The ecological and evolutionary stability of interspecific territoriality

Interspecific territoriality may play an important role in structuring ecological communities, but the causes of this widespread form of interference competition remain poorly understood. Here, we investigate the phenotypic, ecological and phylogenetic correlates of interspecific territoriality in wood warblers (Parulidae). Interspecifically territorial species have more recent common ancestors and are more similar phenotypically, and are more likely to hybridise, than sympatric, non‐interspecifically territorial species. After phylogenetic corrections, however, similarity in plumage and territorial song are the only significant predictors of interspecific territoriality besides syntopy (fine‐scale geographic overlap). Our results do not support the long‐standing hypothesis that interspecific territoriality occurs only under circumstances in which niche divergence is restricted, which combined with the high incidence of interspecific territoriality in wood warblers (39% of species), suggests that this interspecific interaction is more stable, ecologically and evolutionarily, than commonly assumed.     

Multiple routes to interspecific territoriality in sister species of North American perching birds

Behavioral interference between species can influence a wide range of ecological and evolutionary processes. Here, we test foundational hypotheses regarding the origins and maintenance of interspecific territoriality, and evaluate the role of interspecific territoriality and hybridization in shaping species distributions and transitions from parapatry to sympatry in sister species of North American perching birds (Passeriformes). We find that interspecific territoriality is pervasive among sympatric sister species pairs, and that interspecifically territorial species pairs have diverged more recently than sympatric noninterspecifically territorial pairs. None of the foundational hypotheses alone explains the observed patterns of interspecific territoriality, but our results support the idea that some cases of interspecific territoriality arise from misdirected intraspecific aggression while others are evolved responses to resource competition. The combination of interspecific territoriality and hybridization appears to be an unstable state associated with parapatry, whereas species that are interspecifically territorial and do not hybridize are able to achieve extensive fine- and coarse-scale breeding range overlap. In sum, these results suggest that interspecific territoriality has multiple origins and impacts coexistence at multiple spatial scales.     

Increased Response of the Short-Toed Treecreeper Certhia brachydactyla in Sympatry to the Playback of the Song of the Common Treecreeper C. familiaris

The two European species of treecreepers, the short-toed treecreeper Certhia brachydactyla and the common treecreeper C. familiaris, occupy different habitats, but are sympatric to a certain extent. The response to specific and heterospecific song was studied both in a sympatric and in an allopatric population of short-toed treecreeper in the Iberian Mountains (Burgos, Castilla-León, Spain). Short-toed treecreepers in sympatry showed a more aggressive response towards the song of the common treecreeper than those in allopatry. Although strict interspecific territoriality was not found, the experimental data suggest that the altitudinal distribution pattern of the two species may be a consequence of the increased aggression. This interspecific aggression may be advantageous for the short-toed treecreeper, as it may achieve a loose interspecific territoriality resulting in some spacing. However, a non-adaptive hypothesis for this behaviour cannot be rejected.     

Resource control by territorial male cichlid fish in Lake Malawi

1. The rocky habitat communities of Lake Malawi contain a high diversity of ecologically similar, predominantly herbivorous, cichlid fish species. How so many species can coexist is still unknown.
2. Adult males of the majority of these species hold permanent territories which form dense multispecies mosaics across the shores.
3. The study examined six coexisting species of cichlids from the rocky habitat where males are known to control access to the algal food resources within their territorial space. These included four sibling species of the Pseudotropheus ( Tropheops ) species complex, which are specialized epilithic algal herbivores, and two sibling species of the trophically more generalist P . ( Maylandia ) species complex which feed upon both epilithic algae and plankton.
4. The hypotheses that both intraspecific and interspecific territoriality occur in these communities and that interspecific aggression initiated by territorial males is preferentially directed at species with the most similar diets to their own were tested.
5. It was found that males preferred to exclude conspecific intruders, while they tolerated intruders with different diets to their own. Also, male P . ( Tropheops ) preferentially excluded similar heterospecific dietary specialists, while male P. ( Maylandia ) tolerated similar heterospecific dietary generalists.
6. Based on this study, it is proposed that interspecific territoriality may be reducing competition between species with different diets and promoting their coexistence, while it may be enhancing competition between species with the same diet. Furthermore, it suggests that ecological generalism may be reducing the intensity of interspecific competition, while specializations for the same resources may be increasing competition.     

