Evolution of brain–body allometry in Lake Tanganyika cichlids

Brain size is strongly associated with body size in all vertebrates. This relationship has been hypothesized to be an important constraint on adaptive brain size evolution. The essential assumption behind this idea is that static (i.e., within species) brain–body allometry has low ability to evolve. However, recent studies have reported mixed support for this view. Here, we examine brain–body static allometry in Lake Tanganyika cichlids using a phylogenetic comparative framework. We found considerable variation in the static allometric intercept, which explained the majority of variation in absolute and relative brain size. In contrast, the slope of the brain–body static allometry had relatively low variation, which explained less variation in absolute and relative brain size compared to the intercept and body size. Further examination of the tempo and mode of evolution of static allometric parameters confirmed these observations. Moreover, the estimated evolutionary parameters indicate that the limited observed variation in the static allometric slope could be a result of strong stabilizing selection. Overall, our findings suggest that the brain–body static allometric slope may represent an evolutionary constraint in Lake Tanganyika cichlids.     

Mating systems of two midwater-spawning cichlids,Cyprichromis microlepidotus andParacyprichromis brieni,in Lake Tanganyika

The breeding habits of 2 maternal mouthbrooding cichlids,Cyprichromis microlepidotus andParacyprichromis brieni, were investigated in Lake Tanganyika. Although spawning on the substrate in the male's mating territory is prevalent in maternal mouthbrooders, bothC. microlepidotus andP. brieni spawned in the water column. MaleC. microlepidotus established their mating territories in the open water column, while maleP. brieni did so around fixed spawning sites near a vertical rock surface. In both species, females visited male mating territories, departing soon after spawning and collecting the eggs. Sneaking, which was observed only inP. brieni, may be attributed to the presence of refuges for sneakers in this species. FemaleC. microlepidotus deposited their entire clutch of about 9 eggs in one male territory. In contrast, femaleP. brieni divided their clutch of about 11 eggs among several males. After the final egg-release, femaleC. microlepidotus repeatedly approached their mate, with the mouth near the abdomen of the latter (nuzzling), but femaleP. brieni often departed without nuzzling. Males may eject sperm during nuzzling to fertilize eggs inside the female's mouth. However, maleP. brieni is also known to eject sperm near spawning females when the females are not nuzzling. Such behavior seems to be a male countermeasure against female mate infidelity, because males could not ensure paternity of eggs by ejecting sperm only during female nuzzling.     

Helping opportunities and space segregation in cooperatively breeding cichlids

Studies of cooperative breeding have largely ignored the roleof conflict among helpers and how it shapes group dynamics andhelping behavior. In the present study, performing laboratoryexperiments with cooperatively breeding cichlids from Lake Tanganyika,we show that secondary group members (potential helpers) occupyhome ranges within the group territory and may be aggressiveto one another. Experimental removal of secondary group membersallowed the individual next in rank to move closer to the removedindividual's home range. In the field, dominant secondary groupmembers stayed closer to the brood chamber than did subordinategroup members of similar size, and proximity to the brood chamberwas related to the length of time spent inside. We suggest thatspace segregation and competition among secondary group membersis common in these cichlids, and may limit the opportunitiesto provide help.     

Lunar synchronization of spawning in cichlid fishes of the tribe Lamprologini in Lake Tanganyika

Lunar synchronization of spawning was investigated for eight substrate brooding cichlid fishes belonging to the tribe Lamprologini. Their spawning activities all peaked during the second quarter of the lunar cycle. Comparison between their breeding styles and the degrees of spawning synchronization suggested that the nocturnal guarding-efficiency of eggs, especially exposed ones, is improved by the maximal amount of moonlight during full moon and, in species whose young leave the breeding sites about 2 weeks after spawning, the survival of dispersing young is enhanced by the darkness of moonless nights.     

Intestinal coiling pattern in the epilithic algal-feeding cichlids (Pisces, Teleostei) of Lake Tanganyika, and its phylogenetic significance

This paper suggests that, on the basis of ontogenetic change in intestinal coiling pattern, four types and four subtypes could be recognized within 20 species of epilithic algal-feeding cichlids in Lake Tanganyika. The phylogenetically useful anatomical features characterizing each type are as follows. Petrochromis type: torsion of the intestinal tract generally occurs at Stage 5 Intersecting Point (IP), and the haplochromine loop and a two-loop coiling system are observable. Cyathopharynx type: the value of IP becomes 7 before torsion occurs, and the coiling system changes qualitatively from one-loop to two-loop types. Asprotilapia type: torsion occurs at Stage 5 IP, and a one-loop coiling system is retained throughout ontogeny. Telmatochromis type: no torsion occurs, and a one-loop coiling system is retained throughout ontogeny. Within the Petrochromis type, each of the four subtypes is characterized by the following features. Petrochromis subtype: the rostrally directed haplochromine loop has its rostral part situated to the right side of the stomach in the adult stage. Pseudosimochromis subtype: a circular haplochromine loop is present. Eretmodus subtype: the haplochromine loop tends to be prolonged and bends through 90 or 180 further counter-clockwise. Tropheus subtype: torsion occurs at Stage 7 IP.     

