Sex-specific defence behaviour against brood parasitism in a host with female-only incubation

Nest protection against intruders is an indispensable component of avian parental care. In species with biparental care, both mates should evolve nest defence behaviour to increase their reproductive success. In most host-parasite systems, host females are predicted to have more important roles in nest defence against brood parasites, because they typically are primarily responsible for clutch incubation. Male antiparasitic behaviour, on the other hand, is often underestimated or even not considered at all. Here we investigated sex-specific roles in four aspects of great reed warbler (Acrocephalus arundinaceus) nest defence against a brood parasite—the cuckoo (Cuculus canorus), namely (1) mobbing, (2) nest attendance/guarding, (3) nest checking and (4) egg ejection. Using dummy experiments, simulating brood parasitism and by video-monitoring of host nests we found that males took the key roles in cuckoo mobbing and nest guarding, while females were responsible for nest checking and egg ejection behaviours. Such partitioning of parental roles may provide a comprehensive clutch protection against brood parasitism.     

Cooperation in defence against a predator

The origin and the evolutionary stability of cooperation between unrelated individuals is one of the key problems of evolutionary biology. In this paper, a cooperative defence game against a predator is introduced which is based on Hamilton's selfish herd theory and Eshel's survival game models. Cooperation is altruistic in the sense that the individual, which is not the target of the predator, helps the members of the group attacked by the predator and during defensive action the helper individual may also die in any attack. In order to decrease the long term predation risk, this individual has to carry out a high risk action. Here I show that this kind of cooperative behaviour can evolve in small groups. The reason for the emergence of cooperation is that if the predator does not kill a mate of a cooperative individual, then the survival probability of the cooperative individual will increase in two cases. If the mate is non-cooperative, then—according to the dilution effect, the predator confusion effect and the higher predator vigilance—the survival probability of the cooperative individual increases. The second case is when the mate is cooperative, because a cooperative individual has a further gain, the active help in defence during further predator attacks. Thus, if an individual can increase the survival rate of its mates (no matter whether the mate is cooperative or not), then its own predation risk will decrease.     

Persistence of host defence behaviour in the absence of avian brood parasitism

The fate of host defensive behaviour in the absence of selection from brood parasitism is critical to long-term host-parasite coevolution. We investigated whether New World Bohemian waxwings Bombycilla garrulus that are allopatric from brown-headed cowbird Molothrus ater and common cuckoo Cuculus canorus parasitism have retained egg rejection behaviour. We found that egg rejection was expressed by 100 per cent of Bohemian waxwings. Our phylogeny revealed that Bohemian and Japanese waxwings Bombycilla japonica were sister taxa, and this clade was sister to the cedar waxwing Bombycilla cedrorum. In addition, there was support for a split between Old and New World Bohemian waxwings. Our molecular clock estimates suggest that egg rejection may have been retained for 2.8-3.0 Myr since New World Bohemian waxwings inherited it from their common ancestor with the rejecter cedar waxwings. These results support the 'single trajectory' model of host-brood parasite coevolution that once hosts evolve defences, they are retained, forcing parasites to become more specialized over time.     

Chemically mediated predator inspection behaviour in the absence of predator visual cues by a characin fish

Animals commonly approach (i.e. 'inspect') potential predators. Glowlight tetras, Hemigrammus erythrozonus, have previously been shown to inspect the combined chemical and visual cues originating from novel predators and to modify their inspection (approach) behaviour depending upon the predator's diet. We conducted two experiments to determine whether tetras would inspect the chemical cues of injured prey or the dietary cues of a novel predator in the absence of any visual cues. Shoals of glowlight tetras were exposed to either distilled water (control) or the skin extract of swordtail (lacking ostariophysan alarm pheromones) or the skin extract of tetra (with alarm pheromones). There was no significant difference in the frequency of predator inspection behaviour towards swordtail or tetra skin extract compared to the distilled water controls. In the second experiment, we exposed shoals of tetras to either distilled water or the odour of Jack Dempsey cichlids, Cichlasoma octofasciatum, which had been food deprived, or fed a diet of swordtails or tetras. There was no significant difference in the frequency of predator inspection behaviour towards the odour of the starved cichlids and the odour of the fed cichlids in either of the two diet treatments. However, when tetras were exposed to the odour of cichlids fed tetras, they took significantly longer to initiate an inspection visit, remained further from the source of the chemical cues and inspected in smaller groups, compared with the odour of a starved cichlid or a cichlid fed swordtails. These data strongly suggest that tetras will inspect chemical cues alone, but only if the cue contains information about the predator. Copyright 2000 The Association for the Study of Animal Behaviour.     

