Intra- and interspecific social organization among three herbivorous cichlid fishes in Lake Tanganyika

Intra- and interspecific social organization among 3 coexisting herbivorous cichlids,Tropheus moorii, Petrochromis trewavasae andP. orthognathus, were studied at a rocky shore in Lake Tanganyika. Individuals of all species maintained discrete intra- and interspecific territories, exceptT. moorii andP. trewavasae whose feeding territories overlapped. Territory owners attacked smaller individuals but exhibited displays towards larger neighbours, irrespective of species. This observation suggests that both intra- and interspecifically, these cichlids interact in the context of size-dependent dominance hierarchies. Removal experiments showed that smallerT. moorii benefitted fromP. trewavasae in the maintenance of their territory borders against largerP. orthognathus. Interspecific dominance hierarchy and commensalism among these cichlids are not species specific, but change dynamically in relation to the difference in body size between component individuals. Mating territories of maleP. orthognathus andT. moorii are compared to their feeding territories, and territory forms are discussed. It is suggested that an approach incorporating the concept of an interspecific society may play an important role in the elucidation of guild structure and function.     

Depth-related differences in territory size and defense in the herbivorous cichlid,neolamprologus moorii, in lake tanganyika

Neolamprologus moorii, a small herbivorous cichlid of Lake Tanganyika, defends its territory against food competitors. Territory size and territorial defense were compared between two sites at different depths. In the shallow site, more fishes intruded into the territories than in the deep site, owing to the higher density fish population in the former. The territory holders in the shallow site attacked intruders more often, although the ratio of attacks to number of intruders was similar to that of the deep site. The territories in the deep site were larger and contained a greater amount of algae than the shallow site. The larger territory size in the deep site may have resulted from the lower cost of territory defense due to the lower competitor density and from the need to maintain a greater amount of food resources to compensate for lower algal productivity.     

Increased energy investment in testes following territory acquisition in a maternal mouthbrooding cichlid

Underwater observations conducted in Lake Tanganyika showed that males of a maternal mouthbrooding cichlid, Petrochromis fasciolatus, defended a mating territory for at least several months, but left at approximately noon every day. After the experimental removal of males from their territories, new owners occupied the vacated territories within several days. New owners exhibited higher body-condition factors and fat indices than the original owners. These results suggest that the new owners had not previously occupied a territory, considering the physical exhaustion of owners owing to the energy expenditure on territory maintenance and mating behaviors and a limited time for feeding. Original owners had heavier testes than new owners, despite similar body sizes. This disparity in testis weight suggests that energy investment in testes increases following territory acquisition. Among males that newly occupied a territory during the observation period, larger individuals occupied territories sooner, suggesting that large body size facilitates competition for territory. It is concluded that energy investment in testes is delayed for somatic growth until territory acquisition.     

Overlapping territories ofPseudosimochromis curvifrons males and other herbivorous cichlid fishes in Lake Tanganyika

Males of the herbivorous cichlid fishPseudosimochromis curvifrons established mating territories 3–10 m in diameter, which included both spawning and feeding sites. Territorial males attacked conspecific males and also other species. Only conspecific males were chased out of the territories. Territorial males attacked other species at the spawning sites while courting or waiting for females and at other sites in their territories while patrolling and foraging. Attacks against herbivorous species were more frequent than against non-herbivores since herbivores were much more abundant. Territories of four abundant herbivorous cichlids largely overlapped the territories ofP. curvifrons males. Symbiotic relationships were not detected between the species. Instead, they were aggressive towards each other but coexisted by segregating feeding sites in the overlapping areas. The aggressive coexistence ofP. curvifrons males with other herbivores may have resulted from the energetic costs of defending their relatively large territories against all food competitors. Interspecific food-resource partitioning may also have facilitated the coexistence.     

