed the greatest decline in abundance in response to low pH and elevated temperature. Nematodes increased in abundance in response to low pH and temperature rise, possibly caused by decreased predation and competition for food owing to the declining macrofauna density. Nematode species composition changed significantly between the different treatments, and was affected by both seawater acidification and warming. Estimated nematode species diversity, species evenness, and the maturity index, were substantially lower at 16 °C, whereas trophic diversity was slightly higher at 16 °C except at pH 6.7. This study has demonstrated that the combination of elevated levels of CO2 and ocean warming may have substantial effects on structural and functional characteristics of meiofaunal and nematode communities, and that single stressor experiments are unlikely to encompass the complexity of abiotic and biotic interactions. At the same time, ecological interactions may lead to complex community responses to pH and temperature changes in the interstitial environment.

The state and known impacts of fish species transferred to or within continental Africa

Fish introductions have been made from small fish ponds to the largest lakes in Africa. The primary intent of these introductions has been to sustain or increase fish production, although some introductions have been made to develop sport fisheries and to control unwanted organisms. Some of these introductions have fulfilled their objective in the short term, but several of these "successful" introductions have created uncertainties about their long term sustainability. Lates niloticus, Oreochromis niloticus, O. leucostictus, Tilapia melanopleura and T. zilli were introduced into lakes Victoria and Kyoga in 1950s and early 1960s. By the 1980s O. niloticus and O. niloticus dominated the fisheries of these lakes, virtually eliminating a number of endemic fish species. The loss of genetic diversity of the fish in the worlds second largest lake has also been accompanied by a loss of trophic diversity. The transformation of the fish community has, in Lake Victoria coincided with a profound eutrophication (algal blooms, fish kills, hypolimnetic anoxia) which might be related to alterations of the lake's food-web structure. In contrast, the introduction of a planktivore, Limnothrissa miodon into Lake Kivu and the Kariba reservoir has established highly successful fisheries with little documented effect on the pre-existing fish community or trophic ecology of the lakes. The highly endemised species-rich African Great lakes may be particularly sensitive to species introductions and require special consideration and caution when introductions are contemplated because species extinctions, introgressive hybridization and ecosystem alterations may occur following fish introductions.

Análise biogeográfica da dieta de um predador de topo ao longo da região Paleártica

Existem vários estudos sobre a dieta de predadores de topo a uma escala local, mas o estudo da dieta de uma espécie ao longo de diferentes regiões geográficas poderá permitir a deteção de certos padrões e variações ecológicas. Para estudar a variação biogeográfica da dieta do bufo-real (Bubo bubo) ao longo do Paleártico criou-se uma base de dados baseada em 192 estudos. Pretendia-se analisar padrões em macro-escala de descritores da dieta e avaliar o efeito de diferentes condições ambientais, de descritores da paisagem e da disponibilidade de presas. Registaram-se 346813 presas de 698 espécies distintas. As principais presas são os lagomorfos e os roedores, enquanto que as aves constituem importantes presas alternativas. Temperaturas mais elevadas favorecem a diversidade trófica, mas esta diminui com a latitude e a longitude. Os resultados revelam a natureza oportunista do bufo-real e a sua elevada adaptabilidade trófica a diferentes habitats e condições ambientais; Biogeographic analysis of a top predator's diet across the Paleartic Region Summary: There are several studies about top predators' diet at a local scale, but studying a species diet across different geographic regions may allow the detection of certain patterns and variations which might influence its ecological features. We created a database of 192 papers to study the diet of the Eurasian Eagle Owl (Bubo bubo) across the Palearctic region. We analysed large scale biogeographical patterns of diet descriptors in relation to different environmental conditions, including climate, landscape and prey availability. We recorded 346816 preys from 698 different species. The main prey groups are lagomorphs and rodents, whereas birds represent important alternative prey. Higher temperatures favour a higher trophic diversity, which in turn decreases at lower latitudes and longitudes. The results reveal the opportunistic nature of the Eurasian Eagle Owl and its high trophic adaptability to different habitats and environments.

The meso-scale response of subarctic North Pacific seabird community structure to lower trophic level abundance and diversity

Understanding the mechanisms that structure communities and influence biodiversity are fundamental goals of ecology. To test the hypothesis that the abundance and diversity of upper-trophic level predators (seabirds) is related to the underlying abundance and diversity of their prey (zooplankton) and ecosystem-wide energy availability (primary production), we initiated a monitoring program in 2002 that jointly and repeatedly surveys seabird and zooplankton populations across a 7,500 km British Columbia-Bering Sea-Japan transect. Seabird distributions were recorded by a single observer (MH) using a strip-width technique, mesozooplankton samples were collected with a Continuous Plankton Recorder, and primary production levels were derived using the appropriate satellite parameters and the Vertically Generalized Production Model (Behrenfeld and Falkowski 1997). Each trophic level showed clear spatio-temporal patterns over the course of the study. The strongest relationship between seabird abundance and diversity and the lower trophic levels was observed in March/April ('spring') and significant relationships were also found through June/July ('summer'). No discernable relationships were observed during the September/October ('fall') months. Overall, mesozooplankton abundance and biomass explained the dominant portion of seabird abundance and diversity indices (richness, Simpson's Index, and evenness), while primary production was only related to seabird richness. These findings underscore the notion that perturbations of ocean productivity and lower trophic level ecosystem constituents influenced by climate change, such as shifts in timing (phenology) and synchronicity (match-mismatch), could impart far-reaching consequences throughout the marine food web.

