F.A. Sibbing

Specializations and limitations in the utilization of food resources by the carp, Cyprinus carpio: a study of oral food processing

SynopsisThe wide variety of aquatic food is considered to be instrumental for the diversification in fish species. Yet their abilities and inabilities of handling food are poorly known. For these reasons the food processing and feeding repertoire of the adult carp, Cyprinus carpio, fed on a variety of food types, were analyzed by light and X-ray cinematography of the head parts and by electromyography of the head and body muscles during feeding. Nine stereotyped movement patterns (particulate intake, gulping, rinsing, spitting, selective retention of food, transport, crushing, grinding and deglutition) compose the feeding process, their sequence and frequency were adjusted to the type of food. Following quantitative morphological analysis at macroscopic, light- and electronmicroscopical level, the relations between the functioning and architecture of the feeding apparatus were established. The structure and dimensions of the mouth opening, the protrusible upper jaw, the slit-shaped pharyngeal cavity, the palatal and postlingual organ, the branchial sieve, the pharyngeal masticatory apparatus and the distribution of taste buds, mucous cells and muscle fibers along the oropharyngeal surface were the directive structural characters used for estimating the abilities in food processing. The specializations for utilizing food items and its limitations, derived from structural and functional data, are compared with diet data found in the literature in order to evaluate the relative position of the carp in competition for food in the aquatic environment. It is established that the ‘omnivorous’ carp is specialized in effective handling of several categories of aquatic food, even when these are mixed with non-food (bottom invertebrates <4% SL in diameter) since the palatal organ enables the carp to separate food from non-food. This includes very hard-skinned food items, processed with the powerful pharyngeal jaws of the fish, and to a lesser extent zooplankton (>250 μm). The carp is at the same time very limited in processing long and struggling prey (e.g. fish) as well as vegetable matter, due to the lack of oral teeth and the specialized morphology of its pharyngeal chewing apparatus. These feeding abilities agree with diet data from literature. The reported herbivorism of carp illustrates its opportunism in feeding behaviour. Specialization in feeding is discussed and the necessity to take into account the total series of post-capture feeding actions for a more complete view on trophic specialization. Food intake and the intra-oral food processing of carp are bound to the structures of its sensory, central processing and effector apparatus and to the plasticity in their functioning. These together determine its feeding efficiency in exploiting the available aquatic food resources. Next to ethological and ecological studies functional morphology is another important tool to explain the trophic interactions of fish.

Resource partitioning by Lake Tana barbs predicted from fish morphometrics and prey characteristics

We develop a food-fish model (FFM), whichquantitatively relates properties of aquaticfood types (size, shape, escape velocity,habitat, mechanical properties and chemicalquality) to feeding structures of cyprinidfish. The model is based on functionalmorphology and experiments on search, capture,selection, and internal processing of food byfish. The FFM shows which food properties aremost critical in feeding and how fish canoptimise coping with them. Relative food sizeimposes the highest demands, followed by preyvelocity, food habitat and mechanicalproperties. These overrule taxonomic affinitiesof food types. Highly demanding food types(large, fast prey, suspensions of plankton,benthic prey and mechanically tough items)impose incompatible morphological requirementson fish. We apply the FFM to the endemic Barbus species flock of Lake Tana (Ethiopia),since the structural diversity of its 14species reflects recent adaptations to trophicniches. We predict their potentials inutilising different food types by quantitativecomparisons of 35 parameters, measured for eachspecies, with the values for each foodspecialist derived from the FFM. These dietpredictions are tested against gut contentsfrom 4,711 fish, sampled over seasons andhabitats. Gut contents and predictions show agood overall fit. The value of the model isshown by its resolution in predicting resourcepartitioning among the barbs. For the 14 barbsa trophic hierarchy with six major trophicgroups is reconstructed which closely matchesthe predictions. Trophic specialists (> 65%by volume of a single food type) are alsostructurally specialised, whereas less extremeanatomical structures characterise trophicgeneralists, allowing them to switch betweenfeeding modes. Trophic generalists are bestdefined by behavioral flexibility, sincefeeding modes integrate both fish and foodcharacters. The FFM is of practical use inevaluating the role of morphological diversityin an ecosystem and enables the analysis oftrophic interactions in fish communities and ofthe cascading effects by environmental change.Such an approach can be instrumental in thedevelopment of management strategies forfisheries and in conservation of biodiversity.

