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.     

Intraspecific phenotypic variation in a fish predator affects multitrophic lake metacommunity structure

Contemporary insights from evolutionary ecology suggest that population divergence in ecologically important traits within predators can generate diversifying ecological selection on local community structure. Many studies acknowledging these effects of intraspecific variation assume that local populations are situated in communities that are unconnected to similar communities within a shared region. Recent work from metacommunity ecology suggests that species dispersal among communities can also influence species diversity and composition but can depend upon the relative importance of the local environment. Here, we study the relative effects of intraspecific phenotypic variation in a fish predator and spatial processes related to plankton species dispersal on multitrophic lake plankton metacommunity structure. Intraspecific diversification in foraging traits and residence time of the planktivorous fish alewife (Alosa pseudoharengus) among coastal lakes yields lake metacommunities supporting three lake types which differ in the phenotype and incidence of alewife: lakes with anadromous, landlocked, or no alewives. In coastal lakes, plankton community composition was attributed to dispersal versus local environmental predictors, including intraspecific variation in alewives. Local and beta diversity of zooplankton and phytoplankton was additionally measured in response to intraspecific variation in alewives. Zooplankton communities were structured by species sorting, with a strong influence of intraspecific variation in A. pseudoharengus. Intraspecific variation altered zooplankton species richness and beta diversity, where lake communities with landlocked alewives exhibited intermediate richness between lakes with anadromous alewives and without alewives, and greater community similarity. Phytoplankton diversity, in contrast, was highest in lakes with landlocked alewives. The results indicate that plankton dispersal in the region supplied a migrant pool that was strongly structured by intraspecific variation in alewives. This is one of the first studies to demonstrate that intraspecific phenotypic variation in a predator can maintain contrasting patterns of multitrophic diversity in metacommunities.     

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.     

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.     

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.     

The effects of competitor density on aggressive behaviour and resource defence in a Poeciliid fish

To test the hypothesis that competitor density and time spent at a food resource influences aggressive behaviour in male swordtail, Xiphophorus sp., fish were kept at two densities (low: 18 fish m^?3; high: 54 fish m^?3) and aggressive behaviour was recorded with the time at the food resource used as a covariate. Competitor density is likely to affect the cost:benefit ratio of food defence. When density is low, there should be sufficient food for all individuals and intra‐specific interactions are expected to be rare, while at high densities, increased intra‐specific encounter rates mean that individuals may spend more time defending a resource than utilising it. Food resource defence should occur at intermediate densities. The frequency of aggression, i.e. bites, chases, and display behaviour was significantly positively influenced by time spent at the food resource (Bites: F_1,31 = 8.186, P = 0.007; Chases: F_1,31 = 6.439, P = 0.016; Displays: F_1,31 = 4.435, P = 0.043) suggesting that food resource defence occurred. Competitor density had no effect on food resource defence and minimal effect on the frequency of aggression with only one type of aggressive behaviour, male–male displays, showing a difference between densities (F_1,31 = 6.975, P = 0.013). This finding is suggested to be a result of the formation of dominance hierarchies in this species. Aggression from the dominant individual may be directed at only subordinates of the next dominance rank and subordinate behaviour may be restricted by status rather than immediate threat. In such a situation, aggression may be independent of competitor density.     

Introduced invertebrates are important prey for a generalist predator

Concern over biological invasions has drawn increased attention to the impacts of introduced predators or competitors, but not to the importance of introduced prey. North American forests are rich in introduced invertebrates, including species that represent relatively novel taxonomic or trophic guilds and show biased distributions among forest types. We analysed the diets of red‐backed salamanders, Plethodon cinereus, from three upland and three lowland forests to determine whether introduced prey are important contributors to geographical or temporal variation in salamander food resources. We found several introduced species were volumetrically important salamander prey, and were responsible for resource differences between forest types and much of the seasonal fluctuation in food resources in both forest types. In lowland forests, rain had a stronger effect on salamander predation on non‐native earthworms than native taxa, creating more dynamic resource fluctuations in resource levels than was observed in upland forests where earthworms were absent. With one exception, predation on non‐native species was positively associated with predation on native species, suggesting non‐native prey have added to salamander resources rather than replaced salamander predation on native taxa. We hypothesize that the novel resource gradients created by non‐native prey introductions are contributing to patterns of geographical and temporal phenotypic variation among salamander populations.     

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.     

Parental response to brood size in a cichlid fish

This paper reports on an experiment designed to investigate whether guarding females of the cichlid Aequidens coeruleopunctatus adjust their parental behaviour in response to changes in brood size. It was found that females were easier to scare away from their broods when threatened, and stayed away for longer, after some of their fry had been removed, and that the reverse occurred when broods were experimentally augmented with foreign fry. Brood size manipulations change the expected benefits of future investment. These results therefore indicate that parental care strategies in A. coeruleopunctatus reflect future prospects for reproductive success rather than past investment.     

