Mimicry as a novel pathway linking biodiversity functions and individual behavioural performances

The feedback of biodiversity on individual trait variation is a poorly explored mechanistic pathway in ecological research. We analysed the relationship between biodiversity and individual performance by focusing on vocal mimicry, a widespread interaction that may serve in intra- and interspecific communication. We studied the songs of two lark species (genus Galerida) that increase the complexity of their song displays by mimicking other birds, and analysed the influence of bird species richness on individual song performance. The diversity of mimicked species and the prevalence of mimicry increased in areas characterized by great α and γ diversity (i.e. where larks experience more diverse encounters with community members, many of them being highly vocal owing to breeding). Conversely, the variability in species-specific song components peaked where larks were abundant, probably matching the complexity of conspecific social milieu. Some trade-offs existed between homo- and heterospecific complexity, suggesting that larks could change from population- to community-driven song variation by tracking the composition of the auditory environment. Mimicry, which serves communication with conspecifics or predators, may mediate interactions, ultimately cascading to aspects of ecological diversity other than those promoting its complexity.     

Limited Phylogeographic Signal in Sex-Linked and Autosomal Loci Despite Geographically,Ecologically, and Phenotypically Concordant Structure of mtDNA Variation in the Holarctic Avian Genus Eremophila

Phylogeographic studies of Holarctic birds are challenging because they involve vast geographic scale, complex glacial history, extensive phenotypic variation, and heterogeneous taxonomic treatment across countries, all of which require large sample sizes. Knowledge about the quality of phylogeographic information provided by different loci is crucial for study design. We use sequences of one mtDNA gene, one sex-linked intron, and one autosomal intron to elucidate large scale phylogeographic patterns in the Holarctic lark genus Eremophila. The mtDNA ND2 gene identified six geographically, ecologically, and phenotypically concordant clades in the Palearctic that diverged in the Early - Middle Pleistocene and suggested paraphyly of the horned lark (E. alpestris) with respect to the Temminck''s lark (E. bilopha). In the Nearctic, ND2 identified five subclades which diverged in the Late Pleistocene. They overlapped geographically and were not concordant phenotypically or ecologically. Nuclear alleles provided little information on geographic structuring of genetic variation in horned larks beyond supporting the monophyly of Eremophila and paraphyly of the horned lark. Multilocus species trees based on two nuclear or all three loci provided poor support for haplogroups identified by mtDNA. The node ages calculated using mtDNA were consistent with the available paleontological data, whereas individual nuclear loci and multilocus species trees appeared to underestimate node ages. We argue that mtDNA is capable of discovering independent evolutionary units within avian taxa and can provide a reasonable phylogeographic hypothesis when geographic scale, geologic history, and phenotypic variation in the study system are too complex for proposing reasonable a priori hypotheses required for multilocus methods. Finally, we suggest splitting the currently recognized horned lark into five Palearctic and one Nearctic species.     

Interspecific interactions drive cultural co-evolution and acoustic convergence in syntopic species

1.?Antagonistic interactions have been favourite subjects of studies on species co-evolution, because coexistence among competing species often results in quantifiable character displacement. A common output for competitive interactions is trait divergence, although the opposite phenomenon, convergence, has been proposed to evolve in some instances, for example in the communication behaviour of species that maintain mutually exclusive territories. 2.?I use here experimental and observational evidence to study how species interactions drive heterospecific signal convergence and analyse how convergence feeds back to the interaction itself, in the form of aggressive behaviour. I recorded the learned territorial signals of two non-hybridizing larks, Galerida cristata and G.?theklae, and used allopatric populations as controls for evaluating acoustic convergence in syntopy. Acoustic variation was analysed with respect to social conditions controlling for other potential agents of natural selection, habitat and climate. 3.?Interspecific convergence of Galerida calls peaked in syntopy. Although call acoustic structure was affected by climate and habitat, it matched gradients of density and proximity to congeners even at small local scales. The process of cultural transmission, in which individuals may acquire components of behaviour by copying neighbours, enhances the correlation between call acoustics and the local social milieu. 4.?Territories were defended against both species, but playback stimuli of convergent congener calls elicited a stronger aggressive reaction than congener calls from allopatric locations. 5.?This study shows that learned behaviours may co-evolve as a consequence of antagonistic interactions, determining reciprocal cultural evolution or cultural co-evolution. As for (biological) co-evolution, the distribution of competing species influences whether a particular area becomes a syntopic environment in which convergence is occurring, or an allopatric environment lacking interactions and reciprocal change. Because of their plastic nature, cultural coadaptations may rapidly shift in response to fluctuating social selection, thus propelling dynamic interactions and fine adjustments to the local environment.     