Temporal and spatial distribution of microcrustacean zooplankton in relation to turbidity and other environmental factors in a large tropical lake (L. Tana, Ethiopia)

The spatial and seasonal distribution of microcrustacean zooplankton of Lake Tana (Ethiopia) was monthly studied for 2 years. Concurrently, various environmental parameters were measured and related to zooplankton distribution. Canonical Correspondence Analysis (CCA) was used to estimate the influence of abiotic factors and chlorophyll a content in structuring the zooplankton assemblage. Among the environmental factors, zooplankton abundance correlated most strongly with turbidity. Turbidity was negatively correlated with species abundance, especially for Daphnia spp. and to the least extent for Diaphanosoma spp. Analysis of variance (ANOVA) was used to determine spatial (littoral, sublittoral and pelagic zone) and temporal (four seasons) variation in zooplankton abundance. We observed significant temporal differences in zooplankton abundance, with highest densities during dry season (November–April). Only cladocerans showed significant differences in habitat use (highest densities in the sublittoral zone). %

Evolution of MHC class II &; chain-encoding genes in the Lake Tana barbel species flock (Barbus intermedius complex)

Abstract Major histocompatibility complex (MHC) class II protein polymorphism is maintained in allelic lineages which evolve in a trans-specific manner, passing from one species to descendant species. Selection pressure on peptide binding residues should be greatest during speciation, when organisms move into new environments and their MHC molecules encounter new pathogens. The isolation of MHC genes from teleost fishes, the most diverse group of vertebrates, has created possibilities for testing this hypothesis. The large barbels of Lake Tana have undergone an adaptive radiation within the last 5 million years, producing 14 morphotypes which inhabit different ecological niches within the lake. We studied the variability in class II beta chain-encoding genes of four of these morphotypes using polymerase chain reaction amplification and DNA sequencing. The sequences obtained were orthologous to four of the known class II genes from the common carp, from which barbels diverged approximately 32 million years ago. When subjected to phylogenetic analysis, the 48 sequences clustered into groups which represent allelic lineages. A comparison of nonsynonymous and synonymous substitutions between the peptide binding region codons and non-peptide binding region codons of these sequences revealed that they are under strong selective pressure.

Declining stocks of Lake Tana's endemic Barbus species flock (Pisces, Cyprinidae): natural variation or human impact?

The only remaining species flock of endemic, large cyprinid fishes is found in Lake Tana, Ethiopia. A monthly experimental trawl program was conducted in 1991-1993 and 1999-2001, sampling 12 stations distributed over three habitats differing in depth and distance to shore. The aim was to compare the total abundance, spatial distribution and proportion of juveniles of the most common Barbus species in the Bahar Dar Gulf between both periods. We found a sharp reduction (75%) in total abundance, both in number and biomass of the Barbus species and even more (90%) in the number of juveniles between the two periods. However, the spatial distribution of the different Barbus species over the three habitats had not changed. High natural variability in fish stocks might be expected in environmentally unstable lakes. Although strongly pulsed (seasonal), Lake Tana is a relatively stable system. No major differences were found in abiotic parameters in 1990s that could have caused the dramatic changes in abundance. The most likely explanation is the negative impact of the motorised, commercial gillnet fishery targeting the spawning aggregations of these barbs. The drastic decline in juveniles points especially towards serious recruitment over-fishing. The results stress the need for the immediate development of a sound management plan focussing on fishing effort restrictions during the Bat-bus breeding season

Adaptive radiation of Lake Tana's (Ethiopia) Labeobarbus species flock (Pisces, Cyprinidae)

Studying species flocks (e.g. Darwins finches, Caribbean anoline lizards, East African cichlid fishes) has proven to be highly successful in understanding the forces driving speciation. The only known, intact species flock of cyprinid fishes, the 15 Labeobarbus species in Lake Tana (Ethiopia), includes eight piscivorous species. Piscivory is a rare specialisation among the highly successful (>2000 species) but mostly benthivorous Cyprinidae. The extent and mechanisms of diversification of this remarkable Labeobarbus species flock, particularly among the unexpected piscivorous species, are still largely unknown. In the present study we demonstrate that all 15 Labeobarbus species are segregated to a great extent along spatial, trophic and/or temporal dimensions. The spatial distribution, diet (prey species but not prey size), time of active feeding and predation techniques differed significantly among the eight piscivores. Lake Tanas cyprinids displayed their retained potential for ecological diversification and speciation, including the uncommon specialisation of piscivory. The latter is probably a result of the absence of common African specialist piscivores in Lake Tana. We suggest that the evolution of Lake Tanas Labeobarbus species flock at this stage is predominantly structured by ecological selection models. The labeobarbs most likely underwent sequential stages of radiation and speciation: habitat divergence followed by trophic divergence.

In Lake Tana, a unique fish fauna needs protection.

tion of biodiversity usually focus on terrestrial habitats, especially on rain forest ecosystems (Myers 1979, Simberloff 1984, Wilson 1989). Because of this bias, the value of aquatic communities, less accessible for direct observations, is often not fully appreciated. The purpose of this article is to draw attention to a unique freshwater systemLake Tana-and especially to its fishes. Thus we want to urge the international scientific community to increase knowledge on the origin and state of the current biodiver-

Structure, development and function of the branchial sieve of the common bream, Abramis brama, white bream, Blicca bjoerkna and roach, Rutilus rutilus.