Lack of habitat segregation and no interspecific territoriality in three syntopic cryptic species of the golden‐spectacled warblers Phylloscopus (Seicercus) burkii complex

It is widely accepted that exploitative competition prevents the coexistence of any two or more closely‐related species unless differences exists in their ecological niches and resource use. In sibling bird species, exploitative competition is reduced mainly by spatial segregation of competing species. Spatial segregation can be achieved in two basic ways: by using different habitats or microhabitats that each species is predominately exploiting, or by interspecific territoriality. To our knowledge, either habitat segregation or interspecific territoriality or both have been found in all dyads and groups of sympartic sibling bird species studied so far. In this study, we investigated the coexistence of three cryptic sibling species of the golden‐spectacled warblers Phylloscopus burkii complex in Hunan Province, China: Ph. tephrocephalus, Ph. omeiensis and Ph. valentini. We analyzed their habitat preferences, and spatial interrelations between the species. In order to analyze either presence or absence of interspecific territoriality, we additionally performed con‐ and heterospecific playback experiments. Contrary to expectations, we found no evidences for either habitat segregation or interspecific territoriality in these three species.     

Interspecific aggressive behaviour by long-toed lapwings (Vanellus crassirostris)

In this paper interspecific aggression is related to ecological relationships between bird species. Three kinds of relationships are distinguished according to the presence or absence of aggression and the degree to which resources are exclusively utilized: interspecific territoriality (aggression, exclusive use), partial exclusion (aggression, no exclusive use), and tolerance (no aggression, no exclusive use). Aggression by long-toed lapwings (Vanellus crassirostris) is examined within this framework, using quantitative data gathered through observational sampling. Lapwings were interspecifically territorial with one species, partially excluded five species, and tolerated four species. The results indicate that the function of interspecific aggressive behaviour may be related to competition, predation, or both.     

Interspecific aggression,not interspecific mating,drives character displacement in the wing coloration of male rubyspot damselflies (Hetaerina)

Traits that mediate intraspecific social interactions may overlap in closely related sympatric species, resulting in costly between-species interactions. Such interactions have principally interested investigators studying the evolution of reproductive isolation via reproductive character displacement (RCD) or reinforcement, yet in addition to reproductive interference, interspecific trait overlap can lead to costly between-species aggression. Previous research on rubyspot damselflies (Hetaerina spp.) demonstrated that sympatric shifts in male wing colour patterns and competitor recognition reduce interspecific aggression, supporting the hypothesis that agonistic character displacement (ACD) drove trait shifts. However, a recent theoretical model shows that RCD overshadows ACD if the same male trait is used for both female mate recognition and male competitor recognition. To determine whether female mate recognition is based on male wing coloration in Hetaerina, we conducted a phenotype manipulation experiment. Compared to control males, male H. americana with wings manipulated to resemble a sympatric congener (H. titia) suffered no reduction in mating success. Thus, female mate recognition is not based on species differences in male wing coloration. Experimental males did, however, experience higher interspecific fighting rates and reduced survival compared to controls. These results greatly strengthen the case for ACD and highlight the mechanistic distinction between ACD and RCD.     

Reproductive interference explains persistence of aggression between species

Interspecific territoriality occurs when individuals of different species fight over space, and may arise spontaneously when populations of closely related territorial species first come into contact. But defence of space is costly, and unless the benefits of excluding heterospecifics exceed the costs, natural selection should favour divergence in competitor recognition until the species no longer interact aggressively. Ordinarily males of different species do not compete for mates, but when males cannot distinguish females of sympatric species, females may effectively become a shared resource. We model how reproductive interference caused by undiscriminating males can prevent interspecific divergence, or even cause convergence, in traits used to recognize competitors. We then test the model in a genus of visually orienting insects and show that, as predicted by the model, differences between species pairs in the level of reproductive interference, which is causally related to species differences in female coloration, are strongly predictive of the current level of interspecific aggression. Interspecific reproductive interference is very common and we discuss how it may account for the persistence of interspecific aggression in many taxonomic groups.     

