The social and ecological costs of an ‘over-extended' phenotype

Extended phenotypes offer a unique opportunity to experimentally manipulate and identify sources of selection acting on traits under natural conditions. The social cichlid fish Neolamprologus multifasciatus builds nests by digging up aquatic snail shells, creating an extended sexual phenotype that is highly amenable to experimental manipulation through addition of extra shells. Here, we find sources of both positive sexual selection and opposing natural selection acting on this trait; augmenting shell nests increases access to mates, but also increases social aggression and predation risk. Increasing the attractiveness of one male also changed social interactions throughout the social network and altered the entire community structure. Manipulated males produced and received more displays from neighbouring females, who also joined augmented male territories at higher rates than unmanipulated groups. However, males in more attractive territories received more aggression from neighbouring males, potentially as a form of social policing. We also detected a significant ecological cost of the ‘over-extended'' phenotype; heterospecific predators usurped augmented nests at higher rates, using them as breeding sites and displacing residents. Using these natural experiments, we find that both social and ecological interactions generate clear sources of selection mediating the expression of an extended phenotype in the wild.     

山鶺鴒繁殖期的领域鸣唱特征分析

山(Ｄｅｎｄｒｏｎａｎｔｈｕｓｉｎｄｉｃｕｓ)为雀形目科的小型鸟类,在杭州地区为夏候鸟。4月中、下旬抵达,5～7月为其繁殖季节。多活动于林地及林间空旷地。主食多种农林害虫[1],为著名的益鸟,但数量很少。以往对其生态学和行为方面的研究也不多。在繁殖初期,山为了占领域,往往站立于屋脊或高树上,头左右顾盼,不时地发出尖细的“ｚｈｉｊ敶,ｚｈｉｊ敶,…”如拉锯似的鸣声,因而在野外极易识别。1996年5月上旬至6月中旬,我们对栖息于本校园内的山的占区鸣叫行为进行了观察,同时记录其鸣声,对其鸣声的特征进行了分析。并根…     

Territory acquisition in a polymorphic lizard: An experimental approach

Males of the colour polymorphic Australian painted dragon lizard Ctenophorus pictus occur in red or yellow head colouration. In a previous experiment, we showed that red is associated with a higher probability of winning staged contests for resources (females or space) and that manipulation of male colouration by painting males in the opposing morph changed the dynamics of staged interactions by prolonging them 30‐fold. Thus, colour is linked to behavioural differences between males and is involved in information transfer between competing males. This inherent red dominance could result in yellow males being marginalized to poorer quality territories in terms of access to females, food, perch sites or shade. With an experiment in the wild, we test to what extent this prediction is upheld, and how colour manipulation affects morph‐specific success in territory acquisition when male body size, territory quality and emergence time from hibernation are controlled through manipulation or randomization. There was no significant effect of colour category per se, although on average red males remained closer to the release sites (our proxy for territory acquisition ability) than yellow males and artificially altered morphs moved the furthest away. There was a significant interaction effect between colour category and experimental release position, which may be linked to differences in how exposed (or not) these positions were and morph‐specific ability to cope with such exposure (e.g. ‘boldness’). Our data show that territory acquisition success is not merely a function of competitive ability but a composite outcome of a suite of factors, including signal perception.     

Investigating patterns and processes of demographic variation: environmental correlates of pre-breeding survival in red-billed choughs Pyrrhocorax pyrrhocorax

1. Quantifying the pattern of temporal and spatial variation in demography, and identifying the factors that cause this variation, are essential steps towards understanding the structure and dynamics of any population. 2. One critical but understudied demographic rate is pre-breeding survival. We used long-term colour-ringing data to quantify temporal (among-year) and spatial (among-nest site) variation in pre-breeding survival in red-billed choughs (Pyrrhocorax pyrrhocorax) inhabiting Islay, Scotland, and identified environmental correlates of this variation. 3. Random-effects capture-mark-recapture models demonstrated substantial temporal and spatial process variance in first-year survival; survival from fledging to age 1 year varied markedly among choughs fledged in different years and fledged from different nest sites. Spatial variance exceeded temporal variance across choughs fledged from well-studied nest sites. 4. The best-supported models of temporal variation suggested that first-year survival was higher in years following high tipulid larvae abundance and when weather conditions favoured increased invertebrate productivity and/or availability to foraging choughs. These variables explained up to 80% of estimated temporal process variance. 5. The best-supported models of spatial variation suggested that first-year survival was higher in choughs fledged from nest sites that were further from exposed coasts and closer to flocking areas, and surrounded by better habitat and higher chough density. These variables explained up to 40% of estimated spatial process variance. 6. Importantly, spatio-temporal models indicated interactive effects of weather, tipulid abundance, local habitat and local chough density on first-year survival, suggesting that detrimental effects of poor weather and low tipulid abundance may be reduced in choughs fledged from nest sites surrounded by better foraging habitat and lower chough density. 7. These analyses demonstrate substantial temporal and small-scale spatial variation in pre-breeding survival, a key demographic rate, and indicate that this variation may reflect interactive effects of weather, prey abundance, habitat and geography. These patterns illustrate the value of holistic models of demographic variation, and indicate environmental factors that may limit the growth rate of Islay's protected chough population.     