Brood size and the economy of brood defence: examining Lack's hypothesis in a biparental cichlid fish

Synopsis We tested the explanatory value of two hypotheses reviewed by Lack (1954) in the maintenance of brood size in free-ranging convict cichlidsCichlasoma nigrofasciatum: (1) physiological constraints on egg production, and (2) behavioural constraints imposed by brood defence. Number of free-swimming young in 13 experimental (E) broods was augmented to the upper limit of the size distribution of natural broods (150 young); 18 control (C) broods were handled in the same way but brood size was not changed (mean ± SE = 69.5 ± 11.0). E and C brood sizes were measured at 5 day intervals. At day 20 (just before independence from parental care), 50.3 ± 9.4 (n = 9) young remained in E broods and 30.8 ± 7.8 (n = 8) young remained in C broods (p> 0.05). Offspring number did not differ significantly (p> 0.05) between C and E broods after day 10. Mean growth rate of offspring was significantly lower in E broods than in C broods, perhaps in response to increased density of young in the former. Both the convergence of offspring number in E and C broods and suppression of growth in E broods support a behavioural constraint; that during the first 10 days in which the young are free swimming, two parents are unable to defend large broods as successfully as small broods. A trade-off exists in parental investment between current and future reproduction. Extra-parental investment in current reproduction (eggs) does not result in an increased number of young at independence, therefore a behavioural constraint during brood defence should stabilize the evolution of clutch size.     

Who dares, learns: chemical inspection behaviour and acquired predator recognition in a characin fish

Individuals that dare approach predators (predator inspection behaviour) may benefit by acquiring information regarding the potential threat of predation. Although information acquisition based on visual cues has been demonstrated for fish, it is unknown whether fish will inspect predators on the basis of chemical cues or whether such inspection behaviour results in information acquisition. Here, we first ascertained whether predator inspection behaviour can be mediated by chemical cues from predators by exposing groups of predator-naive glowlight tetras (Hemigrammus erythrozonus) to the chemical cues of a potential fish predator (convict cichlid Cichlasoma nigrofasciatum) that had been fed either tetras (which possess an alarm pheromone) or swordtails (Xiphophorus helleri, which lack Ostariophysan alarm pheromones). Tetras showed a significant increase in antipredator behaviour when exposed to the tetra-diet cue, but not when exposed to the swordtail-diet cue. Chemically mediated predator inspection behaviour was also affected. Both the latency to inspect and the minimum approach distance to the predator significantly increased, and the mean number of inspectors per predator inspection visit significantly decreased when tetras were exposed to the tetra-diet versus the swordtail-diet chemical cues. We then examined a potential benefit associated with chemically mediated predator inspection behaviour. Only tetras that were initially exposed to the tetra-diet cue and that had inspected the predator acquired the visual recognition of a convict cichlid as a predation threat. Our results thus demonstrate that (1) predator inspection behaviour in the glowlight tetra can be initiated by chemical cues, (2) chemically mediated inspection behaviour is affected by the presence of alarm pheromone, and (3) inspectors benefit by acquiring the recognition of novel predators. Copyright 1999 The Association for the Study of Animal Behaviour.     

Reticulate sympatric speciation in Cameroonian crater lake cichlids

BACKGROUND: Traditionally the rapid origin of megadiverse species flocks of extremely closely related species is explained by the combinatory action of three factors: Disruptive natural selection, disruptive sexual selection and partial isolation by distance. However, recent empirical data and theoretical advances suggest that the diversity of complex species assemblages is based at least partially on the hybridization of numerous ancestral allopatric lineages that formed hybrids upon invasion of new environments. That reticulate speciation within species flocks may occur under sympatric conditions after the primary formation of species has been proposed but not been tested critically. RESULTS: We reconstructed the phylogeny of a complex cichlid species flock confined to the tiny Cameroonian crater lake Barombi Mbo using both mitochondrial and nuclear (AFLP) data. The nuclear phylogeny confirms previous findings which suggested the monophyly and sympatric origin of the flock. However, discordant intra-flock phylogenies reconstructed from mitochondrial and nuclear data suggest strongly that secondary hybridization among lineages that primarily diverged under sympatric conditions had occurred. Using canonical phylogenetic ordination and tree-based tests we infer that hybridization of two ancient lineages resulted in the formation of a new and ecologically highly distinct species, Pungu maclareni. CONCLUSIONS: Our findings show that sympatric hybrid speciation is able to contribute significantly to the evolution of complex species assemblages even without the prior formation of hybrids derived from allopatrically differentiated lineages.     