Predator diversity enhances secondary production and decreases the likelihood of trophic cascades

We manipulated the diversity of top predators in a three trophic level marine food web. The food web included four top benthic marine fish predators (black goby, rock goby, sea scorpion and shore rockling), an intermediate trophic level of small fish, and a lower trophic level of benthic invertebrates. We kept predator density constant and monitored the response of the lower trophic levels. As top predator diversity increased, secondary production increased. We also observed that in the presence of the manipulated fish predators, the density of small gobiid fish (intermediate consumers) was suppressed, releasing certain groups of benthic invertebrates (caprellid amphipods, copepods, nematodes and spirorbid worms) from heavy intermediate predation pressure. We attribute the mechanism responsible for this trophic cascade to a trait-mediated indirect interaction, with the small gobiid fish changing their use of space in response to altered predator diversity. In the absence of top fish predators, a full-blown trophic cascade occurs. Therefore the diversity of predators reduces the likelihood of trophic cascades occurring and hence provides insurance against the loss of an important ecosystem function (i.e. secondary production).

Effects of genetic diversity of grass on insect species diversity at higher trophic levels are not due to cascading diversity effects

Genetic diversity in plant populations has been shown to affect the species diversity of insects. In grasses, infection with fungal endophytes can also have strong effects on insects, potentially modifying the effects of plant genetic diversity. We manipulated the genetic diversity and endophyte infection of a grass in a field experiment. We show that diversity of primary parasitoids (3rd trophic level) and, especially, secondary parasitoids (4th trophic level) increases with grass genetic diversity while there was no effect of endophyte infection. The increase in insect diversity appeared to be due to a complementarity effect rather than a sampling effect. The higher parasitoid diversity could not be explained by a cascading diversity effect because herbivore diversity was not affected and the same herbivore species were present in all treatments. The effects on the higher trophic levels must therefore be due to a direct response to plant traits or mediated by effects on traits at intermediate trophic levels.

Ecological communities in variable environments : dynamics and diversity under coloured environmental stochasticity

While environmental variation is an ubiquitous phenomenon in the natural world which has for long been appreciated by the scientific community recent changes in global climatic conditions have begun to raise consciousness about the economical, political and sociological ramifications of global climate change. Climate warming has already resulted in documented changes in ecosystem functioning, with direct repercussions on ecosystem services. While predicting the influence of ecosystem changes on vital ecosystem services can be extremely difficult, knowledge of the organisation of ecological interactions within natural communities can help us better understand climate driven changes in ecosystems. The role of environmental variation as an agent mediating population extinctions is likely to become increasingly important in the future. In previous studies population extinction risk in stochastic environmental conditions has been tied to an interaction between population density dependence and the temporal autocorrelation of environmental fluctuations. When populations interact with each other, forming ecological communities, the response of such species assemblages to environmental stochasticity can depend, e.g., on trophic structure in the food web and the similarity in species-specific responses to environmental conditions. The results presented in this thesis indicate that variation in the correlation structure between species-specific environmental responses (environmental correlation) can have important qualitative and quantitative effects on community persistence and biomass stability in autocorrelated (coloured) environments. In addition, reddened environmental stochasticity and ecological drift processes (such as demographic stochasticity and dispersal limitation) have important implications for patterns in species relative abundances and community dynamics over time and space. Our understanding of patterns in biodiversity at local and global scale can be enhanced by considering the relevance of different drift processes for community organisation and dynamics. Although the results laid out in this thesis are based on mathematical simulation models, they can be valuable in planning effective empirical studies as well as in interpreting existing empirical results. Most of the metrics considered here are directly applicable to empirical data.

Intersexual and spatial heterogeneity in trophic attributes of the sleeper, Bostrychus africanus (Eleotridae) in the Qua Iboe Estuary, Nigeria

There was variation in the ingestion of the food objects by the sexes. Despite the similarity in rank-order of the food objects, the ingestion of the objects vary significantly (rg=0.320, P>0.05). Dipterans adult and Hymenoptera were the only food objects not eaten by the males whereas insect remains and unidentified bivalves were absent from the trophics spectrum of the females. There was significant increase in feeding intensity by females than males. There was significant increase in GRI by specimens from Nipa Creek whereas individuals from mangrove creek recorded higher MGF and vice-versa. Dipterans adult. Hymenoptera, insect remains, Neritina glabrata and unid bivalves were absent from dietaries for nipa creek whereas a complete array of the food objects were eaten in the mangrove creek. The present findings highlights the importance of the mangrove ecosystem as the native vegetation encompassing great diversity of food resources and living conditions than the succeeding alien nipa vegetation

The trophic dynamics of summer flounder (Paralichthys dentatus) in Chesapeake Bay