Intra- and interspecific aggression do not modulate androgen levels in dusky gregories,yet male aggression is reduced by an androgen blocker

Discussions about social behavior are generally limited to fitness effects of interactions occurring between conspecifics. However, many fitness relevant interactions take place between individuals belonging to different species. Our detailed knowledge about the role of hormones in intraspecific interactions provides a starting point to investigate how far interspecific interactions are governed by the same physiological mechanisms. Here, we carried out standardized resident–intruder (sRI) tests in the laboratory to investigate the relationship between androgens and both intra- and interspecific aggression in a year-round territorial coral reef fish, the dusky gregory, Stegastes nigricans. This damselfish species fiercely defend cultivated algal crops, used as a food source, against a broad array of species, mainly food competitors, and thus represent an ideal model system for comparisons of intra-and interspecific territorial aggression. In a first experiment, resident S. nigricans showed elevated territorial aggression against intra- and interspecific intruders, yet neither elicited a significant increase in androgen levels. However, in a second experiment where we treated residents with flutamide, an androgen receptor blocker, males but not females showed decreased aggression, both towards intra- and interspecific intruders. Thus androgens appear to affect aggression in a broader territorial context where species identity of the intruder appears to play no role. This supports the idea that the same hormonal mechanism may be relevant in intra- and interspecific interactions. We further propose that in such a case, where physiological mechanisms of behavioral responses are found to be context dependent, interspecific territorial aggression should be considered a social behavior.     

Differential Water Mite Parasitism,Phenoloxidase Activity,and Resistance to Mites Are Unrelated across Pairs of Related Damselfly Species

Related host species often demonstrate differences in prevalence and/or intensity of infection by particular parasite species, as well as different levels of resistance to those parasites. The mechanisms underlying this interspecific variation in parasitism and resistance expression are not well understood. Surprisingly, few researchers have assessed relations between actual levels of parasitism and resistance to parasites seen in nature across multiple host species. The main goal of this study was to determine whether interspecific variation in resistance against ectoparasitic larval water mites either was predictive of interspecific variation in parasitism for ten closely related species of damselflies (grouped into five “species pairs”), or was predicted by interspecific variation in a commonly used measure of innate immunity (total Phenoloxidase or potential PO activity). Two of five species pairs had interspecific differences in proportions of individuals resisting larval Arrenurus water mites, only one of five species pairs had species differences in prevalence of larval Arrenurus water mites, and another two of five species pairs showed species differences in mean PO activity. Within the two species pairs where species differed in proportion of individuals resisting mites the species with the higher proportion did not have correspondingly higher PO activity levels. Furthermore, the proportion of individuals resisting mites mirrored prevalence of parasitism in only one species pair. There was no interspecific variation in median intensity of mite infestation within any species pair. We conclude that a species’ relative ability to resist particular parasites does not explain interspecific variation in parasitism within species pairs and that neither resistance nor parasitism is reflected by interspecific variation in total PO or potential PO activity.     