Inter-specific association and habitat use in a farmland passerine assemblage

We studied the pattern of inter-specific association of breeding territories in a passerine assemblage of dry cereal farmland in central Spain and evaluated the role of the presence of heterospecifics in the habitat use patterns exhibited by different species. Bird territories showed a non-random inter-specific spatial aggregation pattern. We studied territory abundance variation in the three more abundant species: the corn bunting, the crested lark, and the fan-tailed warbler. Crested lark and fan-tailed warbler territories were more abundant in plots where corn bunting territories were present and vice versa, while their respective abundances did not vary with the breeding presence of the other species. We used landscape and agricultural management variables to analyze the relationships between habitat and each species?? breeding territories by means of classification trees. While the corn bunting showed a marked pattern of nesting habitat use, the crested lark and the fan-tailed warbler exhibited a much more generalist one. Corn Bunting presence was affected negatively by intensification-related variables, such as field size and percent cover of cereal crops. Similarly, the presence of crested larks was negatively related to high yielding areas. However, when the presence of hetero-specific territories was considered, the presence of corn bunting territories was the most important variable explaining the occurrence of breeding fan-tailed warblers, and the second most important in the case of the crested lark. These results suggest that inter-specific attraction could play a role in the formation of farmland bird assemblages, while adding further evidence for the detrimental effect of agricultural intensification at the community level.     

Rapid phenotypic change in a native bird population following conversion of the Colorado Desert to agriculture

Humans are modifying our planet's ecosystems with increasing frequency and intensity. Exploring population responses to anthropogenic modifications of natural habitat provides insights into how species persist in the Anthropocene. Here, we leverage natural history collections to document rapid phenotypic change within a native bird population following ~80 yr of agriculture in the Colorado Desert of southeastern California. By comparing spectrophotometric measurements of horned lark Eremophila alpestris specimens collected in the Imperial Valley from 1918 to 1934 to those collected from 1984 to 2014, we found that more recent birds have darker backs, napes, and crowns. This dorsal darkening may have resulted from a shift in selective pressures for camouflage induced by land use: previously, the lark population nested on light‐colored desert flats, whereas contemporary larks occupy darker soil associated with agricultural fields. Adaptation and/or introgression may have contributed to this instance of rapid phenotypic change following the rise of agriculture in the Imperial Valley.     

Physiological adjustments to arid and mesic environments in larks (Alaudidae)

Because deserts are characterized by low food availability, high ambient temperature extremes, and absence of drinking water, one might expect that birds that live in these conditions exhibit a lower basal metabolic rate (BMR), reduced total evaporative water loss (TEWL), and greater ability to cope with high air temperatures than their mesic counterparts. To minimize confounding effects of phylogeny, we compared the physiological performance of four species of larks at ambient temperatures (T(a)'s) ranging from 0 degrees to 50 degrees C: hoopoe larks (Alaemon alaudipes) and Dunn's larks (Eremalauda dunni) live in hot and dry deserts, whereas skylarks (Alauda arvensis) and woodlarks (Lullula arborea) occur in temperate mesic areas. Mass-adjusted BMR and TEWL were indistinguishable between hoopoe lark and Dunn's lark and between skylark and woodlark. When grouping the data of the two desert larks in one set and the data of the two mesic larks in another, desert larks are shown to have 43% lower BMR levels and 27% lower TEWL values than the mesic species. Their body temperatures (T(b)'s) were 1.1 degrees C lower, and the minimal dry heat transfer coefficients (h) were 26% below values for the mesic larks. When T(a) exceeded T(b), the h of hoopoe larks and Dunn's larks was high and indistinguishable from h at 40 degrees C, in contrast to the prediction that h should be decreased to minimize heat gain through conductance, convection, or radiation from the environment when T(a) exceeds T(b).     

