Territorial defense, territory size, and population regulation

The carrying capacity of an environment is determined partly by how individuals compete over the available resources. To territorial animals, space is an important resource, leading to conflict over its use. We build a model where the carrying capacity for an organism in a given environment results from the evolution of territorial defense effort and the consequent space use. The same evolutionary process can yield two completely different modes of population regulation. Density dependence arises through expanding and shrinking territories if fecundity is low, breeding success increases gradually with territory size, and/or defense is cheap. By contrast, when fecundity is high, breeding success sharply saturates with territory size, and/or defense is costly, we predict fixed territory sizes and regulation by floaters. These "surplus" individuals form a buffer against population fluctuations. Yet floaters can also harm breeder performance, and by comparing population growth of a territorial population to a nonterritorial (and individually suboptimal) alternative, we can quantify the harmful effect of evolutionary conflict on population performance. Territoriality has often been found to increase population stability, but this may come at a cost of reduced equilibrium densities.     

Histomorphology of preorbital gland in territorial and non-territorial male blackbuck Antelope cervicapra, a critically endangered species

The preorbital gland is a specialized dermal gland of antelopes which plays an important role in territorial marking behavior and pheromonal communication. To our knowledge, there is little information available on the role of preorbital gland marks in Indian antelopes (blackbucks). Males are seen averting the gland during behavioural display and territorial marking but the functional aspect of this gland has not been examined. Hence, the aim of this study was to describe the histomorphology of the preorbital gland in territorial and non-territorial male blackbucks to determine its morphology and secretory function. The results showed that the preorbital gland is composed of modified sebaceous and apocrine glands. The apocrine gland is lined by simple cuboidal epithelial cells; the serous parts of the secretory products are often seen in the apical portions of the cells. The myoepithelial cells contain actin filaments lying on the basal membranes of the apocrine gland. There are some considerable histological changes in the presence of the sebaceous and apocrine glands in territorial males in comparison to non-territorial males. The following histological changes associated with occurrence of the sebaceous and apocrine glands have been observed in territorial and non-territorial male blackbucks: (1) increase of size of sebaceous and apocrine glands and (2) increase in density of sebaceous and apocrine glands in territorial males compared to non-territorial males. It is suggested that the higher development (i.e., size) and density of sebaceous and apocrine glands in territorial males could depend on hormone production (i.e., testosterone). Based on the histological observation and the role of sebaceous and apocrine glands in the preorbital gland supported by literature, it is possible to conclude that both territorial and non-territorial blackbuck males may produce pheromonal substances through preorbital gland (secretion) for olfactory communication.     

Competitive asymmetry, foraging area size and coexistence of annuals

Individuals allocate resources to the expansion of their foraging area and those resources are no longer available for the traits that determine how well those individuals are able to protect their foraging area against competitors. The resulting trade‐off between foraging area size and the traits associated with the ability to compete for the resources within the foraging area applies to ecological scenarios as different as territorial defence by individuals and colonies, and light competition in plants. Whether the trade‐off affects species performance in competition for resources at the area of overlap between foraging areas depends on the symmetry of resource division. In symmetric competition resources are divided equally between the competitors, while in asymmetric competition the individual with the smallest foraging area, and consequently the greatest competitive ability, gains all the resources. Competition may also be a combination of the symmetric and asymmetric processes. I studied the effects of competitive asymmetry on population dynamics and coexistence of two annual species with different sized foraging areas using an individual‐based spatially explicit simulation model. Symmetric competition favoured the species with the larger foraging area and did not allow coexistence. Competitive asymmetry favoured the species with smaller foraging area and allowed coexistence, which was due to the consequences of losing an asymmetric competition being more severe than losing a symmetric competition. The mechanism of coexistence is the larger foraging area's superiority in low population densities (little competition) and the smaller foraging area's ability to win a large foraging area when competition was intense. Competitive asymmetry and small size of both foraging areas led to population dynamics dominated by long‐term fluctuations of small intensity. Symmetric competition and large size of the foraging areas led to large short‐term fluctuations, which often resulted in the extinction of one or both of the species due to demographic stochasticity.     