Differences in chemical signals may explain species recognition between an island lizard, Podarcis atrata, and related mainland lizards, P. hispanica

Chemical signals can be the basis of interspecific recognition and speciation in many animals. The Columbretes Islands wall lizard, Podarcis atrata is very close genetically to the mainland Iberian wall lizard Podarcis hispanica. However, a previous study suggested that chemosensory interspecific recognition would avoid reproductive interactions and hybridization between these two species. These results suggested that chemicals used in intraspecific communication might differ in composition and/or proportions between these two species. In this paper, we used gas chromatography–mass spectrometry (GC–MS) to characterize the chemical composition of the lipophilic fraction from femoral gland secretions of male P. atrata and P. hispanica. The analysis showed that chemicals found in femoral secretions varied in composition and proportions between species and between populations. Seven steroids and two unidentified waxy esters, were exclusive of P. atrata lizards from the islands. In contrast, nine steroids and other six compounds were only found in mainland P. hispanica. There were also differences in proportions of shared compounds between species. Moreover, all these differences were higher between P. atrata and P. hispanica than between any population of P. hispanica. Chemical differences might be consequence of genetic differences, but they could also be explained by adaptation to different habitats with different climatic conditions or diet resources. Compounds that are specific of each species, or differences in the pattern of compounds, could explain species recognition. Therefore, these results of chemical composition and previous studies of chemosensory recognition reinforce the fact that the genetic differences between P. hispanica and P. atrata may result in an effective reproductive isolation between these two taxa.     

Interspecific territoriality in gibbons (Hylobates lar and H. pileatus) and its effects on the dynamics of interspecies contact zones

We investigated the ecology and interspecific interactions of the two gibbon species (Hylobates lar and H. pileatus) that overlap in distribution within a narrow zone of contact in the headwaters of the Takhong River at Khao Yai National Park in central Thailand. The zone is about 10-km wide, with phenotypic hybrids comprising 6.5% of the adult population. We compared species with respect to diet, territory size, intra- and interspecific group encounters, and territory quality. The two gibbon species exploited the same types of resources within their territories despite variation in the relative abundance of food-plant species between territories. The gibbons were interspecifically territorial, and males of both species displayed aggressive behaviors at both intraspecific and interspecific territorial boundaries. There was no marked difference in the amount of overlap between territories of conspecific and heterospecific pairs of groups. Although the habitat was not homogeneous, territory quality did not vary significantly between species. The species have not diverged in habitat preference or in diet. Neither species dominated in interspecific encounters, and both were reproducing well in the contact zone. We analyzed the potential consequences of several types of interspecific interactions on individual dispersal options and on the structure of the contact zone. Interference competition through interspecific territoriality affects the dispersal of individuals into the range of the other species. In general, territorial competition coupled with limited hybridization leads to predictions of a narrow contact zone or parapatry between species; thus, behavioral and ecological interactions between species need to be considered as potential factors in explaining range borders of primate species.     

Species cohesion of an extremophyte (Carex angustisquama,Cyperaceae) in solfatara fields maintained under interspecific natural hybridization

Background and AimsHybridization is the main driver of plant diversification, and gene flow via hybridization has multifaceted effects on plant evolution. Carex angustisquama is an extremophyte that grows on soils heavily acidified by volcanism. Despite its habitat distinct from that of other species, this species is known to form interspecific hybrids, implying interspecific gene flow. It is crucial to verify the extent and direction of interspecific gene flow between C. angustisquama and closely related species to understand the evolutionary process of an extremophyte in solfatara fields.MethodsIn this study, expressed sequence tag–simple sequence repeat markers were utilized to infer the extent and direction of interspecific gene flow between C. angustisquama and closely related species.Key ResultsBayesian clustering and simulation analyses revealed that all individuals of the three hybrid species were classified into the first hybrid generation or first backcross to C. angustisquama; therefore, current interspecific gene flow is limited. Moreover, in the Bayesian inference of historical gene flow based on multispecies samples, the model that assumed no interspecific gene flow was the most strongly supported across all species pairs, including phylogenetically close but ecologically distinctive species pairs.ConclusionsOur results revealed that interspecific gene flow between C. angustisquama and its related species has been limited both currently and historically. Moreover, our results of Bayesian inference of historical gene flow indicated that extrinsic, rather than intrinsic, factors probably act as isolating barriers between Carex species, with hybrid breakdown via microhabitat segregation being the probable potential barrier. Overall, our findings provide insights into the evolutionary process of an extremophyte in solfatara fields and offer an important example of the mechanisms of diversification of the speciose genus Carex.     