Overlapping territory of the benthophagous cichlid fish, Lobochilotes labiatus, in Lake Tanganyika

Synopsis Feeding territories do not usually overlap in the same fish species. We studied overlapping territories and feeding behaviours of a benthophagous cichlid fish Lobochilotes labiatus Boulenger in a coastal area of Lake Tanganyika. Forty-four individuals of both sexes, 6–31 cm in total length, maintained territories in a 21 m × 25 m area. These could be regarded as feeding territories. The territories of similar-sized owners (usually different by less than 5 cm) did not overlap each other, but those of different-sized owners overlapped widely. Aggressive interactions were rarely observed between fish distinctly different in body size. Both large and small fish took benthic animals of the same kinds and of similar size. In contrast, the size of foraging sites (crevices) was different between them: the large fish foraged at large crevices and small fish at small crevices. This partitioning of foraging sites may enable the overlap of territories in this benthivore.     

Territorial behavior of threespot damselfish (Eupomacentrus planifrons) increases reef algal biomass and productivity

Synopsis Algal growth and damselfish (Eupomacentrus planifrons) territories were studied in two reef habitats at Discovery Bay, Jamaica. Damselfish territories were contiguous in the reef flat (0 to 2.5 m), where the algal composition and biomass varied from territory to territory. In contrast, on the lower reef terrace (22 m), damselfish territories were often spatially segregated. While the algal composition of the territories was more uniform on the reef terrace, the total algal biomass was lower than in the territories on the reef flat. Damselfish are largely herbivorous, and they defend their territories against most intruding fish, including a number of herbivorous species. Areas of the reef terrace outside of damselfish territories were heavily grazed by herbivorous fishes and contained only small quantities of non-crustose algae.The reef terrace territories were characterized by a multispecific turf of algae (greens, blue-greens, and reds) covering the Acropora cervicornis framework and by the leafy, brown alga, Lobophora variegata. A rapid reduction in the biomass of brown algae and filamentous algae was noted when damselfish were permanently removed from their territories. Only calcified, encrusting algae — plants apparently somewhat undesirable as fish food sources — would be common on the terrace zone of this reef if damselfish territories were absent. Damselfish territoriality may significantly influence the dynamics of some reefs by increasing the biomass of the algal turf thereby increasing; reef productivity. Since blue-green algae, potential nitrogen fixers, occur in these algal turfs, the fish may also be indirectly affecting reef nutrition.     

Coexistence of permanently territorial cichlids of the genus Petrochromis through male-mating attack

Coexistence mechanisms of permanently territorial fishes proposed hitherto have been mainly based on interspecific competition. To test the hypothesis of the coexistence through male-mating aggression (Kohda, 1995a), spacing patterns of feeding territories of three coexisting herbivorous cichlids, Petrochromis polyodon, P. trewavasae and P. famula, were investigated in a homogeneous habitat at a shallow rocky shore in Lake Tanganyika. All three species maintained individual feeding territories that were defended against both conspecifics and congenerics and rarely overlapped either intra- or interspecifically. Territories of three species were scattered and covered almost all rocky bottoms. Territorial attacks usually occurred near the border of feeding territories. But male P. polyodon, the largest and socially most dominant fish, attacked conspecific males far beyond their feeding territories, which were widely separated. Removal of some territory owners suggested that such attacks result in the separate distribution of male P. polyodon territories. The interstitial space between P. polyodon feeding territories resulting from their mating aggression can be occupied by subordinate congeneric fishes. Male-mating attack of dominant males of Petrochromis will facilitate the coexistence of other congeners.     

Factors in habitat partitioning in Pseudotropheus aurora (Pisces: Cichlidae), an introduced species to a species-rich community of Lake Malawi

The territorial behaviour of a small rock-dwelling cichlid, Pseudotropheus aurora , was studied at Thumbi Island West at the southern end of Lake Malawi. P. aurora is one of several species that have been introduced to an already diverse fish community. Many of the endemic and introduced species are highly territorial. The coexistence of an introduced territorial species within a speciesrich community of other territorial species is examined. Territories from which P. aurora individuals were removed were invariably re-occupied by conspecifics. Similarly, when all territories of a certain area were emptied of all territory owners of any existing species, the former P. aurora territories were re-occupied by conspecifics.
The possible roles of site specificity, competitive exclusion and behaviourally transmitted traditions regarding territories are discussed.     

