Competitive habitat utilization in the damselfly,Mnais nawai (Zygoptera: Calopterygidae) coexisting with a related species,Mnais pruinosa

Reproductive behaviors related to habitat utilization were studied in males of the damsefly,Mnais nawai, which has two male forms, territorial orange-winged males (nawai) and non-territorial pale-orange-winged males (sahoi), at the upper part of a mountain stream where they partiallycoexist with a related species,Mnais pruinosa, which also has two male forms, territorial orange-winged males (esakii) and non-territorial hyaline-winged males (strigata). These two species showed parapatric distribution; the lower part of the stream was occupied byM. nawai, and the upper part byM. pruinosa. In the present study, cross-matings occurred between bothMnais species, although normal intraspecific matings occurred more frequently than cross-matings. Territorial males of both species copulated with conspecific females that entered their territory and guarded the ovipositing females, probably to avoid sperm displacement resulting from subsequent copulations. Severe competition for oviposition sites by territorial males even occurred between the two species. On the other hand, non-territorial males of both species have alternative mating strategies (including several tactics such as sneaking, takeover and interception). The possible benefits from conflict among territorial males of both species is discussed.     

Space use and genetic structure do not maintain color polymorphism in a species with alternative behavioral strategies

Space use including territoriality and spatial arrangement within a population can reveal important information on the nature, dynamics, and evolutionary maintenance of alternative strategies in color polymorphic species. Despite the prevalence of color polymorphic species as model systems in evolutionary biology, the interaction between space use and genetic structuring of morphs within populations has rarely been examined. Here, we assess the spatial and genetic structure of male throat color morphs within a population of the tawny dragon lizard, Ctenophorus decresii. Male color morphs do not differ in morphology but differ in aggressive and antipredator behaviors as well as androgen levels. Despite these behavioral and endocrine differences, we find that color morphs do not differ in territory size, with their spatial arrangement being essentially random with respect to each other. There were no differences in genetic diversity or relatedness between morphs; however, there was significant, albeit weak, genetic differentiation between morphs, which was unrelated to geographic distance between individuals. Our results indicate potential weak barriers to gene flow between some morphs, potentially due to nonrandom pre‐ or postcopulatory mate choice or postzygotic genetic incompatibilities. However, space use, spatial structure, and nonrandom mating do not appear to be primary mechanisms maintaining color polymorphism in this system, highlighting the complexity and variation in alternative strategies associated with color polymorphism.     

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.     

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.     

Simulation study of territory size distributions in subterranean termites

In this study, on the basis of empirical data, we have simulated the foraging tunnel patterns of two subterranean termites, Coptotermes formosanus Shiraki and Reticulitermes flavipes (Kollar), using a two-dimensional model. We have defined a territory as a convex polygon containing a tunnel pattern and explored the effects of competition among termite territory colonies on the territory size distribution in the steady state that was attained after a sufficient simulation time. In the model, territorial competition was characterized by a blocking probability P_block that quantitatively describes the ease with which a tunnel stops its advancement when it meets another tunnel; higher P_block values imply easier termination. In the beginning of the simulation run, N=10, 20,…,100 territory seeds, representing the founding pair, were randomly distributed on a square area. When the territory density was less (N=20), the differences in the territory size distributions for different P_block values were small because the territories had sufficient space to grow without strong competitions. Further, when the territory density was higher (N>20), the territory sizes increased in accordance with the combinational effect of P_block and N. In order to understand these effects better, we introduced an interference coefficient γ. We mathematically derived γ as a function of P_block and N: γ(N,P_block)=a(N)P_block/(P_block+b(N)). a(N) and b(N) are functions of N/(N+c) and d/(N+c), respectively, and c and d are constants characterizing territorial competition. The γ function is applicable to characterize the territoriality of various species and increases with both the P_block values and N; higher γ values imply higher limitations of the network growth. We used the γ function, fitted the simulation results, and determined the c and d values. In addition, we have briefly discussed the predictability of the present model by comparing it with our previous lattice model that had been used to explain the territory size distributions of mangrove termites on the Atlantic coast of Panama.