Mismatches between habitat preferences and risk avoidance for birds in intensive Mediterranean farmland

Land use intensification may create habitats that organisms perceive as suitable, but where reproduction or survival is insufficient to maintain self-sustaining populations. Such conditions may qualify as ecological traps, but their existence is often hard to prove. Here, we provide a practical framework to evaluate a potential ecological trap resulting from mismatch between habitat preferences and predation risk, focusing on ground-nesting farmland birds of conservation concern. The framework is based on species-specific associations with safe or unsafe habitat types (i.e. field and landscape types with high or low nest survival), and the occurrence of risk avoidance (i.e. negative responses to predator abundances or to nest failure rates after controlling for habitat effects). Bird densities were far more influenced by field characteristics than landscape context. Corn bunting and fan-tailed warbler were associated with tall swards (safe habitats), and did not show risk avoidance. Tawny pipit and and Galerida larks were associated with short swards (unsafe habitats), with the former avoiding fields with high nest predation rates, and the later avoiding high mongoose abundances. Short-toed lark was associated with fields with short swards and low nest trampling rates. Results suggest that short-toed lark may be the most vulnerable to ecological trapping, because it nests on unsafe habitats and did not show predation risk avoidance. Our approach provides a practical first step to infer vulnerability to a potential ecological trap, though further research is needed to confirm this effect. Management actions increasing nest survival in short sward fields will likely favour grassland bird conservation in intensive Mediterranean farmland.     

The geographic and seasonal dimensions of habitat use in Galerida larks: implications for species coexistence and range limits

In this paper, we investigated the patterns of habitat use in three sibling species of larks: the Thekla lark Galerida theklae which is sympatric with either of two parapatric species of crested larks (G. macrorhyncha replacing G. cristata in arid areas of Morocco). Specifically, we addressed the following questions: 1) do species show niche conservatism in space (France vs Morocco) and time (breeding vs winter period)? 2) Does habitat use change at different interspecific densities? 3) Which factors drive the narrow range limit between G. cristata and G. macrorhyncha? We found that G. theklae was strongly associated with sloping and rocky (rugged) habitats in comparison with crested larks, but habitat use was not entirely consistent in space and time. Evidences for ecological displacement were mitigated. In the only habitat shared by G. cristata and G. theklae in France (vineyards surrounded by low shrubland), we observed a one‐third increase of G. theklae from nearby garrigues in winter, which coincided with a significant decrease of G. cristata after we controlled for density changes in non‐shared habitats. However, in Morocco, the use of rugged habitats by G. theklae was not affected by the prevalence of G. cristata or G. macrorhyncha in surrounding areas, at least at the spatial scales investigated. Our results suggest that interspecific competition plays a role in the parapatric distribution of G. cristata and G. macrorhyncha, because: 1) the alternative of a recent contact receives little support; 2) transition between species coincides with aridity gradients but not with topographic barriers or a sharp ecotone; 3) G. cristata also occurs in arid areas wherein G. macrorhyncha is rare or absent. Overall, our results add support to the view that a better integration of biotic interactions and spatio‐temporal variations in ecological niches might increase the predictive power of ecological niche models.     