Determinants of territory size in the pomacentrid reef fish,Parma victoriae

Summary Factors governing the size of territories defended by the pomacentrid reef fish, Parma victoriae, were investigated, prompted by contradictory predictions in the literature concerning the effects of food supply and competitors. Observations were carried out over the non-breeding period (March–October) on a medium density population in which territories were partially contiguous. The territory size of adult fish varied between 3 and 26 m², and was inversely correlated with local densities of conspecifics. The same range in territory size was found for both males and females, which did not differ in the time they spent on territory defence and foraging activities. No correlation existed between territory size and the abundance of algal food, body size, age or time spent on territory defence. Also, there was little variation in territory size over time, despite seasonal changes in the abundance of food algae.Experimental reduction of food supplies on isolated territories of males and females had no effect on territory size. In a higher density habitat an experiment was carried out in which population density and food abundance were simultaneously manipulated. This showed that territory size was primarily determined by intraspecific interactions, as territories exhibited considerable increases in size upon removal of neighbours. No changes in the size of defended areas resulted from either artificial increases or decreases of food levels. There were also no changes in the time spent on defence of territories, foraging time or feeding rates associated with food manipulations or territory expansion, which suggested that food was not a limited resource. This conflicted with current theories proposed to explain territory defence and expansion. It is hypothesized that intraspecific interactions constrain territory size well below the optimum in terms of the abundance of preferred food algal species.     

Testes Investment along a Vertical Depth Gradient in an Herbivorous Fish

Males are expected to adjust testes investment according to the varying level of sperm competition that they experience. Spatial and temporal variation in population density likely influences sperm competition. In herbivorous aquatic organisms, densities often decrease along a vertical depth gradient, because their food is photosynthetic and thus becomes less abundant in deeper regions where less light penetrates. This decrease should be dramatic on a steep slope, which allows testing of the association between density and testes investment at the within‐population level. We tested the effect in the socially monogamous herbivorous cichlid fish Variabilichromis moorii living on a steep slope in Lake Tanganyika. We examined competitor density and food abundance as ecological factors, territory defense behaviors and phenotypic traits (testes investment and somatic investment), and compared them between shallow (4–6 m depth) and deep habitats (10–13 m depth) separated by several dozen meters. We found that food availability drastically decreased with increasing depth and that V . moorii was much more abundant in shallower habitats. Males in shallower habitats were in better physical condition (based on fat and liver mass) despite experiencing greater costs in terms of territory defense. Testes investment differed in areas with different competitor density and food abundance along a vertical depth gradient, but competitor density was the most explanatory factor of the difference. This suggests that this herbivorous fish would change testes investment in response to population density.     

The feeding habits of a group of herbivorous rock-dwelling cichlid fishes (Cichlidae: Perciformes) from Lake Malawi,Africa

Synopsis The feeding; habits of a group of tropical herbivorous rock-dwelling cichlid fishes from Lake Malawi, Africa, are investigated using stomach content analyses. The various species fed selectively on the periphyton of the rocky shores. Blue-green alga of the genus Calothrix was the most common item ingested by the group. Diatoms (Chrysophyta) also were abundant food items. Discriminant analysis showed that dietary items were good variables to identify species. Interspecific dietary differences showed a continuum from those species feeding primarily on Calothrix to those feeding primarily on diatoms. Algal resources exhibit distinct patterns of spatial variation. Diet was correlated with foraging behavior and trophic morphology. Interspecific differences in diet could possibly facilitate ecological coexistence among various species. Such coexistence would contribute to the maintenance of the high diversity fish faunas characteristic of the Great Rift Lakes of Africa.     

