Contribution of cultural heritage values to steppe conservation on ancient burial mounds of Eurasia.

Civilizations, including ancient ones, have shaped global ecosystems in many ways through coevolution of landscapes and humans. However, the cultural legacies of ancient and lost civilizations are rarely considered in the conservation of the Eurasian steppe biome. We used a data set containing more than 1000 records on localities, land cover, protection status, and cultural values related to ancient steppic burial mounds (kurgans); we evaluated how these iconic and widespread landmarks can contribute to grassland conservation in the Eurasian steppes, which is one of the most endangered biomes on Earth. Using Bayesian logistic generalized regressions and proportional odds logistic regressions, we examined the potential of mounds to preserve grasslands in landscapes with different levels of land-use transformation. We also compared the conservation potential of mounds inside and outside protected areas and assessed whether local cultural values support the maintenance of grasslands on them. Kurgans were of great importance in preserving grasslands in transformed landscapes outside protected areas, where they sometimes acted as habitat islands that contributed to habitat conservation and improved habitat connectivity. In addition to steep slopes hindering ploughing, when mounds had cultural value for local communities, the probability of grassland occurrence on kurgans almost doubled. Because the estimated number of steppic mounds is about 600,000 and similar historical features exist on all continents, our results may be applicable at a global level. Our results also suggested that an integrative socioecological approach in conservation might support the positive synergistic effects of conservation, landscape, and cultural values.

Contribución de los valores culturales para la conservación esteparia en los antiguos montículos funerarios de Eurasia Resumen Las civilizaciones modernas y antiguas han moldeado de muchas maneras los ecosistemas globales mediante la coevolución del paisaje y la humanidad. Sin embargo, pocas veces se considera el legado cultural de las civilizaciones perdidas o antiguas para la conservación del bioma de la estepa euroasiática. Usamos un conjunto de datos que contiene más de 1,000 registros de las localidades, cobertura del suelo, estado de protección y valores culturales relacionados con los antiguos montículos funerarios de esta estepa (kurgans). Después analizamos cómo estos símbolos icónicos y distribuidos extensamente pueden contribuir a la conservación de los pastizales en la estepa euroasiática, uno de los biomas en mayor peligro de extinción. Analizamos el potencial de conservación de los montículos en paisajes con diferentes niveles de transformación en el uso de suelo mediante regresiones logísticas generalizadas bayesianas y regresiones logísticas de probabilidades proporcionales. También comparamos el potencial de conservación de los montículos dentro y fuera de las áreas protegidas y evaluamos si los valores culturales locales conservan los pastizales dentro de estas mismas áreas. Los kurgans fueron de gran importancia para la conservación de los pastizales en los paisajes transformados ubicados fuera de las áreas protegidas, en donde llegaron a fungir como hábitats aislados que contribuyeron a la conservación y conectividad del hábitat. Además de que las pendientes pronunciadas impiden el arado, cuando los montículos contaban con valor cultural para las comunidades locales, la probabilidad de que el pastizal se ubicara sobre un kurgan casi se duplicó. Ya que se estima que el número de montículos esteparios ronda los 6,000 y que rasgos históricos similares existen en todos los continentes, nuestros resultados pueden aplicarse a nivel global. Nuestros resultados también sugieren que una estrategia socio-ecológica integradora para la conservación podría respaldar los efectos sinérgicos positivos de la conservación, el paisaje y los valores culturales.

Female-female aggression and male responses to the two colour morphs of female common cuckoos.

Female-only colour polymorphism is rare in birds, but occurs in brood parasitic cuckoos (Cuculidae). Obligate brood parasites leave incubation and parental care to other species (hosts), so female-female interactions can play a role in how parasites guard critical resources (host nests) within their laying areas. The plumage of adult female common cuckoos (Cuculus canorus) is either rufous (typically rare) or grey (common), whereas adult male conspecifics are monochromatic (grey). In previous studies, hosts and conspecific males responded with less intensity toward the rare female morph in support of a negative frequency-dependent benefit of female plumage polychromatism. Here, we assessed responses of both conspecific females and males to vocal playbacks of female calls, coupled with one of two 3D models of the different morphs of female cuckoos. At our study population, the rufous female morph was as common as the grey morph; therefore, we predicted similarly high rates of conspecific responses in both treatments. Both female and male cuckoos responded to playbacks acoustically, which demonstrated the primary role of acoustic communication in social interactions amongst cuckoos. Following this, some cuckoos flew closer to the models to inspect them visually. As predicted, no significant differences were detected between the live cuckoos' responses toward the two colour morphs in this population. We conclude that dichromatism in female cuckoos evolved to serve one or more functions other than conspecific signalling.

Naïve hosts of avian brood parasites accept foreign eggs, whereas older hosts fine-tune foreign egg discrimination during laying.