Genetic and phenotypic variation among geographically isolated populations of the globally threatened Dupont's lark Chersophilus duponti

Identifying genetically and phenotypically distinct populations of threatened species is critical if we are to delineate appropriate plans for their conservation. We conducted an integrated analysis of population genetic structure, historical demographic events, current gene flow (all based on mtDNA sequences) and morphological variation of three geographically separated groups of populations of Dupont's lark Chersophilus duponti, located in the Iberian Peninsula (three populations), Morocco (two populations), and Tunisia (one population). Unusually, this lark species is the only one among the genus Chersophilus. Our results revealed the early historical divergence of an eastern Dupont's lark lineage (in Tunisia) and a western lineage (in Morocco and Spain), consistent with subspecies taxonomy and distribution. The western lineage subsequently split into two lineages, following the isolation of Iberian and African populations. Such pattern of historical differentiation caused great population genetic structure, with differences among geographic areas explaining more than 80% of total genetic variation. Mismatch distributions and coalescent estimates of divergence time showed that lineage divergence was associated with sudden population expansion events, which apparently took place during the last glaciation, when steppe habitats were widespread across the Mediterranean region. Extant populations from different geographic areas hardly shared any haplotype (only one out of 16 ND2 haplotypes was shared by Tunisian and Moroccan Dupont's larks), and consequently gene flow between geographic areas was found to be virtually absent. Apart from showing great genetic differentiation, Dupont's larks from different geographic areas were morphologically distinct, showing substantial variation in body size and feeding-related traits (length of feet and bill). We conclude that Dupont's lark populations isolated in the Iberian Peninsula, Morocco, and Tunisia are distinct evolutionary entities and should be considered as such in conservation plans. Such circumstance sets a daunting conservation challenge that exemplifies the need of incorporating knowledge of historical processes to our general understanding of the demography of threatened species.     

Ontogenetic and gender-related variation in the isotopic niche within and between three species of fur seals (genus <Emphasis Type="Italic">Arctocephalus</Emphasis>)

Differences in aggressiveness when competing for environmental resources are the main factor leading to social hierarchy in group living fish. Social status acquired is related to changes in physiological parameters, as metabolic rate. Habitat variation can interfere with aggressive behaviour and promote changes in physiological parameters associated with social status. The primary goal of our study was to investigate how differences in habitat complexity affect the relationship between resting metabolic rate (RMR) and social status in the Amazonian dwarf cichlid Apistogramma agassizii. We compared agonistic interactions between pairs of males in aquaria with different habitat enrichment levels, manipulated by adding shelters. RMR was measured before and after hierarchy establishment. Habitat enrichment promotes changes in aggressive behaviour and influences differences in metabolic rate between dominant and subordinate fish. We observed an increase in biting by dominant fish at high enrichment habitat, which could be related to the increase in territory value. We observed an increase in metabolic rate in dominant fish after hierarchy establishment. However, it occurs only in enriched habitats. We concluded that habitat structure interfere with behavioural characteristics in social hierarchies, as aggressiveness, and changes in aggressive interactions affect metabolic rate in different social ranks in the dwarf cichlid Apistogramma agassizii.     

Influence of learning on range expansion and adaptation to novel habitats

Learning has been postulated to ‘drive’ evolution, but its influence on adaptive evolution in heterogeneous environments has not been formally examined. We used a spatially explicit individual‐based model to study the effect of learning on the expansion and adaptation of a species to a novel habitat. Fitness was mediated by a behavioural trait (resource preference), which in turn was determined by both the genotype and learning. Our findings indicate that learning substantially increases the range of parameters under which the species expands and adapts to the novel habitat, particularly if the two habitats are separated by a sharp ecotone (rather than a gradient). However, for a broad range of parameters, learning reduces the degree of genetically‐based local adaptation following the expansion and facilitates maintenance of genetic variation within local populations. Thus, in heterogeneous environments learning may facilitate evolutionary range expansions and maintenance of the potential of local populations to respond to subsequent environmental changes.     