Phytoplankton pigments and community composition in Lake Tanganyika

1. A 2‐year (2002–2003) survey of chlorophyll and carotenoid pigments is reported for two off‐shore stations of Lake Tanganyika, Kigoma (Tanzania) and Mpulungu (Zambia), and from three cruises between those sites. Chlorophyll a concentrations were low (0.3–3.4 mg m^?3) and average chlorophyll a integrated through the 100 m water column were similar for both stations and years (36.4–41.3 mg m^?2). Most pigments were located in the 0–60 m layer and decreased sharply downward. Chlorophyll a degradation products (phaeophytins and phaeophorbides) were detected at 100 m depth, whereas carotenoids became undetectable. Temporal and seasonal variation of the vertical distribution of pigments was high. 2. The biomass of phytoplankton groups was calculated from marker pigment concentrations over the 0–100 m water column using the CHEMTAX software. On average for the study period, chlorophytes dominated in the northern station, followed by cyanobacteria T1 (type 1, or Synechococcus pigment type), whereas cyanobacteria T1 dominated in the south. Cyanobacteria T2 (type 2, containing echinenone), presumably corresponding to filamentous taxa, were detected in the rainy season. Diatoms (and chrysophytes) developed better in the dry season conditions, with a deep mixed layer and increased nutrient availability. Very large variation in the vertical distribution of algal groups was observed. 3. Our observations on phytoplankton composition are broadly consistent with those from previous studies. Our pigment data provide evidence for the lake‐wide importance of picocyanobacteria and high interannual variation and spatial heterogeneity of phytoplankton in Lake Tanganyika, which may render difficult assessment of long‐term changes in phytoplankton driven by climate change.     

Evolution of the tribe Tropheini from Lake Tanganyika: synchronized explosive speciation producing multiple evolutionary parallelism

One of the most surprising outcomes of recent molecular studies on cichlid fishes of the three Great East African Lakes Victoria, Malawi and Tanganyika, was the stunning rapidity of speciation and cladogenesis at early stages of adaptive radiation. Despite their rapid pace, speciation events were so far intuitively assumed to proceed in a bifurcating and tree-like fashion, even if they could not be resolved by gene phylogenies due to a lack of resolution. On the basis of phylogenetic analyses of the Tropheini, a lineage of endemic rock-dwelling cichlid fishes from Lake Tanganyika, we suggest a pathway of explosive speciation that accounts for a non-bifurcating manner of cladogenesis. This pattern is likely to be the result of the contemporaneous origin of a multitude of founder populations in geographically isolated rock habitats among which gene flow was interrupted simultaneously by a major change of the lake habitat in the form of a rapid rise of the lake level. As a consequence, all new species arising from that vicariance event must exhibit almost equal genetic distances to each other, within the scope of genetic diversity of the founder population(s), even if the actual processes of subsequent speciation and eco-morphological diversification followed independent routes. Our phylogeny also suggests a high frequency of parallel evolution of equivalent trophic specialization in the Tropheini. This phenomenon seems to be an inherent feature of this pathway of speciation, due to the action of similar selective forces on the same set of species colonizing isolated habitats of the same type. Explosive speciation via synchronization of genetic divergence triggered by rapid environmental changes seems to be particularly likely to occur at advanced stages of adaptive radiation, when species are already adapted to particular habitats and have a reduced ability for dispersal.     

An initial measurement of photosynthetic productivity in Lake Tanganyika

Measurements of ¹⁴C uptake and oxygen evolution by phytoplankton in Lake Tanganyika provide the first direct estimates of the lake's primary productivity. These measurements at the expected nadir of the annual range suggest that Lake Tanganyika is as productive as other large, deep tropical lakes.     