Feeding selectivity in relation to territory size in a herbivorous reef fish

Summary Territory size in the herbivorous reef fish, Parma victoriae, (Pisces: Pomacentridae) is primarily determined by local density patterns and is independent of changes in food levels (Norman and Jones 1984). However, while individual readily expand territories in response to having their neighbours removed, they appear not to increase total food consumption. Individuals with large territories may benefit from improved food quality. Here we test the hypothesis that expanded territories increase access to preferred food types.A preliminary comparison of the relative proportions of algal species consumed and those present in the field indicated that feeding was highly selective. Strong feeding preferences were indicated by a field cafeteria experiment, in which algal species were presented to individuals in equal amounts. Selectivity indices calculated from diet and availability measures were positively correlated with the feeding preference rankings.Individuals examined from two territory size categories (<10 m² and > 10 m²) did not exhibit significant differences in feeding prefernces. However, they differed greatly in terms of the makeup of their diets. Higher ranking algae such as Champia zostricola and Rhodoglossum sp. were consumed in greater proportions on large territories. Algal abundances within territories also differed for the two groups. Some of the less abundant, high ranking food species were found in greater quantities per unit area on large territories. The differences in diet were not solely explained by changes in availability. The appropriate comparison showed that whereas individuals on large territories fed in a highly selective manner, those on small territories consumed algae in proportion to their availability. A neighbour removal experiment supported the hypothesis that territory expansion results in an increase in the consumption of preferred algal species.     

Diet disparity among sympatric herbivorous cichlids in the same ecomorphs in Lake Tanganyika: amplicon pyrosequences on algal farms and stomach contents

Background

Lake Tanganyika, an ancient lake in the Great Rift Valley, is famous for the adaptive radiation of cichlids. Five tribes of the Cichlidae family have acquired herbivory, with five ecomorphs: grazers, browsers, scrapers, biters and scoopers. Sixteen species of the herbivorous cichlids coexist on a rocky littoral slope in the lake. Seven of them individually defend feeding territories against intruding herbivores to establish algal farms. We collected epiphyton from these territories at various depths and also gathered fish specimens. Algal and cyanobacteria community structures were analysed using the amplicon-metagenomic method.

Results

Based on 454-pyrosequencing of SSU rRNA gene sequences, we identified 300 phototrophic taxa, including 197 cyanobacteria, 57 bacillariophytes, and 31 chlorophytes. Algal farms differed significantly in their composition among cichlid species, even in the same ecomorph, due in part to their habitat-depth segregation. The algal species composition of the stomach contents and algal farms of each species differed, suggesting that cichlids selectively harvest their farms. The stomach contents were highly diverse, even between species in the same tribe, in the same feeding ecomorph.

Conclusions

In this study, the amplicon-metagenomic approach revealed food niche separation based on habitat-depth segregation among coexisting herbivorous cichlids in the same ecomorphs in Lake Tanganyika.

     

Variation of territory size and defense behavior in breeding pairs of the endemic Lake Tanganyika cichlid fish Variabilichromis moorii

Lake Tanganyika harbors the ecologically, morphologically, and behaviorally most diverse species flock of cichlid fishes. It is comprised by substrate breeding and mouthbrooding species, most of which live in littoral habitats. Species communities are characterized by complex behavioral and trophic interactions, resulting in a dense pattern of partially overlapping territories, depending on the degree of ecological distinctness. We studied territorial behavior of breeding pairs in a substrate breeding species, with respect to territory size and defense behavior. The study species Variabilichromis moorii belongs to the tribe Lamprologini, the most species rich tribe of cichlids in Lake Tanganyika. Our study shows that breeding pairs of V. moorii can have highly complex territories, in which both parents hold separate sub-territories which are shifted slightly according to the movements of the fry, but the outer borders are conjointly defended. The size of the total defended territory varied from <1 to almost 4 m², averaging at about 2 m². Depending on presence of competitors or fry-predators evoking agonistic interactions, the territory size varied quite substantially over the day. Attack rates and size of the defended area decreased with water depth. Agonistic behavior was observed toward heterospecifics as well as conspecifics, with heterospecific attacks mostly concerning territorial neighbors and potential fry-predators in about equal frequencies. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Guest editors: T. Wilke, R. V?in?l? & F. Riedel Patterns and Processes of Speciation in Ancient Lakes: Proceedings of the Fourth Symposium on Speciation in Ancient Lakes, Berlin, Germany, September 4–8, 2006     