Dissolved oxygen and thermal regimes of a Ugandan crater lake

This paper quantifies the temporal pattern of thermal stratification and deoxygenation in Lake Nkuruba, a small (3 ha), deep (maximum depth = 38 m) crater lake in western Uganda. Dissolved oxygen penetrated to an average depth of 9 m and a maximum depth of 15 m below which the lake was permanently anoxic over the 2 years of study. Although surface oxygen levels were correlated with both surface water temperature and rainfall, seasonal cycles of dissolved oxygen were not well-defined and may have been obscured by the high frequency of short-term fluctuations and by inter-annual variations caused by shifts in rainfall. Surface water temperature averaged 23.3±0.7 °C (S.D.) and varied directly with air temperature. Both diurnal changes and top-bottom temperature differentials were small averaging 1.7±0.7 °C and 1.6±0.8 °C, respectively. Thermal stability ranged from 101.3 to 499.9 g-cm cm^-2 and was positively related to surface water temperature suggesting that this small protected lake responds rapidly to short-term meteorological changes. Because contribution to the annual heat exchange cycle was confined to upper waters, the lake's annual heat budget was low, 1,073.8 cal cm^-2 yr^-1. However, net primary productivity was relatively high averaging 1.3 g C m^-2d^-1. The region where Lake Nkuruba is situated experienced a very strong earthquake (6.2 on the Richter scale) on 4 February, 1994. Subsequently, water levels dropped markedly in the lake, falling 3.14 m over a 5-month period. This revised version was published online in September 2006 with corrections to the Cover Date.     

Parental behaviour in Copris lunaris (Coleoptera, Scarabaeidae): care and defence of brood balls and nest

Abstract. 1. Nesting female beetles righted brood balls (so as to replace the egg or larva in the uppermost position) and repaired damaged balls. This behaviour required the presence of an egg or larva in the ball, or of a short-lasting material found just after oviposition. The shape of the ball was also a righting stimulus since artificial ellipsoids were stood on end.
2. Balls containing dichloromethane extracts of C.lunaris brood were righted and repaired. Eggs and larvae of several other Scarabaeidae did not release these responses but were destroyed.
3. Righting behaviour was released when brood was absent from the top of the ball. The beetle then crawled vertically downwards and, if it encountered the displaced apex, a novel rolling action followed which automatically turned the ball towards the correct position.
4. An opening made in the nest was repaired with soil excavated from the chamber floor. Clunaris adults and Aphodius fossor larvae were attacked if they were encountered in the nest.     

Temporal variation of phytoplankton in a small tropical crater lake, Costa Rica

The temporal variation in lake's phytoplankton is important to understand its general biodiversity. For tropical lakes, it has been hypothesized that they follow a similar pattern as temperate ones, on a much accelerated pace; nevertheless, few case studies have tried to elucidate this. Most studies in Costa Rica have used a monthly sampling scheme and failed in showing the expected changes. In this study, the phytoplankton of the small Barvas's crater lake was followed for more than three years, first with monthly and later with weekly samplings, that covered almost two years. Additional information on temperature and oxygen vertical profiles was obtained on a monthly basis, and surface temperature was measured during weekly samplings around noon. Results showed that in spite of its shallow condition (max. depth: 7m) and low surface temperature (11 to 19 degrees C), the lake stratifies at least for brief periods. The phytoplankton showed both, rapid change periods, and prolonged ones of relative stasis. The plankton composition fluctuated between three main phases, one characterized by the abundance of small sized desmids (Staurastrum paradoxum, Cosmarium asphaerosporum), a second phase dominated by equally small cryptomonads (Chryptochrysis minor, Chroomonas sp.) and a third phase dominated by the green alga Eutetramorus tetrasporus. Although data evidenced that monthly sampling could miss short term events, the temporal variation did not follow the typical dry and rainy seasons of the region, or any particular annual pattern. Year to year variation was high. As this small lake is located at the summit of Barva Volcano and receives the influence from both the Caribbean and the Pacific weather, seasonality at the lake is not clearly defined as in the rest of the country and short term variations in the local weather might have a stronger effect than broad seasonal trends. The occurrence of this short term changes in the phytoplankton of small tropical lakes in response to weather variations needs to be further explored in other lakes.     