A global biodiversity fund to implement distributive justice for genetic resources

This article examines the question of who has a right to control and benefit from genetic resources globally. To this end it draws on different accounts in the resource rights literature with a focus on the specific features that distinguish genetic resources from other types of natural resources. It will be argued that due to the intangible and non‐territorial nature of genetic resources, territorial rights over these resources are difficult to maintain. Moreover, the vulnerability of genetic resources implies that much cost and effort is required to protect them. I will argue that not only benefits resulting from the use of genetic resources but also these costs associated with their protection should be an object of distributive justice. To accommodate these two points I will introduce the model of a global biodiversity fund that could replace the bilateral access and benefit sharing negotiations suggested by the Convention on Biological Diversity.     

Territorial spatial vulnerability assessment based on PSO-BP neural network: A case study in Shenzhen,China

Studies on vulnerability, a fairly new research paradigm inspired by global environmental change, are expanding to support scientific decision-making for developing sustainable, resilient territorial space at the regional level. The goal of this research is to reveal the vulnerability of a human-environment coupled system and its interaction mechanisms amidst a changing environment. This study started with the vulnerability scoping diagram (VSD) conceptual model and built up a research framework of territorial spatial vulnerability according to the three structural elements of ecological protection, agricultural production and urban construction. Following the requirements for intelligent computing, we introduced particle swarm optimization and BP neural network learning algorithms to evaluate the vulnerability of Shenzhen's territory. We also introduced flow cytometry to analyze the mechanism of Shenzhen's territorial spatial vulnerability driven by human and natural forces. The results showed that the territorial spatial vulnerability of Shenzhen was low under various functional orientations. Among them, ecological protection-oriented territorial spatial vulnerability was high in the west and low in the east and followed an inverted U-shaped trend in the north-south direction. Agricultural production-oriented territorial spatial vulnerability was high in the east and low in the west, high in the north and low in the south. Urban construction-oriented territorial spatial vulnerability was high in the west and low in the east, high in the north and low in the south. Under the complex orientation of the gigantic human-environment coupled system, the vulnerability of territory was symmetrical and balanced in the north-south direction and fluctuated with an overall declining trend in the east-west direction. In vulnerability, ecological protection, agricultural production and urban construction systems independently accounted for 35.642%, 38.209%, and 26.149% of the territorial spatial vulnerability, respectively.     

The role of androgens in the trade-off between territorial and parental behavior in the Azorean rock-pool blenny, Parablennius parvicornis

Androgen hormones have been shown to facilitate competitive ability in courtship and territorial behavior, while suppressing paternal behavior. The rock-pool blenny, Parablennius parvicornis, provides an excellent model to study the proximate regulation of such a trade-off between territorial and parental behavior, because nest-holder males of this species display these behaviors simultaneously. A field study was carried out in which territorial nest holder males were either treated with long-lasting implants filled with 11-ketotestosterone (11-KT) or with control implants. Males treated with 11-KT showed a higher frequency of aggressive behavior, were more responsive to aggressive challenges, and were more persistent in aggressive behavior than control males. In addition, territories were larger in males treated with 11-KT than in controls. We found evidence for incompatibility between defense of a large territory and high levels of parental behavior. However, contrary to expectation, 11-KT did not suppress parental behavior. We suggest that trade-offs between territorial and parental behavior may not be regulated by androgen hormones but may result from a time constraint in the individual's activity budget.     

Strategies, Conflict, and the Emergence of Territoriality: The Case of the Maine Lobster Industry

In this article, we use game theory to understand the emergence of various kinds of territorial arrangements in the Maine lobster fishery during the past century. Using the Nash equilibria of models of the fishery as our theoretical framework, we show that informal territorial arrangements in this fishery went through three sequential stages. These stages are the result of decisions by groups of lobster fishermen to defend fishing areas or invade those of other groups. A large number of factors influence these defensive and offensive strategies: concentrations of lobsters, the adoption of better technology, transportation costs, ecological changes, trap monitoring costs, the ability to organize defensive and offensive groups, and better law enforcement---all of which are captured by crucial parameters of our model. We argue that this technique can be applied to elucidate territorial changes more generally.     

