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     

Farming-out of offspring is a predominantly male tactics in a biparental mouthbrooding cichlid <Emphasis Type="BoldItalic">Perrisodus mircolepis</Emphasis>

Synopsis Intraspecific brood-mixing of a Tanganyikan biparental cichlid fish Perissodus microlepis has been attributed to farming-out, a behaviour in which a parent transport some or all of their free-swimming young in the mouth and put them under the care of other guarding parents. We conducted observations under natural conditions to understand how often farming-out occurs and which sex is the primary performer. At most brooding sites, young in a school entirely disappeared before attaining to the size of independence. At many of these sites, farming-out and its precursory behaviours by males preceded disappearance of their young. Females never performed farming-out. We proposed that farming-out by males is an alternative tactics to brood desertion to evade paternal roles for remating.     

Contrasting geographic structure in evolutionarily divergent Lake Tanganyika catfishes

Geographic isolation is suggested to be among the most important processes in the generation of cichlid fish diversity in East Africa's Great Lakes, both through isolation by distance and fluctuating connectivity caused by changing lake levels. However, even broad scale phylogeographic patterns are currently unknown in many non‐cichlid littoral taxa from these systems. To begin to address this, we generated restriction‐site‐associated DNA sequence (RADseq) data to investigate phylogeographic structure throughout Lake Tanganyika (LT) in two broadly sympatric rocky shore catfish species from independent evolutionary radiations with differing behaviors: the mouthbrooding claroteine, Lophiobagrus cyclurus, and the brood‐parasite mochokid, Synodontis multipunctatus. Our results indicated contrasting patterns between these species, with strong lake‐wide phylogeographic signal in L. cyclurus including a deep divergence between the northern and southern lake basins. Further structuring of these populations was observed across a heterogeneous habitat over much smaller distances. Strong population growth was observed in L. cyclurus sampled from shallow shorelines, suggesting population growth associated with the colonization of new habitats following lake‐level rises. Conversely, S. multipunctatus, which occupies a broader depth range, showed little phylogeographic structure and lower rates of population growth. Our findings suggest that isolation by distance and/or habitat barriers may play a role in the divergence of non‐cichlid fishes in LT, but this effect varies by species.     

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     

A new diminutive genus and species of catfish from Lake Tanganyika (Siluriformes: Clariidae)

The examination of material representing one of Lake Tanganyika's six previously recognized endemic catfish lineages, has revealed the presence of an additional genus of clariid, described here as Pseudotanganikallabes new genus. This genus is represented by a single species, Pseudotanganikallabes prognatha sp. nov., which is distinguished from all other clariids by its lack of an infraorbital series, the presence of multiple osseous connections between the swim bladder capsules and elements of the neurocranium, the absence of an ethmoid notch, the presence of a very large, egg‐shaped occipital fontanelle and the extension of the lower lip beyond the margin of the upper jaw. A combination of additional external and molecular characters serves to further distinguish this taxon from all currently recognized clariid species. Phylogenetic analysis of mitochondrial (cytb) and nuclear (18S‐ITS1‐5.8S‐ITS2‐28S) sequence data supports the creation of a new genus for this species, as it appears to represent an independent, monophyletic lineage within the family Clariidae.     

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.     

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.     

Diatoms from surface sediments of the northern part of Lake Tanganyika

227 Diatom taxa were observed in the surface sediments of the northern part of Lake Tanganyika, including 1 new to science: Amphora tanganyikae. The diatom community of these sediments is mainly composed of benthic organisms while planktonic diatoms are rather rare. Many brackish-water and a few marine organisms were observed. Cosmopolitan organisms (77.1%) dominate the diatom flora but tropical, tropical African and African taxa are also well represented (22.9%)Deceased.Deceased.     

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.     

Mating system and kin relationship between adults and young in the shell-brooding cichlid fish <Emphasis Type="Italic">Neolamprologus meeli</Emphasis> in Lake Tanganyika

Mating system and parental behavior of ten monogamous pairs and two polygynous groups of the Tanganyikan cichlid Neolamprologus meeli were observed in their natural habitat. The home ranges of males and females overlapped with each other. Most groups included one to six young. Paternal and maternal relationships were determined for 22 young from DNA microsatellite markers. Three types of kinship were found: (I) kinship to both the male and female; (II) kinship to females only; and (III) non-kinship to both sexes. In the groups with type II young, step-fathering or sneaking may have occurred. Type III young were larger than type I, suggesting that the former were of sufficient size to leave their birth nest and settle in the territories of foster parents. Both males and females drove out potential predators of young (including three species of Lepidiolamprologus) as a parental behavior. Adults with type III young attacked approaching predators with as much frequency as those with type I young only, indicating that they provided alloparental care. Adults and young swam together, but, a significant difference existed in the frequencies of interactions between adults versus kin young and adults versus non-kin young. The results suggest that both adults and young recognized kin. Electronic Publication     

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.     