Socioecological Factors Affecting Range Defensibility Among Howler Monkeys

Range defensibility is defined as the ability of animals to efficiently move over an area to monitor and defend it. Therefore, range defensibility can help us understand the spatial structure of animal territoriality. We used howler monkeys (Alouatta spp.), a genus for which no agreement on the extent of their territoriality exists, to investigate the factors mediating range defensibility. We compared the defensibility index (D) across 63 groups of howler monkeys, representing 8 different species, based on a literature review. All species, except Alouatta palliata, were classified as potentially territorial according to D, although there was high variability within and among species. Group size had a positive effect on D, probably owing to the greater ability of groups to defend a territory as they become larger. Study area had a negative effect on D, perhaps suggesting that unlike small areas, large areas allow groups to have territories that do not require significant defense from neighbors. However, population density was the factor with the strongest effect on D, with greater monitoring of home ranges under high levels of competition. Our results suggest that howler monkeys are theoretically capable of maintaining a territory and suggest that animals can show a gradient in territoriality, which can be mediated by the competitive context in which it occurs.     

Coexistence in Mediterranean warblers: ecological differences or interspecific territoriality?

We studied the coexistence of four species of Sylvia warblers living in Mediterranean matorral in order to identify the respective role of ecological segregation and of interspecific territoriality in explaining the local distribution of these four species. Data on habitat use, foraging behaviour and interspecific spacial segregation were collected on Corsica and on Spargi (Sardinia) islands. Despite large overlap in patch selection and in foraging behaviour the four species did segregate ecologically and behaviourally while foraging (differences in the choice of plant species used for foraging, in the height of the plant selected, in the selection of the portion of the plant volume used and in the selection of the plant structure explored). Complementarity in foraging behaviour was observed in the morphologically and ecologically closest species: the Dartford ( Sylvia undata ) and the Marmora's ( Sylvia sarda ) warblers. We did not observe any evidence of direct interspecific interactions in song, alarm, or aggressive behaviour. Nor did we observe patterns of spatial distribution that would support the idea of coexistence by interspecific territorial exclusion. These results contrast with the results of Cody & Walter (1976) suggesting interspecific territoriality in Mediterranean Sylvia warblers. They are consistent with other published results emphasizing ecological differences as explanation for species coexistence.     

An Extreme Case of Interspecific Territoriality: Male Anthidium manicatum (Hymenoptera,Megachilidae) Wound and Kill Intruders

Anthidium manicatum males defend patches of flowering plants as mating territories and copulate with females that enter the territory to obtain nectar or pollen. Territorial males defend their territories not only against conspecific males but also against many other insect species that attempt to utilize the flowers inside the territory. In our study area, a territorial male defended its territory against conspecific males on average only twice per h but it attacked other species of insects on average 70 times per h. Territorial males distinguished between different species of intruders and attacked them unequally. During the hour following experimental removal of the territory holder the number of intruders of some insect species tripled but the number of intruders of those species that were rarely attacked remained approximately the same. A. manicatum males, but not females, bear long spines on the last two segments of the abdomen. Territorial males rammed intruding insects at high speed and shortly before the moment of impact they curved the abdomen forward to hit with the spines. Intruders could be seriously damaged or killed by these attacks. Abdominal spines of A. manicatum males may have evolved as weapons to increase the effectiveness of interspecific territoriality.     