Elevated testosterone levels during nonbreeding-season territoriality in a fall-breeding lizard,Sceloporus jarrovi

Summary In many vertebrates, seasonal activation of sexual and territorial behaviors coincides with seasonal gonadal activation and is caused by the increase in sex steroid hormones. Both male and femaleSceloporus jarrovi are territorial, but in this species territorial behavior is seasonally activated in late April, months before seasonal gonadal maturation, which occurs in August prior to the fall mating season. Measurements of seasonal changes in circulating levels of the sex steroid hormones testosterone, progesterone, and estradiol indicated that testosterone levels in both sexes are elevated when territorial behavior is expressed, even during the period of nonbreeding-season territoriality during the summer. This suggests that a nonbreeding season behavior is activated by a sex steroid hormone in this species.     

Territorial intrusion risk and antipredator behaviour: a mathematical model

In territorial animals that hide to avoid predators, a predatory attack creates a conflict because a hiding animal cannot defend its territory from conspecific intruders. When intruders are persistent, a past conspecific intrusion informs a territorial resident that future intrusions by the same animal are likely. Using a mathematical model, I examine the effects that past territorial intrusions can have on antipredator behaviour. Past territorial intrusions rarely affect a resident animal's time to hide (the optimal behaviour is to hide as soon as the predator initiates its attack). In contrast, past intrusions should shorten the length of time during which territory holders remain in hiding, with the magnitude of this effect depending on the time of the predator's attack, the re-intruder's pattern of return, and the intrusion rates of other conspecifics. The results of the model show that we need more information on patterns of re-intruders' behaviour, and emphasize that a similar functional explanation could underlie other behavioural changes following territorial and/or aggressive encounters (such as winner/loser effects or changes in display frequency and territorial vigilance). Differences between my findings and those from previous studies suggest that the trade-off between antipredator behaviour and territorial defence can involve different costs from the trade-off between antipredator behaviour and foraging.     

Male dimorphism, territoriality and mating success in the tropical damselfly, Paraphlebia zoe Selys (Odonata: Megapodagrionidae)

The tropical damselfly Paraphlebia zoe has two male morphs: a black-winged (BW) male which is associated with territorial defense of oviposition sites; and a hyaline-winged (HW) male similar in appearance to females, and, compared to the black morph, less frequently found defending territories. In a wild population of this species, we first assessed the relationship between phenotypic traits [male morph, size and territorial status (being territorial or non-territorial)], their role on mating success, and the degree to which a particular territory may contribute to male mating success. Second, to relate a physiological basis of being territorial we compared both morphs in terms of muscular fat reserves and thoracic muscle, two key traits related to territory defense ability. Males of both morphs defended territories although the BW males were more commonly found doing this. BW males were larger than HW males and size predicted being territorial but only within HW males (territorial males were larger) but not in BW males. Male mating success was related to territorial status (territorial males achieved a higher mating success), but not to morph or size. Furthermore, territory identity also explained mating success with some territories producing more matings than others. The BW morph stored more fat reserves which may explain why this morph was more likely to secure and defend a place than the HW morph. However, the HW morph showed higher relative muscle mass which we have interpreted as a flexible strategy to enable males to defend a territory. These results are distant to what has been found in another male dimorphic damselfly, Mnais pruinosa, where the advantage of the non-territorial morph relies on its longevity to compensate in mating benefits compared to the territorial morph.     