SynopsisThe filter feeding organ of cyprinid fishes is the branchial sieve, which consists of a mesh formed by gill rakers and tiny channels on the gill arches. In order to establish its possible role during growth we measured the following morphological gill raker parameters over a range of sizes in three cyprinid fishes, bream, white bream and roach: inter raker distance, bony raker length, raker width, cushion length and channel width. At any given standard length common bream has the largest inter raker distance, roach the lowest and white bream is intermediate. In the ‘comb model’ of filter feeding the inter raker distance is considered to be a direct measure of the mesh size and retention ability (= minimal size of prey that can be retained) of a filter. For the three species under study there is a conflict between the comb model and experimental data on particle retention. Lammens et al. (1987) found that common bream has a large retention ability whereas roach and white bream have a much smaller one. A new model, the ‘channel model’ (Hoogenboezem et al. 1991) has been developed for common bream; in this model the lateral gill rakers can regulate the mesh size of the medial channels on the other side of the gill slit. The present data indicate that this model is not appropriate for white bream and roach. At any given standard length white bream and roach only reach 70% of the raker length of common bream, which means that in this model the gill slits should to be very narrow during filter feeding. The gill rakers consist of a bony raker and a fleshy cushion. The bony rakers have a rather long needle-like part outside the cushion in bream, but not in white bream and roach which have blunt gill rakers. Blunt gill rakers are not suited to reduce the diameter of the medial channels. The comb model seems more appropriate for white bream and roach, but doubts about the validity of this simple model remain. The sum of the areas of the medial channels is an approximation of the area through which water flows in the filter. This channel area therefore gives an impression of the capacity or flow rate of the filter. With this capacity estimation and an estimation of energy consumption we calculated an energy ratio of filter feeding. The energy ratio decreases with increasing standard length with an exponent close to the expected exponent of -0.40. The energy ratio is highest in bream, intermediate in white bream and lowest in roach.

Preliminary insight into the age and origin of the Labeobarbus fish species flock from Lake Tana (Ethiopia) using the mtDNA cytochrome b gene

The high diversity of Cyprinid fish in Ethiopias Lake Tana appears to be an example of ecological differentiation and assortative mating leading to rapid sympatric speciation. Lake Tanas Labeobarbus species flock consists of 15 morphological and ecological distinct species. This is the first attempt to determine the age and origin and inter-species relationships of Lake Tanas Labeobarbus species using the mtDNA cytochrome b gene. Analysis of cytchrome b sequences shows that Lake Tanas species flock appears to be young but the present dataset did not unequivocally support monophyly of Lake Tanas species. Additional markers are needed to determine whether Lake Tanas labeobarbs originated from a single or multiple incursion(s) of ancestral L. intermedius in the Lake Tana drainage basin, or the disruption of an ancient continuous riverine population by the emergence of the Tissisat waterfalls. Adaptive radiation and speciation within Lake Tanas Labeobarbus species flock may have occurred in the last 10,000-25,000years, following the desiccation of Lake Tana around 17,000years ago, at the same time as Lake Victoria, however, obtaining more data using other (nuclear) markers is urgently required.

Barbus tanapelagius, A New Species from Lake Tana (Ethiopia): its Morphology and Ecology

The endemic cyprinid species flock in Lake Tana consists of 15 species of large hexaploid barbs, eight of which are piscivorous. Previously, it was assumed that all piscivores preyed on the same small barb species, Barbus trispilopleura. In this paper we present a description of morphology and ecology of a new abundant small barb species, Barbus tanapelagius sp. nova (holotype RMNH 33731) from Lake Tana, Ethiopia, which appears to be the major prey species for the large pelagic piscivorous barbs. B. tanapelagius differs clearly in morphology from the other 3 small, diploid Barbus species known from Lake Tana, B. trispilopleura Boulenger, 1902, B. humilis Boulenger, 1902 and B. pleurogramma Boulenger, 1902. Conspicuous differences are its elongated body, large eye diameter, prominent and hooked lower jaw contour and colouration. Preliminary data suggest that B. tanapelagius also differs ecologically from the other small Barbus spp. by its pelagic, strictly zooplanktivorous feeding and its occurrence mainly in the deeper, offshore waters. The other small Barbus species are most probably largely benthic feeders and dominant in the shallow inshore waters. Previous views about the evolution of the present 8 endemic piscivorous large barb species therefore require reconsideration, as the present paper shows a more complex scenario including several prey species.