Trophic Niche in a Raptor Species: The Relationship between Diet Diversity,Habitat Diversity and Territory Quality

Recent research reports that many populations of species showing a wide trophic niche (generalists) are made up of both generalist individuals and individuals with a narrow trophic niche (specialists), suggesting trophic specializations at an individual level. If true, foraging strategies should be associated with individual quality and fitness. Optimal foraging theory predicts that individuals will select the most favourable habitats for feeding. In addition, the “landscape heterogeneity hypothesis” predicts a higher number of species in more diverse landscapes. Thus, it can be predicted that individuals with a wider realized trophic niche should have foraging territories with greater habitat diversity, suggesting that foraging strategies, territory quality and habitat diversity are inter-correlated. This was tested for a population of common kestrels Falco tinnunculus. Diet diversity, territory occupancy (as a measure of territory quality) and habitat diversity of territories were measured over an 8-year period. Our results show that: 1) territory quality was quadratically correlated with habitat diversity, with the best territories being the least and most diverse; 2) diet diversity was not correlated with territory quality; and 3) diet diversity was negatively correlated with landscape heterogeneity. Our study suggests that niche generalist foraging strategies are based on an active search for different prey species within or between habitats rather than on the selection of territories with high habitat diversity.     

Pale and dark dichromatism related to microhabitats in a herbivorous tanganyikan cichlid fish,Telmatochromis temporalis

Dichromatism and habitat utilization of a small herbivorous cichlid fish,Telmatochromis temporalis, were studied in a littoral rocky area of Lake Tanganyika. Individuals of both sexes had either pale or dark body coloration that did not change for long periods. Both sexes defended territories from consexuals, and a male had a harem of several females. Pale fish had territories in exposed areas and dark fish in shaded ones. Larger fish were dominant over smaller consexuals and occupied well-illuminated areas. When pale and dark fish were kept in areas with shaded and lighted backgrounds, respectively, they changed body colour within a few weeks. When pale fish were in well-illuminated areas and dark ones in shaded areas, they were cryptic. BecauseTelmatochromis temporalis are very vulnerable prey in coastal areas, dichromatism may function as antipredator camouflage.     

Concentration of toxic metals and potential risk assessment in edible fishes from Congo River in urbanized area of Kinshasa,DR Congo

Abstract

The objective of this research is to determine the concentration and potential health risks of toxic metals in six most consumed freshwater fish species (Auchenoglanis occidentalis, Bagrus ubangensis, Citharinus gibbosus, Distichondus fasciolatus, Marcusenius moorii, and Schilbe mystus). A total of 126 samples of these fish species were collected from seven sites of Congo River in the vicinity of Kinshasa City, Democratic Republic of the Congo. Toxic metals (Cr, Co, Ni, Zn, Se, Ag, Cd, Sn, Sb, and Pb) levels in fish muscle tissues were determined using Inductively Coupled Plasma Mass Spectrometry whereas Hg analysis was carried out using atomic absorption spectrophotometry. The maximum metal concentration (mg kg^?1 wet weight) of Zn (59.72), Sb (5.98), and Pb (4.96) was detected in muscle tissues of M. moorii, C. gibbosus, and D. fasciolatus, respectively. Pb, Zn, and Hg values in many fish samples exceeded the permissible levels set by Food and Agriculture Organization and World Health Organization for human consumption. Given the concentrations of Pb in all fish species and Zn in M. moorii, human health implications are likely to occur on continuous consumption. Addition, the evaluation of potential health risks for fish consumers indicates that there may or not be adverse effects.     