繁殖期两种百灵科鸟类对捕食风险的行为响应

生活史理论认为亲代对捕食风险的不同响应影响亲代资源在繁殖中的分配比例,为检验这一理论,于2002—2004年的4—9月以青藏高原高寒草甸的角百灵(Eremophilaalpestris)和小云雀(Alaudagulgula)为研究对象,通过捕食风险处理实验,研究了两种鸟对不同强度捕食压的响应。结果表明在自然条件下,二者繁殖行为存在显著差异,而在捕食压力下二者的行为反应没有差异。在捕食风险下,二者亲鸟缩短伴巢时间,减少递食率,延长消失时间;二者雄性的风险容忍度均低于雌性。此外,角百灵亲代对人类活动的响应较小云雀敏感。以上结果表明,这两种百灵科鸟类通过降低当前的繁殖投入和提高未来存活概率来响应捕食风险,支持上述生活史理论的观点。     

Absence of egg discrimination in a suitable cuckoo Cuculus canorus host breeding away from trees

There is at present considerable variation in the level of antiparasite defences among different host species of avian brood parasites, but in many potential hosts some individuals reject poorly matching parasite eggs. Here we present unique absence of egg discrimination behaviour backed up by a lack of egg recognition abilities in a suitable common cuckoo Cuculus canorus host, the skylark Alauda arvensis. Skylarks did not show any clear rejection response to experimentally added highly non‐mimetic foreign eggs in any behavioural context, even before they had started laying or when the whole clutch was exchanged with foreign eggs. This absence of antiparasite defence can be explained by the breeding habitat of larks consisting of largely treeless open landscapes where cuckoos have little access to the nests, thereby eroding the possibility of coevolutionary interactions. Our results are strikingly consistent with the spatial habitat structure hypothesis proposed to explain the occurrence and extent of avian host‐parasite co‐adaptation.     

A Fossil Species of Eremophila and Other Larks (Aves,Alaudidae) from the upper Pliocene of the Selenga River Valley (Central Asia)

We describe lark (Alaudidae) fossils from the upper Pliocene of the Beregovaya (southern Transbaikalia) and Shaamar (northern Mongolia) localities. The presence of 4 extinct forms in these localities is established, including the new fossil horned lark Eremophila orkhonensis (Zelenkov et Kurochkin, 2012), comb. nov. This is the oldest member of Eremophila in the fossil record, indicating a possible Central Asian origin of the genus. Two other larks Alaudala aff. A. rufescens and Calandrella aff. C. brachydactyla also probably represent extinct forms. The paper describes in detail the osteology of larks and compares it with other passerines in its size class. The evolutionary history of Eremophila is discussed, and the environmental preferences of larks and their relationship to the late Pliocene landscapes of Central Asia are considered.     

The influence of habitat and behavioural interactions on the local distribution of the wrasse,Pseudolabrus celidotus

Synopsis The temperate wrasse, Pseudolabrus celidotus (Labridae), was studied in a shallow rocky reef habitat to asses how ecological and behavioural factors influence its within habitat patterns of distribution. Emphasis was placed on examining the mechanisms underlying size/age related changes in dispersion and habitat use. Juveniles (< 100 mm SL) were found to be aggregated whereas adults tended to be more randomly dispersed. This pattern appeared to be a product of both (a) ecological, and (b) behavioural factors. (a) Juveniles were closely associated with shelter (macro-algae) which was located in patches throughout the habitat. Their local distribution was probably not related to the distribution of their major prey items. Shelter dependence appears to decrease with growth and fish switched from feeding on micro-crustaceans present in macro-algae to those associated with bare rock — coralline turf areas. Adults foraged throughout larger home ranges, but fed preferentially in microhabitats where their prey items (e.g. bivalves) were most abundant. (b) Juveniles tended to aggregate into loose foraging schools. This tendency declined with age and was associated with a corresponding increase in the rate of aggressive interactions among individuals of a cohort. Consequently, individuals of the same size became more spaced out as they grew. Between cohort association or aggression was rare and apparently unimportant in determining local patterns of distribution. Many interspecific foraging associations and aggressive interactions were recorded. However, with the probable exception of interspecific territoriality by the blenny Forsterygion varium, these behaviours probably had only a transitory influence on the distribution of the wrasse.     