Spatial arrangement and diet overlap between colonies of desert ants

Summary The spatial patterns and diets of three desert ant species were examined. The results indicate that food competition may account for the spatial arrangement of these species, and that only intraspecific interactions may be required. Each ant species was significantly overdispersed, and the average intraspecific nearest neighbor distances were greater than the interspecific nearest neighbor distances. A test of pairwise spatial arrangment showed that all three species pairs were aggregated interspecifically. The level of the interspecific aggregation was related to the diet similarity of the species. The two species pairs with the lowest diet overlaps were significantly aggregated, and the species pair with the most similar diets was not significantly aggregated. Pairwise dietary overlaps between colonies showed that average intraspecific overlaps were significantly greater than interspecific diet overlaps. Furthermore, the diet overlap was significantly positively correlated to the mean nearest neighbor distance for the three intraspecific and three interspecific comparisons. These data indicate competition for food, especially within species, may be regulating the intercolony distances of these ant species. A computer simulation tested whether only intraspecific territoriality is necessary to produce the observed nearest neighbor distances. A simulation that placed colonies randomly on a patch confirmed that these colonies are intraspecifically overdispersed. By adding intraspecific territoriality, the simulation nearest neighbor distances fit the empirical data reasonably well. Thus interspecific competitive interactions seem unnecessary to account for the spatial arrangement of these species.     

Discrimination of cis-trans sex pheromone components in two sympatric Lepidopteran species

Pheromone-binding proteins (PBPs) play an important role in the recognition of pheromones by insects. However, the abilities of these PBPs to discriminate pheromone components and recognize the isomers are unclear. Dendrolimus houi and Dendrolimus kikuchii are two sympatric coniferous pests whose pheromones have cis-trans isomers. We used these insect species to detect the precise recognition abilities of PBPs. The four PBPs examined showed male-biased antenna-intensive expression patterns, whereas PBP1 showed higher expression than PBP2 in the antenna. DhouPBP1 only bound to a minor interspecific pheromone component, whereas DhouPBP2 bound to all three intraspecific components and another minor interspecific component. DkikPBP1 and DkikPBP2 could recognize all three intraspecific components with affinities negatively correlated with their ratios, and they bound to interspecific pheromones with affinity that was positively correlated with the ratios. The four PBPs have different cis-trans isomer discrimination abilities, i.e., DhouPBP1 and DkikPBP1 could not discriminate the two cis-trans isomer pairs of pheromones from the two species, whereas DhouPBP2 could discriminate between both pairs, and DkikPBP2 could only discriminate one pair. Overall, PBPs from D. houi and D. kikuchii use different strategies to help the moths to discriminate the intra- and interspecific pheromone components. Our work will contribute to better understanding of the sex pheromone recognition mechanism in these two sister species of moths and provide insights into more effective management practices of these pest species.     

Observations on Male Territories in a Carpenter Bee,Xylocopa (Neoxylocopa) hirsutissima Maidl (Hymenoptera,Anthophoridae)

Observations were made on territoriality of ♂♂ of the Brazilian carpenter bee Xylocopa hirsutissima. These territories are found on mountain tops where the ♂♂ hover from 4–6 p.m. in close vicinity of an optical beacon, being a protruding shrub or little tree. Territories are defended against other ♂♂Production of a pheromone by the mandibular glands is supposed. This territoriality as well as that of other species of the genus is considered to be an important aspect of the mating biology.     

Altitudinal gradients,plant hybrid zones and evolutionary novelty

Altitudinal gradients are characterized by steep changes of the physical and biotic environment that present challenges to plant adaptation throughout large parts of the world. Hybrid zones may form where related species inhabit different neighbouring altitudes and can facilitate interspecific gene flow and potentially the breakdown of species barriers. Studies of such hybrid zones can reveal much about the genetic basis of adaptation to environmental differences stemming from changes in altitude and the maintenance of species divergence in the face of gene flow. Furthermore, owing to recombination and transgressive effects, such hybrid zones can be sources of evolutionary novelty. We document plant hybrid zones associated with altitudinal gradients and emphasize similarities and differences in their structure. We then focus on recent studies of a hybrid zone between two Senecio species that occur at high and low altitude on Mount Etna, Sicily, showing how adaptation to local environments and intrinsic selection against hybrids act to maintain it. Finally, we consider the potential of altitudinal hybrid zones for generating evolutionary novelty through adaptive introgression and hybrid speciation. Examples of homoploid hybrid species of Senecio and Pinus that originated from altitudinal hybrid zones are discussed.