Experimental analysis of territory size in a population of the fire ant Solenopsis invicta

Alternative models of territoriality are based on contrastingassumptions about the behavioral processes determining territorysize. In a series of controlled field experiments on the fireant Solenopsis invicta, I tested whether territory size is affectedby the availability of food, as predicted by most economic models,and whether territory size is affected by fighting ability,as predicted by models of competition among neighbors. Abundantfood was offered for 30–35 days to selected colonies eitherimmediately next to the nest (experiment 1) or at peripheralsites near the territory boundary (experiment 2). These foodsupplements had no detectable effect on territory size. Furthermore,food placed near the periphery of the territory did not significantlyalter local boundary positions. During both experiments, largecolonies lost more territory than did small colonies, reflectingtemporary declines in worker number due to the seasonal productionof reproductives. Such losses by large colonies during the summermonths create opportunities for newly founded colonies to expandterritories. In a third experiment, colonies from which workerswere removed lost significantly more territory than did unmanipulatedcontrols. These results show that territory areas in S. invictaare strongly affected by the relative fighting ability of neighboringcolonies but provide no evidence that colonies adjust territoryarea in response to short-term changes in the availability offood.     

Reproductive success of Tree Pipits Anthus trivialis in relation to habitat selection in conifer plantations

Aspects of the reproductive success of Tree Pipits Anthus trivialis were examined in relation to broad‐scale habitat and nest‐site selection in Thetford Forest, a coniferous plantation forest in eastern England. Three habitat classes were defined corresponding to previously reported densities of Tree Pipits: clearfell and recently planted stands (habitat class A: low density), stands 2–5 years old (B: high density) and stands 6 years or older (C: low density). The preference for 2–5‐year‐old stands indicated by higher densities was supported by the timing of territory settlement. Tree Pipits also showed distinct preferences for nest‐site characteristics that were relatively consistent across habitat classes and throughout the breeding season. At the ‘habitat scale’, results were consistent with the predictions of the ideal despotic distribution model. First clutches were laid significantly earlier in the preferred habitat class B. Overall nesting success (i.e. the proportion of nests producing fledglings), but not clutch size, also varied between habitats, being greater in habitat classes B and C than in habitat class A. The variation in overall nesting success between habitats was primarily driven by low nest survival rates during the laying/incubation period in clearfell and recently planted stands. Nest survival rates during the nestling period were lower in the preferred 2–5‐year‐old (and older) stands and declined over the course of the study. Preferences for nest‐site characteristics (at least for those that were measured) provided no apparent benefit to nest survival rates. Overall nesting success thus appeared to be determined at the habitat scale, perhaps because the broad differences in cover between habitats affected the likelihood of nest predation (the main cause of nest failure). It is suggested that the very low nesting success experienced by Tree Pipits in clearfell and new stands may be one factor in the species’ relative avoidance of this habitat and preference for 2–5‐year‐old stands.     

The deterrent effect of bird song in territory defense

Using the responses of territory owners to playback to inferthe territorial function of acoustic signals is common practice.However, difficulties with interpreting the results of suchexperiments have obscured our understanding of territorial signalling.For instance, a stronger response to playback is often interpretedas more aggressive, but there is no consensus as to whetherthis should be in response to the least or most threateningsimulated intruder. Rather than following a gradual increaseor decrease, the relationship between signal intensity and responsestrength may instead describe a peaked curve. We manipulatedbanded wren (Thryophilus pleurostictus) songs to simulate low-,median-, and high-performance singers and used these songs asstimuli in playback experiments. Banded wrens were less likelyto approach the high-performance stimulus compared with thelow- and median-performance stimuli. However, the birds thatdid approach the high-performance stimulus sang more than thosethat approached the low-performance stimulus. In addition, birdswere more likely to match the songs when exposed to the median-and high-performance stimuli compared with the low-performancestimuli, and song matching predicted approach behavior. Theseresults are in accordance with theoretical models of aggressiveencounters in which low-performance opponents are challengedwithout further assessment. Median- and high-performance opponents,however, may require further assessment, and the latter maybe perceived as too intimidating for approach.     

Access to mates in a territorial ungulate is determined by the size of a male's territory, but not by its habitat quality

1. Territoriality is commonly associated with resource defence polygyny, where males are expected to gain access to females by anticipating how resources will influence female distribution and competing for resource-rich sites to establish their zone of dominance. 2. We tested this hypothesis in European roe deer (Capreolus capreolus) by simultaneously assessing the influence of resources on female distribution and the influence of female distribution on male distribution and breeding success using paternity analyses. 3. Females did not fully distribute themselves among male territories in relation to resources. As a result, relative female abundance in a male's territory depended on territory size, but not on its habitat quality. In turn, relative female abundance in a male's territory determined, at least partially, his breeding success. 4. Interestingly, male territory size, and hence access to females, was partly determined by male body mass (all males) and by residual antler size (subadults only). The latter result suggests that large antlers may be important to young males for establishing their first territory, which is then usually retained for all subsequent reproductive seasons. 5. To conclude, although territoriality of male roe deer has certainly evolved as a tactic for ensuring access to mates, our results suggest that it does not really conform to a conventional resource defence polygyny strategy, as males seem to gain no obvious benefit from defending a territory in an area of high habitat quality in terms of enhanced access to mates. 6. This may explain the stability of male territories between years, suggesting that male territoriality conforms to an 'always stay' and 'low risk-low gain' mating strategy in roe deer.