Sedimentary diatom assemblages in the northern part of Lake Tanganyika

Surface sediment diatoms from the northern part of Lake Tanganyika were analysed with several ordination and classification techniques. Three sample groups characterized by four diatom assemblages were recognized. The first sample group occurs in a rather shallow area near the delta of the Rusizi River on a coarse-grained sand substrate. The second and most important sample group occurs on the silty bottom of an underwater depression situated off the delta of the Rusizi River in the north-eastern part of Lake Tanganyika; its diatom composition has a great affinity to the Rusizi River diatom population. The third sample group contains all the other samples and covers an area corresponding to the west-, east- and southward continuation of the area of the second group: its diatom composition is also related to the Rusizi River diatom population, but in a less pronounced manner.     

Obligate feeding of cichlid eggs by Caecomastacembelus zebratus in Lake Tanganyika

This study examined food habits of four common species of spiny eels (Mastacembelidae) in Lake Tanganyika to find if they eat cichlid eggs and larvae. Diets of Caecomastacembelus zebratus consisted nearly exclusively of eggs of substratum-brooding cichlids, whereas two other species fed on eggs only facultatively and one species never did. More than half individuals of C. zebratus had empty guts, although the mean weight of their gut contents was highest. This spiny eel ate many eggs in the first and second quarters of the lunar cycle when the spawning of substratum-brooding cichlids was active, while it nearly ceased eating in the latter half of the cycle. It is suggested that this fish is an obligate egg-eater and one of the predators that cause the high mortality of early cichlid broods.     

Alloparental care between catfishes in Lake Tanganyika

A bagrid catfish Auchenoglanis occidentalis cared for its brood, deposited within an accumulation of shells and gravel in the centre of a large saucer-like depression, for up to 2 weeks in Lake Tanganyika. Adults of another catfish Dinotopterus cunningtoni (Clariidae) persistently came to A. occidentalis nests near days of host spawning. Eggs of D. cunningtoni were found in the host nests on the day and within 2 days of host spawning. During and for a few days after the end of host brooding, associate species' young of much older ages than the host brood were often found in the nests together with those of a similar age to the host young. Associate young were rarely found in nests before spawning and in unused nests. These findings suggested that this brood-mixing has two origins: egg dumping by associate adults and voluntary intrusion of large associate young into host nests. Possible benefits to the associate species are to take advantage of nest preparation and parental behaviour of the host species and to feed on the host brood.     

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.     

Paternal mouthbrooding bagrid catfishes in Lake Tanganyika

 This article gives the first information about paternal mouthbrooding in bagrid catfishes, based on analysis of specimens collected while SCUBA diving in Lake Tanganyika, Africa. Lophiobagrus aquilus and L. cyclurus, endemic to the lake, were nocturnally active, and in the daytime were found concealed singly or in pairs beneath rocks. Among 39 L. aquilus and 52 L. cyclurus collected, 8 and 1 solitary males, respectively, were incubating eggs or young in the buccal cavity. Paired males and females and solitary females never took care of the offspring. In both species, females of pairs showed much higher gonadosomatic index values and had larger oocytes in the ovaries than did solitary females. This fact suggests that pair formation takes place at a time near the gonadal maturation of the female and that the pair separate after spawning. We tentatively propose an evolutionary transition route of care patterns in bagrid catfishes. Received: January 25, 2002 / Revised: April 13, 2002 / Accepted: April 17, 2002     

Differential response of phytoplankton to additions of nitrogen, phosphorus and iron in Lake Tanganyika

1. In order to evaluate limitation of different phytoplankton groups by inorganic nutrients, multiple nutrient enrichment bioassays using the addition of iron (Fe) and the combined addition of nitrogen and phosphorus (NP) were carried out in the north and the south of Lake Tanganyika during the rainy and dry seasons in 2003 and 2004. 2. Nutrient additions resulted in an increase in phytoplankton growth rate relative to control treatments in all experiments. HPLC pigment data and epifluorescence microscopy counts indicated differential stimulation of the dominant phytoplankton groups. Iron additions mainly stimulated prokaryotic picophytoplankton, while enrichments with nitrogen and phosphorus stimulated green algae and in some cases diatoms. Extended incubation (3 days) indicated co‐limitation of Fe and NP, in particular for picocyanobacteria.     