基于生态安全格局理论的国土空间生态修复分区模拟——以粤港澳大湾区为例

国土空间生态修复分区有助于推动生态修复工作的整体性和系统性。利用国土空间生态安全格局理论进行生态修复分区是一种较为经典的模式,但现有分区技术方案对高强度人类活动导致的土地利用景观变化考虑不足,不能很好的适应城市群尺度国土空间生态安全格局的塑造需求。设计了一种基于情景模拟的国土空间生态修复分区方案,该框架首先通过耦合空间马尔科夫链与元胞自动机模型模拟城镇化发展末期的土地利用景观格局;然后利用"生态源地-生态廊道-生态网络"理论构建国土空间生态安全格局基本骨架;最后根据最小累积阻力模型划分出生态修复关键区、调节区和双修区等类型。粤港澳大湾区案例研究表明：（1）土地利用景观变化对生态修复分区方案影响较大,不同景观格局导致的分区规模差异可达国土空间的10%左右;（2）国土空间生态修复分区需要充分考虑高强度人类活动对景观格局变化的潜在影响,进而使得生态修复分区政策更有利于国土空间生态安全格局塑造目标的实现。研究构建的国土空间生态修复分区技术方案可进一步深化生态安全格局理论在国土空间规划应用上的认知。     

Reproductive behavior of the dragonfly,Orthetrum japonicum (Odonata: Libellulidae)

The reproductive behavior of the dragonfly,Orthetrum japonicum, is described. Behavioral processes of turnover of territorial males, simultaneous guarding of 2 females by a male, and copulation by non-territorial males are described. The males with longer hind wings won the territorial conflicts more frequently. The total duration of territorial residence of a given male was correlated with the number of his matings, but not correlated with the length of his abdomen or hind wings. The territorial site with the lower degree of vegetation cover was occupied by males more consistently. Males in more consistently occupied territorial sites did not have longer abdomen and hind wings than males in less consistently occupied sites. The territorial site where the larger number of copulations was observed was not occupied more consistently. Selection episode analysis using the method of Arnold & Wade (1984a, b) showed that direct selection on the hind wing length favored the short wing and that direct selection on the abdomen length favored the long abdomen during mating.     

Dual interaction producing both territorial and schooling behavior in fish

Fish schools are frequently observed without a leader and an explicit condition forming school. Several models of schooling have been proposed focusing on the influence of neighbors, and introducing probability distributions, while these models are based on the separation of schooling and territorial behavior. We frequently consider the duality of aggregation of animals, in which behavioral patterns involve both attraction and repulsion, antagonistic with each other. The idea of probability does not explain this duality that can provide both schooling and territorial behavior. From these biological facts, we have constructed a behavior model in which the influence of neighbors is formulated by the interface between the states of neighbors and a map of changes in these states. This interface uses a self-similar nowhere differentiable transition map which is temporally constructed, encompassing a crucial duality of repulsive and attractive forces hidden in the interaction among fishes. We tested it with computer simulations against the biological reality of schooling and territorial behavior. Due to the influence neighbors can have on duality, the same model can show both schooling behavior with a high degree of polarization and territorial behavior.     

Individual Variation in Advertisement Calls of Territorial Male Green Frogs, Rana clamitans: Implications for Individual Discrimination

Individuals of many territorial species discriminate between familiar territorial neighbors and unfamiliar strangers based on individual differences in acoustic signals. Many anuran amphibians are territorial and use primarily acoustic communication during social interactions, but evidence for acoustically mediated individual discrimination is available only for one species. As a first step in research designed to investigate individual discrimination in a second species of territorial frog, we examined patterns of within-male and among-male variability in 198 advertisement calls of 20 male green frogs, Rana clamitans . The aim was to determine which acoustic properties could be used by males to recognize their territorial neighbors and to estimate limits of reliability afforded by these properties for identifying different neighbors. All of the call properties that we examined exhibited significant inter-individual variation. Discriminant function analyses assigned between 52% and 100% of calls to the correct individual, depending on sample size and the call properties included in the model. This suggests that there is sufficient among-male variability to statistically identify individuals by their advertisement calls. The call properties of fundamental frequency and dominant frequency contributed the most towards discrimination among individuals. Based on their natural history and behavior and the results reported here, we suggest that male green frogs likely discriminate between strangers and adjacently territorial neighbors based on individual variation in advertisement calls.     