Calcium carbonate formation by Enteromorpha nana algae in a hypersaline volcanic crater lake

On the black basaltic rock surfaces in shallow water around the margin of Lake Gnotuk, and for several meters above the present lake, there is a thick white coating of aragonite marl. The water in the lake is supersaturated with CaCO₃.From close observation and experimental work, it is evident that the marl is produced by the alga Entermorpha nana which lives in the shallow lake water and grows attached to the basalt. The general mechanism of CaCO₃ precipitation by aquatic plants is outlined, and in Enteromorpha nana it can be demonstrated that the closed tubular thallus absorbs HCO-₃ from the outside water and secretes CO²-₃ inside the tube. This secretion of CO²-₃ greatly increases the ionic product of [Ca²⁺] [CO₂-₃] in the water inside the thallus and leads to the precipitation of aragonite and its accumulation at the base of the tube.     

Recent sympatric speciation involving habitat-associated nuptial colour polymorphism in a crater lake cichlid

Even though the idea that modes of speciation other than allopatric speciation are possible in nature is now widespread, compelling examples of ecological speciation in sympatry remain rare. We studied an undescribed radiation of haplochromine cichlids in a young crater lake in western Uganda, and in the small river that is nearby but has currently no known surface connection to the lake. We describe two different modes of speciation that occurred in this cichlid lineage within the past 1,500–10,000 years. Not constrained by gene flow, allopatric divergence between river and lake cichlids affects many different morphological traits as well as nuptial colouration—muted in the river, but intensified and polymorphic in lake cichlids—and neutral genetic differentiation. More surprisingly, we demonstrate a case for sympatric speciation within the small lake that is associated with dramatic differences in male breeding colouration (yellow with bright red-chest versus bright blue) and subtle differences in microhabitat, feeding regime and morphology. Reproductive isolation by assortative mating is suggested by significant differentiation between yellow and blue males in neutral markers of gene flow despite complete sympatry. We hypothesize speciation is mediated by divergent selection on sexual signalling between microhabitats.

     

Competitor density influences resource defence in a Poeciliid fish

The concept of economic defendability states that individuals will only defend resources when the benefits outweigh the costs. Competitor density is one factor that influences resource defence among fishes, resource defence being predicted to be highest at intermediate competitor densities and reduced at higher and lower densities, producing a dome shaped relationship. We investigated the effects of competitor density using male swordtails, which form dominance hierarchies and have already been demonstrated to defend a food resource. We found that both aggression (F_2,32 = 5.392; P = 0.010) and time at a food resource (F_2,32 = 3.574; P = 0.040) vary with differences in competitor density, and that the frequency of aggression significantly predicts time at the food resource only at an intermediate density (P = 0.033; r²adj = 0.219). The ability to vary the level of defence with respect to the immediate environment may result in fitness benefits to male swordtails. Further studies examining changes in behaviour over time and examining interacting influencing factors would be beneficial.     

Interactions between zooplankton and fish in a fertilized lake

The effects of fish predation on the zooplankton community in an oligotrophic lake, Langvatn, near Trondheim in Central Norway, were investigated during a six-year period (1973–1978), together with the added effects of changes produced by adding artificial fertilizer in 1975 and 1976. The improved nutrient conditions in 1975 resulted in a rapid increase in biomass and production of the largest herbivore zooplankton species and of the fish population. A change in the behaviour and food habits of the arctic char was recorded; they became more pelagic and fed mainly on zooplankton. An increased survival rate of 0-group and biomass of planktivorous fish in 1975 enhanced the degree of fish predation on the zooplankton during subsequent years (1976–1978). As a consequence of fish predation, the composition of the zooplankton changed, from a mainly large-sized to a mainly small-sized community, dominated by Bosmina longirostris and rotifers. Since fish predation is size-selective and visibility-dependent, it induced a decrease in mean size and in body length at onset of maturity of the cladoceran populations and probably also weakened their ability to produce resting eggs.