Data on the trophic dynamics of fishes are needed for management of ecosystems such as Chesapeake Bay. Summer flounder (Paralichthys dentatus) are an abundant seasonal resident of the bay and have the potential to impact foodweb dynamics. Analyses of diet data for late juvenile and adult summer flounder collected from 2002−2006 in Chesapeake Bay were conducted to characterize the role of this flatfish in this estuary and to contribute to our understanding of summer flounder trophic dynamics throughout its range. Despite the diversity of prey, nearly half of the diet comprised mysid shrimp (Neomysis spp.) and bay anchovy (Anchoa mitchilli). Ontogenetic differences in diet and an increase in diet diversity with increasing fish size were documented. Temporal (inter- and intra-annual) changes were also detected, as well as trends in diet reflecting peaks in abundance and diversity of prey. The preponderance of fishes in the diet of summer flounder indicates that this species is an important piscivorous predator in Chesapeake Bay.

The diversity and abundance of zooplankton in the Lake Victoria and Kyoga basins and their relationship to fish production

Biological diversity of an ecosystem is considered a reliable measure of the state of health of the ecosystem. In Uganda's large lakes, the Victoria and Kyoga, the past three decades have been characterized by profound changes in fish species composition following the introduction of the piscivorous Nile perch (Oguto-Ohwayo 1990). Over 300 haplochromine cichlid species comprising a wide range of trophic groups were lost along with a host of non-cichlid fishes which occupied virtually all available ecological niches and in the lakes (Witte 1992). A second major ecological event has been the gradual nutrient enrichment of the water bodies (eutrophication) from diffuse and point sources, while at the same time pollutants have also gained entrance into the water systems in pace with indusfrial development and human population increases in the lake basins. Eutrophication and pollution have drastically altered the physical and-chemical character of the water medium in which different fauna and flora thrive. In Lake Victoria these alterations have resulted in changes of algal species composition from pristine community dominated by chlorophytes and diatoms (Melosira etc) to one composed largely of blue-green algae or Cyanobacteria (Microcystis, Anabaena, Planktolyngbya etc) (Mugidde 1993, Hecky 1993).

Fish species diversity in the Victoria and Kyoga lake basins: their conservation and sustainable use

Introduction of exotic fish species especially the Nile perch Lates niloticus, is believed to be responsible for the decline of fish species diversity in lakes Victoria, Kyoga and Nabugabo.About 60% of the haplochromine cichlids are thought to have become extinct from L. Victoria due to predation by the Nile perch. However there are many lakes satelite to the lakes Victoria and Kyoga basins which still have fish fauna similar to that of the main lakes. many of the satellite lakes are separated from the main lakes in, which Nile perch was introduced by extensive swamps that provide a barrier to Nile perch .A survey was carried out in a number of these satelite lakes and an inventory made of existing fish species. Their distribution and relative abundances were also determined. The lakes studied included Nawampasa, Nakuwa,Kawi Lamwa Gigate, Nyaguo, Agu, Nabugabo. Kayanja, Kaytigi, Mburo, Kachera and Wamala.Some habitats within the main lakes Victoria and Kyoga, especially those with rocky outcrops· and macrophyte cover that provide refugia for endangered species from Nile perch,were also surveyed) Various stations along the River Nile were also sampled to quantify the fish species that are still resent. Kyoga minor lakes were found to have the highest number of fish species especially of haplochromine cichlids. Many haplochromine trophic groups that were thought to be extinct from 1. Victoria still occur in these lakes.!Some of the satellite lakes, especially lakes Kayugi, Mburo and Kachera still contain .healili populations of oreochromis. I esculentus that could be used as brood stock in fish farming. Many of these lakes should .I ( I therefore be protected for conservation offish species diversity

Genetic status of selected fish taxa in relation to conservation of genetic and species diversity in the Victoria and Kyoga Lake basins

Cichlids are known for their explosive radiation especially in the African Great Lakes marked with a high level of lake endemism. These fishes have been characterized mainly along trophic and habitat differences, by variation in morphological structures such as teeth and jaws and by differences in body shape and coloration. Cichlids are important as a microcosm of macroevolution. The explosive radiation, young evolutionary scale, and the isolation of groups characterized with high levels of endemism and presence of living fossils makes the group important for evolutionary and genetic studies. Lake Victoria region cichlids which are isolated and relatively more recent in evolution were the last to be appreciated in their diversity. Recently Ole Seehausen has found scores of rock fishes in Lake Victoria which were up to then thought to be absent from the Lake and only known to occur in Lakes Malawi and Tanganyika. Greenwood put together the species groups of Lake Victoria, and later in the early 1980's revised the classification of haplochromine species to reflect the phyletic origin and interrelationship of the various groups in Lake Victoria region. Melan Stiassny has been interested in early evolution of cichlids while the likes of Paul Fuerst and Lees Kaufman and Axel Meyer have been interested and are working to explain the speciation mechanisms responsible for the explosive radiation and evolution of cichlids. Locally S.B Wandera and his student Getrude Narnulemo are spearheading the biodiversity and taxonomic studies of cichlids in Lake Victoria region