Boldness,Dominance, and Territoriality in the Color Polymorphic Tree Lizard,Urosaurus ornatus

Social dynamics in territorial species often reflect underlying variation in aggression and other aspects of social dominance among individuals. In ornate tree lizards (Urosaurus ornatus), males differing in dewlap color differ in social dominance: while blue males are the dominant, aggressive morph and always territorial, yellow males tend to exhibit a less‐aggressive satellite behavioral tactic. However, in habitats with fewer available territorial resources, yellow males defend territories and increase in relative abundance. These observations suggest that consideration of social dominance alone may be insufficient to explain U. ornatus' territorial dynamics in the wild. Here, we tested how both dominance and another important behavioral trait, boldness, contribute to the outcome of territorial disputes in tree lizards. We recorded the territorial behavior of blue and yellow male tree lizards (entered in pairs) in an experimental arena. At the end of each trial, we then approached each male and recorded whether it fled (shy) or not (bold) in response to our approach. As expected, dominant blue males exploited the higher quality perch more often than yellow males. However, when approached by a simulated predator, blue males were more likely to flee than yellow males. Thus, while blue males are more dominant, yellow is likely the bolder morph. As a result, this morph may be better equipped to defend territories in riskier environments. We conclude that although dominance asymmetries may predictably drive initial territorial interactions among competing males, variation in other behaviors (like boldness) may perturb the long‐term outcome of these interactions across variable environments.     

Evolution of territoriality in Hylinae treefrogs: Ecological and morphological correlates and lineage diversification

Given the diverse nature of traits involved in territorial defence, they may respond to different selective pressures and then exhibit distinct patterns of evolution. These selective pressures also may cause territorial behaviour to be associated with environmental and morphological variables. Such associations, however, have mostly been studied at the intraspecific level, being phylogenetic analyses of territoriality in a broad taxonomic framework rare in the literature. We used the anuran subfamily Hylinae to test (1) whether two territorial-behaviour traits with different levels of aggression—territorial call and physical combat—are evolutionarily more labile than a morphological trait used in physical combat—the spine-shaped prepollex; (2) whether reproduction in lentic waters and phytotelmata, as well as resource scarcity, might favour the occurrence of territoriality; (3) if physical combat is more important than territorial call for the evolution of body size and sexual size dimorphism and (4) the relationships between territorial-behaviour traits and lineage diversification. We mainly used the literature to build two datasets with different levels of certainty. Territorial-behaviour traits exhibited intermediate levels of phylogenetic signal in Hylinae, whereas the phylogenetic signal for the presence of the spine-shaped prepollex was strong. We found support for the hypothesis that reproduction in lentic water favours the occurrence of territorial behaviour, because the expression of territorial-behaviour traits was more associated with reproduction in lentic than in lotic waters. Territorial-behaviour traits were not correlated with annual precipitation nor with habitat complexity. Body size and sexual size dimorphism were not correlated with the presence of territorial call nor with physical combat. We identified negative correlations between diversification rates and physical combat. Relationships of territorial call and physical combat with diversification rates suggest that these territorial behaviours influence evolutionary processes in different ways.     

The asymmetric war of attrition

The paper, which has an informal discussion at the end, provides a game theoretical analysis of the asymmetric “war of attrition” with incomplete information. This is a contest where animals adopt different roles like “owner” and “intruder” in a territorial conflict, and where the winner is the individual prepared to persist longer. The term incomplete information refers to mistakes in the identification of roles. The idea by Parker & Rubenstein (1981) is mathematically worked out and confirmed that there exists only a single evolutionarily stable strategy (ESS) for the model with a continuum of possible levels of persistence and no discontinuities in the increase of cost during attrition. The ESS prescribes to settle the conflict according to “who has more to gain or less to pay for persistence”. The only evolutionarily stable convention is thus to give the player access to the resource who has the role which is favoured with respect to payoffs. By contrast, it was shown earlier (Hammerstein, 1981) for various asymmetric versions of the “Hawks-Doves” model that an ESS can exist which appears paradoxical with respect to payoffs. The nature of this contrast is further analyzed by introducing elements of discreteness in the asymmetric war of attrition. It turns out that some conditions must be satisfied in order to have the possibility of an alternative ESS which is not of the above simple commonsense type. First, a decision to persist (or escalate) further in a contest must typically commit a contestant to go on fighting for a full “round”, before he can give up without danger. Second, such a “discontinuity” must occur at a level of persistence where the contest is still cheap, and, finally, errors in the identification of roles must be rare.     