Biparental mouthbrooding of the catfish Phyllonemus filinemus in Lake Tanganyika

This article reports biparental mouthbrooding of the bagrid catfish Phyllonemus filinemus in Lake Tanganyika, based on analysis of specimens collected during SCUBA diving. This catfish was nocturnally active, and in the daytime it was concealed singly or in pairs beneath rocks. Within a breeding pair, the male or female alone incubated all the brood in the mouth until the offspring attained 12 mm or so in total length, but thereafter joint mouthbrooding or guarding by both parents took place. Most females of nonbrooding pairs showed high values of gonadosomatic index (GSI), whereas all females of brooding pairs and most single females showed low GSI values. This fact indicated that a pair is formed at a time near the gonadal maturation of the female and separates after the brood is reared. No significant difference in body condition among parents of different reproductive states was observed, which suggested that their condition does not deteriorate markedly as the result of foraging by an off-duty parent. Received: September 16, 2000 / Revised: November 18, 2000 / Accepted: January 23, 2001     

Seasonal patterns of sediment loading and benthic invertebrate community dynamics in Lake Tanganyika, Africa

1. This study investigated the spatiotemporal dynamics of benthic invertebrate communities located close to the mouths of the Kalambo and Lunzua Rivers, at the southern end of Lake Tanganyika, and subject to varying degrees of sediment load. 2. Metrics of exposure to riverine sediments were associated inversely with abundance and diversity of benthic organisms at sampling locations adjacent to both river mouths, indicating a major effect of riverine sediments on the structure of near‐shore benthic communities. 3. A lower abundance of benthos was found at the mouth of the Lunzua River, which exports significantly higher sediment loads than the Kalambo. 4. Seasonal cycles of abundance and diversity in the benthos varied with distance from river mouths. This has important implications for monitoring programmes investigating benthic communities close to the mouths of rivers. 5. Our findings suggest that recent increases in the sediment loading of Lake Tanganyika, owing to anthropogenic catchment disturbance, impact significantly on biological diversity throughout the lake littoral, which provides habitat for the majority of species in the lake.     

The impact of the River Ntahangwa, the most polluted Burundian affluent of Lake Tanganyika, on the water quality of the lake

Sources of pollution at the north end of Lake Tanganyika are concentrated around Bujumbura, Burundi. The impact of River Ntahangwa on the water quality of this part of Lake Tanganyika was compared with that of an unpolluted river, the Mugere. A shallow bay, not influenced by a river, was used as a reference station. Chemical analyses were carried out at four week intervals during nearly a year. Samples were taken at different distances from the shore and at different depths. River impact was detected up to 100 m from both river mouths. Even at 50 m from the mouth of the polluted River Ntahangwa, no important decrease of dissolved oxygen was detected. Chlorophyll-a maxima were found 100 m from the Ntahangwa mouth, but this was also the case 100 m from shore in the unpolluted bay. Lake water at 60 m depth was saturated with oxygen during the dry season, as it used to be in 1956. The rivers studied descent initially within the lake due to greater sediment load and lower water temperature. Nevertheless, it appears that these rivers mix completely in the lake within 100–300 m from the river mouths. The direct effects from organic pollution of the Ntahangwa on the lake seem to be rather negligible. Appropriate anti-erosion measures should reduce massive sediment discharge into the lake.     

Ecology and conservation status of endemic freshwater crabs in Lake Tanganyika, Africa

Sedimentation resulting from riparian deforestation has a wide range of detrimental effects on aquatic biodiversity, but predicting the full consequences of such disturbances requires an understanding of the ecosystem’s key functional components. We investigated the ecology and response to sedimentation of the diverse, endemic freshwater crabs of Lake Tanganyika, which may occupy important positions in littoral foodwebs. Our surveys revealed crab distribution patterns to be patchy, and that crabs can be locally abundant (0–28 individuals m⁻²). Crab densities decreased with depth and the dry mass of crab assemblages ranged from 0.0 to 117.7 g m⁻². Comparisons among sites revealed significant effects of sedimentation on crab assemblage evenness, but provided no evidence that sedimentation has altered densities, incidence or species richness. The resilience of crabs to sedimentation might be related to their intraspecific dietary breadth. Stable isotope data (δ¹³C and δ¹⁵N) from crabs and their potential food resources indicated differences in trophic roles among endemic crab species. Overall, crabs occupy higher trophic positions than most other invertebrates, and they draw upon both benthic and planktonic energy pathways. The high biomass and top-predator status of some crab species suggests the potential for cascading effects on organisms lower in the food web.     

Neolamprologus cancellatus, a new cichlid fish from Lake Tanganyika, Africa

Neolamprologus cancellatus, a new cichlid species, is described on the basis of eight specimens from the Zambian coast of Lake Tanganyika. The new species is characterized by a subtruncate caudal fin and a body that is slender (depth 22.3–25.2% in standard length) and easily distinguishable from its congeners by having 7–8 anal fin spines, 34–37 scales in longitudinal line, 33 total vertebrae, and a gridlike body pattern on a pale brownish body color. This species is only known to inhabit the shallow (2–7 m depth), rocky bottom of Wonzye Point in the southern part of the lake.