Resource control by territorial male cichlid fish in Lake Malawi

1. The rocky habitat communities of Lake Malawi contain a high diversity of ecologically similar, predominantly herbivorous, cichlid fish species. How so many species can coexist is still unknown.
2. Adult males of the majority of these species hold permanent territories which form dense multispecies mosaics across the shores.
3. The study examined six coexisting species of cichlids from the rocky habitat where males are known to control access to the algal food resources within their territorial space. These included four sibling species of the Pseudotropheus ( Tropheops ) species complex, which are specialized epilithic algal herbivores, and two sibling species of the trophically more generalist P . ( Maylandia ) species complex which feed upon both epilithic algae and plankton.
4. The hypotheses that both intraspecific and interspecific territoriality occur in these communities and that interspecific aggression initiated by territorial males is preferentially directed at species with the most similar diets to their own were tested.
5. It was found that males preferred to exclude conspecific intruders, while they tolerated intruders with different diets to their own. Also, male P . ( Tropheops ) preferentially excluded similar heterospecific dietary specialists, while male P. ( Maylandia ) tolerated similar heterospecific dietary generalists.
6. Based on this study, it is proposed that interspecific territoriality may be reducing competition between species with different diets and promoting their coexistence, while it may be enhancing competition between species with the same diet. Furthermore, it suggests that ecological generalism may be reducing the intensity of interspecific competition, while specializations for the same resources may be increasing competition.     

Grazing rates of crustacean zooplankton communities on intact phytoplankton communities in Canadian Subarctic lakes and ponds

Quantitative genetic studies in natural populations are of growing interest to speciation research since divergence is often believed to arise through micro-evolutionary change, caused by natural selection on functional morphological traits. The species flock of cichlid fishes in Africa’s oldest lake, Lake Tanganyika, offers a rare opportunity to study this process. Using the cichlid species Tropheus moorii, we assessed the potential for microevolution in a set of morphological traits by estimating their quantitative genetic basis of variation. Two approaches were employed: (1) estimation of trait heritabilities (h ²) in situ from a sample of wild caught fish, and (2) estimation of h ² from first generation offspring produced in a semi-natural breeding experiment. In both cases, microsatellite data were used to infer pedigree structure among the sampled individuals and estimates of h ² were made using an animal model approach. Although power was limited by the pedigree structures estimated (particularly in the wild caught sample), we nonetheless demonstrate the presence of significant additive genetic variance for aspects of morphology that, in the cichlid species Tropheus moorii, are expected to be functionally and ecologically important, and therefore likely targets of natural selection. We hypothesize that traits showing significant additive genetic variance, such as the mouth position have most likely played a key role in the adaptive evolution of the cichlid fish Tropheus moorii.     

Damselfish territories as a refuge for macroalgae on coral reefs

Herbivory is widely accepted as a key process determining the benthic community structure and resilience of coral reefs. Recent studies have mostly focused on the importance of roving herbivorous fishes in ecosystem processes. Here, we examine the role of territorial damselfish in shaping patterns of macroalgal distribution based on benthic surveys and macroalgal bioassays. The territory composition and effect of resident damselfish on the removal of Sargassum bioassays were quantified for six species of damselfish on Lizard Island, a mid-shelf reef in the northern Great Barrier Reef (GBR). The functional composition of algal communities within territories varied markedly among species. The territories of four species (Dischistodus perspicillatus, Dischistodus pseudochrysopoecilus, Plectroglyphidodon lacrymatus, and Stegastes nigricans) were characterized by algal turfs, while the territories of two species (Dischistodus prosopotaenia and Hemiglyphidodon plagiometopon) were characterized by foliose and leathery brown macroalgae. Sargassum, a generally rare alga on mid-shelf reefs, was a particularly common alga within D. prosopotaenia territories on the leeward side of the island but absent within their territories on the windward side of the island. D. prosopotaenia was the only species to retain the transplanted Sargassum, with only a minimal reduction in Sargassum biomass (1.1%) being recorded within their territories at both leeward and windward sites over a 24-h period. In contrast, reductions in Sargassum biomass were high in areas adjacent to D. prosopotaenia territories (83.8%), and within and adjacent to the territories of the five remaining damselfish species (76.2–92.5%). Overall, only one of the six damselfish species provided a refuge for leathery brown macroalgae and may facilitate the development of this macroalgae on mid-shelf reefs of the GBR.