Changes in the sexual tendency accompanying selection for aggressiveness in the three-spined stickleback,Gasterosteus aculeatus L.*

Independent selection experiments for reduced juvenile and reduced territorial aggression levels in male three-spined sticklebacks both resulted in reduced courtship aggression levels. In the low juvenile aggression line this is accompanied by an enhanced sexual activity, but in the low territorial aggression line males display a reduced sexual activity. These results are consistent with a mutually inhibitory relationship between the aggressive and sexual tendencies, if selection for juvenile aggressiveness has only affected the aggressive tendency while selection for territorial aggressiveness has simultaneously affected the aggressive and sexual tendencies in the same direction. This is in agreement with the hormonal changes (pituitary-gonadal axis) suggested by the selection studies.     

Factors Affecting Interspecific Aggression in a Year‐Round Territorial Species,the Jewel Damselfish

Vertebrates live in complex species networks in which interspecific interactions are common. In some contexts, the aggressive behaviours shown in these interspecific interactions are very similar to those shown in intraspecific interactions. It is still an open question whether intra‐ and interspecific aggression share common causality. We studied a year‐round territorial species the jewel damselfish, (Plectroglyphidodon lacrymatus), which cultivate algae they feed on. Territory holders aggressively defend these algae that are an attractive resource for many other species. In this study, we recorded territorial aggression in free‐living individuals and recorded aggressive responses to a standardized territorial intrusion test in captive individuals. Field observations indicated that territorial aggression was selectively targeted towards food competitors. Independent of the size of the species, aggression was more frequent towards common species around their territories. This relationship was confirmed experimentally by confronting the jewel damselfish with novel objects to which the subjects were exposed either frequently or rarely. We suggest that jewel damselfish have to learn which species are competitors and therefore should be chased. In a standardized intrusion test with captive individuals, no significant differences were found in territorial responses towards intra‐ or interspecific intruders. Neither territorial aggression nor the intrusion showed any relationship with plasma androgen levels. Together, these data suggest that experience might be more important in non‐seasonal territorial aggression than circulating hormonal factors.     

Effects of age,territoriality and breeding on survival of Bonelli’s Eagle Aquila fasciata

Survival typically contributes most to population trends in long‐lived birds and its accurate estimation is therefore vital for population management and conservation. We evaluated the effects of age, territoriality and reproduction on survival in Bonelli’s Eagle Aquila fasciata through multistate capture‐mark‐recapture analyses on a long‐term dataset. Monitoring was carried out in southeast France (1990–2008) and involved the surveying of territorial Eagles, the marking of fledged chicks, and the recording of resightings and recoveries of marked non‐territorial and territorial birds. Survival improved with age, but territoriality was not retained in the best model; yearly survival was estimated at 0.479 for fledglings (to 1 year of age), 0.570 for 1‐ and 2‐year‐olds, and 0.870 for 3‐year‐old and older individuals. The second best model supported a further increase in survival from 3‐year‐olds (0.821) to older individuals (0.880). In the third best supported model, territoriality enhanced survival, but only in 2‐year‐olds (0.632 vs. 0.562 for non‐territorial). We found no correlation between the previous breeding stage and future survival, consistent with the long lifespan of the study species. Nevertheless, 4‐year‐old and older successful breeders were more likely to breed the following year than failed adult breeders (0.869 vs. 0.582), suggesting that the cost of reproduction is small in comparison with the variation in quality among individuals or their territories.