Effect of boat noise and angling on lake fish behaviour

The effects of disturbances from recreational activities on the swimming speed and habitat use of roach Rutilus rutilus, perch Perca fluviatilis and pike Esox lucius were explored. Disturbances were applied for 4 h as (1) boating in short intervals with a small outboard internal combustion engine or (2) boating in short intervals combined with angling with artificial lures between engine runs. The response of the fish species was evaluated by high‐resolution tracking using an automatic acoustic telemetry system and transmitters with sub‐minute burst rates. Rutilus rutilus swimming speed was significantly higher during disturbances [both (1) and (2)] with an immediate reaction shortly after the engine started. Perca fluviatilis displayed increased swimming activity during the first hour of disturbance but not during the following hours. Swimming activity of E. lucius was not significantly different between disturbance periods and the same periods on days without disturbance (control). Rutilus rutilus increased their use of the central part of the lake during disturbances, whereas no habitat change was observed in P. fluviatilis and E. lucius. No difference in fish response was detected between the two types of disturbances (boating with and without angling), indicating that boating was the primary source of disturbance. This study highlights species‐specific responses to recreational boating and may have implications for management of human recreational activities in lakes.     

Visual adaptation could aid sympatric speciation in a deep crater lake

Allopatric speciation was originally suggested to be the primary mechanism of animal speciation (Mayr, 1942; Figure 1). During allopatric speciation, populations diverge when gene flow is reduced across significant biogeographic barriers. Sympatric speciation, where species diverge while inhabiting the same location, was thought to be essentially impossible. However, the advent of theoretical models followed by new experimental evidence made sympatric speciation more plausible (Via, 2001). The cichlid fishes of Barombi Mbo, a small crater lake in western Cameroon, became one of the most widely accepted examples of sympatric speciation (Schliewen, Tautz, & Paabo, 1994). Although the phylogenetic history of this clade is not quite as simple as originally thought, it remains one of the best examples of sympatric speciation (Richards, Poelstra, & Martin, 2018). However, little is known about the molecular mechanisms contributing to the splitting of these species in situ. In a From the Cover article in this issue of Molecular Ecology, Musilova et al. (2019) focus on the diversity of visual systems among these fishes. They identify genetic changes associated with several aspects of visual adaptation that may have contributed to the ecological specialization and sympatric speciation of cichlids in this lake.     

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.     

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.     

Referential alarm calling elicits future vigilance in a host of an avian brood parasite

Yellow warblers (Setophaga petechia) use referential ‘seet’ calls to warn mates of brood parasitic brown-headed cowbirds (Molothrus ater). In response to seet calls during the day, female warblers swiftly move to sit tightly on their nests, which may prevent parasitism by physically blocking female cowbirds from inspecting and laying in the nest. However, cowbirds lay their eggs just prior to sunrise, not during daytime. We experimentally tested whether female warblers, warned by seet calls on one day, extend their anti-parasitic responses into the future by engaging in vigilance at sunrise on the next day, when parasitism may occur. As predicted, daytime seet call playbacks caused female warblers to leave their nests less often on the following morning, relative to playbacks of both their generic anti-predator calls and silent controls. Thus, referential calls do not only convey the identity or the type of threat at present but also elicit vigilance in the future to provide protection from threats during periods of heightened vulnerability.     

Shovelnose sturgeon exhibit predator avoidance behaviour in the presence of a hungry predator

An experiment was designed to test whether age‐0 shovelnose sturgeon (Scaphirhynchus platorynchus) exhibited predator avoidance behaviour in response to a channel catfish (Ictalurus punctatus) predator. It was hypothesized that shovelnose sturgeon would not exhibit any innate predator avoidance behaviour because previous reports have shown a congener of the shovelnose sturgeon, the pallid sturgeon (S. albus), to be an unfavourable prey item for channel catfish. The results, however, indicated that shovelnose sturgeon generally avoided space occupied by the catfish predator and spent a greater proportion of time in the predator avoidance zone within the experimental tank. Bitten fish, in particular, spent a greater period of time in the predator avoidance zone. Of all sturgeon used in this experiment (N = 30), 73% swam within the fork length (350 mm) of the catfish predator. The results seem to indicate that shovelnose sturgeon were initially oblivious to the risk of predation by the catfish predator, but after interaction (e.g. being chased or bitten) appeared to display predator avoidance behaviour. Predator avoidance behaviour in shovelnose sturgeon may thus be suggested as a learned rather than an innate behaviour.     

Limnology of Botos Lake, a tropical crater lake in Costa Rica

Botos Lake, located at the Poas Volcano complex (Costa Rica) was sampled eight times from 1994 to 1996 for physicochemical conditions of the water column and phytoplanktonic community composition. Depth was measured at fixed intervals in several transects across the lake to determine its main morphometric characteristics. The lake has an outlet to the north. It is located 2580 m above sea level and is shallow, with a mean depth of 1.8 m and a relative depth of 2.42 (surface area 10.33 ha, estimated volume 47.3 hm3). The lake showed an isothermal water column in all occasions, but it heats and cools completely according to weather fluctuations. Water transparency reached the bottom on most occasions (> 9 m). The results support the idea that the lake is polymictic and oligotrophic. The lake has at least 23 species of planktonic algae, but it was always dominated by dinoflagellates, especially Peridinium inconspicuum. The shore line is populated by a sparse population of Isoetes sp. and Eleocharis sp. mainly in the northern shore where the bottom has a gentle slope and the forest does not reach the shore.