A game theoretical approach to conspecific brood parasitism

We constructed a game theoretical model to predict optimal patternsof egg laying in systems where individuals lay in the nestsof others as well as in their own nests. We show that decreasingthe effect of position within an egg-laying sequence on theworth of an egg should lead to reduced parasitism. Indeed,parasitism can only flourish if the worth of an egg to its biological parent declines with the total number of eggs laid in that nest.Further, we found that increasing the intrinsic costs of eggproduction should lead to an increased propensity for conspecificbrood parasitism. The model also predicts that variation inhosts' ability to reject parasitic eggs has little effect on parasitism until this ability is well developed.     

Tactic for bower acquisition by male cichlids,Cyathopharynx furcifer,in Lake Tanganyika

A tactic for bower acquisition by male cichlids,Cyathopharynx furcifer, was studied in Lake Tanganyika. Males held crater-shaped bowers of sand within their territories, which were located adjacent to each other. Females visited bowers to spawn eggs, subsequently brooding the eggs in their mouths. Some individuals held territories without bowers near those of males with bowers. Although only the males with bowers succeeded in reproduction, they were apparently exhausted by bower maintenance and reproduction, and often deserted their bowers. These were soon occupied by males which had held nearby territories without bowers, such males then engaging in reproductive activity. Although the construction of new bowers by males was observed, such was unsuccessful, probably because too much energy was required and/or the optimal space for bowers was limited. Direct aggression by males without bowers against bower-holding males to deprive the bowers was not observed, possibly due to the high costs of such competition. Therefore, waiting for bower desertion by other males may be the best bower acquisition tactic for males without bowers.     

Social parasitism by workers in queenless and queenright Apis cerana colonies

We examined worker reproduction in queenless and queenright Apis cerana colonies to determine if they are parasitized by workers from other nests. The results demonstrate that 2-6% of workers in queenright colonies are from another nest (non-natal), but these workers are not statistically more likely to have activated ovaries than natal workers, and are therefore unlikely to be active parasites. However, in queenless colonies we found a significant difference between the proportion of non-natal (72.7%) and natal (36.3%) workers with activated ovaries. Non-natal workers also had significantly higher reproductive success than natal workers: 1.8% of workers were non-natal, but these laid 5.2% of the eggs and produced 5.5% of the pupae. Unlike A. florea, the proportion of non-natal workers does not increase in queenless nests.     

Nest destruction elicits indiscriminate con‐ versus heterospecific brood parasitism in a captive bird

Following nest destruction, the laying of physiologically committed eggs (eggs that are ovulated, yolked, and making their way through the oviduct) in the nests of other birds is considered a viable pathway for the evolution of obligate interspecific brood parasitism. While intraspecific brood parasitism in response to nest predation has been experimentally demonstrated, this pathway has yet to be evaluated in an interspecific context. We studied patterns of egg laying following experimental nest destruction in captive zebra finches, Taeniopygia guttata, a frequent intraspecific brood parasite. We found that zebra finches laid physiologically committed eggs indiscriminately between nests containing conspecific eggs and nests containing heterospecific eggs (of Bengalese finches, Lonchura striata vars. domestica), despite the con‐ and heterospecific eggs differing in both size and coloration. This is the first experimental evidence that nest destruction may provide a pathway for the evolution of interspecific brood parasitism in birds.     

A simple model of the eco-hydrodynamics of the epilimnion of Lake Tanganyika

1. The ecosystem response of Lake Tanganyika was studied using a four-component, nutrient–phytoplankton–zooplankton–detritus, phosphorus-based ecosystem model coupled to a nonlinear, reduced-gravity, circulation model. The ecosystem model, an improved version of the earlier eco-hydrodynamics model developed for Lake Tanganyika, was used to estimate the annual primary production of Lake Tanganyika and its spatial and temporal variability. The simulations were driven with the National Centres for Environmental Protection (NCEP) records for winds and solar radiation forcing.
2. The simulated annual cycles of the four ecosystem variables and the daily net primary production were compared with the observations. The comparison showed that simulations reproduced realistically the general features of the annual cycles of epilimnial phosphate, net primary production and plankton dynamics.
3. The climatic simulations for the years 1970–2006 yielded a daily averaged integrated upper layer net production ranging from 0.11 to 1.78 g C m⁻² day⁻¹ and daily averaged chlorophyll- a (chl- a ) from 0.16 to 4.3 mg m⁻³. Although the nutrient concentrations in the epilimnion during the strong wind years were high, the net production was low, which is partly because of the greater vertical mixing, produced by strong winds, exposing the phytoplankton to low light conditions in deeper waters. The simulated annual net production and chl- a agreed quite well with observed production available in the literature.
4. We envisage using this model to predict the future scenarios of primary productivity in the lake.     

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.     

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.     

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.