A comprehensive framework for global patterns in biodiversity

The present study proposes to reconcile the different spatial and temporal scales of regional species production and local constraint on species richness. Although interactions between populations rapidly achieve equilibrium and limit membership in ecological communities locally, these interactions occur over heterogeneous environments within large regions, where the populations of species are stably regulated through competition and habitat selection. Consequently, exclusion of species from a region depends on long‐term regional‐scale environmental change or evolutionary change among interacting populations, bringing species production and extinction onto the same scale and establishing a link between local and regional processes.     

Competition between birds and mammals: A comparison of giving-up densities between crested larks and gerbils

We combined the concept of mechanisms of co-existence with the approach of giving-up densities to study inter-taxon competition between seed-eating birds and mammals. We measured feeding behaviour in food patches to define and study the guild of seed-eating vertebrates occupying sandy habitats at Bir Asluj, Negev Desert, Israel. Despite a large number of putatively granivorous rodents and birds at the site, two gerbil species (Allenbys gerbil, Gerbillus allenbyi, and the greater Egyptian gerbil, G. pyramidum) dominated nocturnal foraging, and a single bird species (crested lark, Galerida cristata) contributed all of the daytime foraging. We used giving-up densities to quantify foraging behaviour and foraging efficiencies. A low giving-up density demonstrates the ability of a forager to profitably harvest food at low abundances and to profitably utilize the foraging opportunities left behind by the less efficient forager. Gerbils had lower giving-up densities in the bush than open microhabitat, and lower giving-up densities in the semi-stabilized than stabilized sand habitats. Crested larks showed the opposite: lower giving-up densities in the open than bush, and on the stabilized than semi-stabilized sand habitats. Despite these patterns, gerbils had substantially lower giving-up densities than crested larks in both microhabitats, all sand habitats, and during each month. Several mechanisms may permit the crested lark to co-exist with the gerbils. Larks may be cream skimmers on the high spatial and temporal variability in seed abundances. Larks may rely on insects, fruit or smaller seeds. Or, larks may rely on adjacent rocky habitats.     

Species persistence decreases with habitat fragmentation: an analysis in periodic stochastic environments

This paper presents a study of a nonlinear reaction–diffusion population model in fragmented environments. The model is set on , with periodic heterogeneous coefficients obtained using stochastic processes. Using a criterion of species persistence based on the notion of principal eigenvalue of an elliptic operator, we provided a precise numerical analysis of the interactions between habitat fragmentation and species persistence. The obtained results clearly indicated that species persistence strongly tends to decrease with habitat fragmentation. Moreover, comparing two stochastic models of landscape pattern generation, we observed that in addition to local fragmentation, a more global effect of the position of the habitat patches also influenced species persistence.      

Reproductive interference hampers species coexistence despite conspecific sperm precedence

Negative interspecific mating interactions, known as reproductive interference, can hamper species coexistence in a local patch and promote niche partitioning or geographical segregation of closely related species. Conspecific sperm precedence (CSP), which occurs when females that have mated with both conspecific and heterospecific males preferentially use conspecific sperm for fertilization, might contribute to species coexistence by mitigating the costs of interspecific mating and hybridization. We discussed whether two species exhibiting CSP can coexist in a local environment in the presence of reproductive interference. First, using a behaviorally explicit mathematical model, we demonstrated that two species characterized by negative mating interactions are unlikely to coexist because the costs of reproductive interference, such as loss of mating opportunity with conspecific partners, are inevitably incurred when individuals of both species are present. Second, we experimentally examined differences in mating activity and preference in two Harmonia ladybird species known to exhibit CSP. These behavioral differences may lead to local extinction of H. yedoensis because of reproductive interference by H. axyridis. This prediction is consistent with field observations that H. axyridis uses various food sources and habitats whereas H. yedoensis is confined to a less preferred prey item and a pine tree habitat. Finally, by a comparative approach, we observed that niche partitioning or parapatric distribution, but not sympatric coexistence in the same habitat, is maintained between species with CSP belonging to a wide range of taxa, including vertebrates and invertebrates living in aquatic or terrestrial environments. Taken together, it is possible that reproductive interference may destabilize local coexistence even in closely related species that exhibit CSP.