BACKGROUND: Many potential hosts of social parasites recognize and reject foreign intruders, and reduce or altogether escape the negative impacts of parasitism. The ontogenetic basis of whether and how avian hosts recognize their own and the brood parasitic eggs remains unclear. By repeatedly parasitizing the same hosts with a consistent parasitic egg type, and contrasting the responses of naïve and older breeders, we studied ontogenetic plasticity in the rejection of foreign eggs by the great reed warbler (Acrocephalus arundinaceus), a host species of the common cuckoo (Cuculus canorus). RESULTS: In response to experimental parasitism before the onset of laying, first time breeding hosts showed almost no egg ejection, compared to higher rates of ejection in older breeders. Young birds continued to accept foreign eggs when they were subjected to repeated parasitism, whereas older birds showed even higher ejection rates later in the same laying cycle. CONCLUSIONS: Our results are consistent with the hypotheses that (i) naïve hosts need to see and learn the appearance of their own eggs to discriminate and reject foreign eggs, whereas (ii) experienced breeders possess a recognition template of their own eggs and reject parasitic eggs even without having to see their own eggs. However, we cannot exclude the possibility that other external cues and internal processes, accumulated simply with increasing age, may also modify age-specific patterns in egg rejection (e.g. more sightings of the cuckoo by older breeders). Future research should specifically track the potential role of learning in responses of individual hosts between first and subsequent breeding attempts by testing whether imprinting on a parasitized clutch reduces the rates of rejecting foreign eggs in subsequent parasitized clutches.

Mowing wet meadows reduces the health of their snail communities.

Wet meadows harbor rich biodiversity, making them pivotal ecosystems worldwide. These habitats are commonly used for grazing or hay production for livestock. However, regular mowing can influence these habitats, potentially leading to significant repercussions for the animals residing within them. In order to investigate the effects of land management practices, we conducted an experimental study to compare snail communities in mowed and unmowed wet meadows in northern Hungary. We found that overall, mowing decreases snail populations, as well as species richness and diversity. Thus, our results suggest that routine mowing of wet meadows is deleterious to their snail communities. Based on these results, we suggest that designated patches of meadows that are regularly managed should be left uncultivated in their natural state. These patches can serve as potential colonization sites, facilitating the restoration of the entire meadow's ecological balance.

Evolution and genetic architecture of sex-limited polymorphism in cuckoos.

Sex-limited polymorphism has evolved in many species including our own. Yet, we lack a detailed understanding of the underlying genetic variation and evolutionary processes at work. The brood parasitic common cuckoo (Cuculus canorus) is a prime example of female-limited color polymorphism, where adult males are monochromatic gray and females exhibit either gray or rufous plumage. This polymorphism has been hypothesized to be governed by negative frequency-dependent selection whereby the rarer female morph is protected against harassment by males or from mobbing by parasitized host species. Here, we show that female plumage dichromatism maps to the female-restricted genome. We further demonstrate that, consistent with balancing selection, ancestry of the rufous phenotype is shared with the likewise female dichromatic sister species, the oriental cuckoo (Cuculus optatus). This study shows that sex-specific polymorphism in trait variation can be resolved by genetic variation residing on a sex-limited chromosome and be maintained across species boundaries.

Discordancy or template-based recognition? Dissecting the cognitive basis of the rejection of foreign eggs in hosts of avian brood parasites.

Many avian hosts have evolved antiparasite defence mechanisms, including egg rejection, to reduce the costs of brood parasitism. The two main alternative cognitive mechanisms of egg discrimination are thought to be based on the perceived discordancy of eggs in a clutch or the use of recognition templates by hosts. Our experiments reveal that the great reed warbler (Acrocephalus arundinaceus), a host of the common cuckoo (Cuculus canorus), relies on both mechanisms. In support of the discordancy mechanism, hosts rejected their own eggs (13%) and manipulated ('parasitic') eggs (27%) above control levels in experiments when manipulated eggs were in the majority but when clutches also included a minority of own eggs. Hosts that had the chance to observe the manipulated eggs daily just after laying did not show stronger rejection of manipulated eggs than when the eggs were manipulated at clutch completion. When clutches contained only manipulated eggs, in 33% of the nests hosts showed rejection, also supporting a mechanism of template-based egg discrimination. Rejection using a recognition template might be more advantageous because discordancy-based egg discrimination is increasingly error prone with higher rates of multiple parasitism.

Host alarm calls attract the unwanted attention of the brood parasitic common cuckoo.

It is well known that avian brood parasites lay their eggs in the nests of other bird species, called hosts. It remains less clear, however, just how parasites are able to recognize their hosts and identify the exact location of the appropriate nests to lay their eggs in. While previous studies attributed high importance to visual signals in finding the hosts' nests (e.g. nest building activity or the distance and direct sight of the nest from vantage points used by the brood parasites), the role of host acoustic signals during the nest searching stage has been largely neglected. We present experimental evidence that both female and male common cuckoos Cuculus canorus pay attention to their host's, the great reed warbler's Acrocephalus arundinaceus alarm calls, relative to the calls of an unparasitized species used as controls. Parallel to this, we found no difference between the visibility of parasitized and unparasitized nests during drone flights, but great reed warblers that alarmed more frequently experienced higher rates of parasitism. We conclude that alarm calls might be advantageous for the hosts when used against enemies or for alerting conspecifics, but can act in a detrimental manner by providing important nest location cues for eavesdropping brood parasites. Our results suggest that host alarm calls may constitute a suitable trait on which cuckoo nestlings can imprint on to recognize their primary host species later in life. Our study contributes to the growing body of knowledge regarding the context-dependency of animal signals, by providing a novel example of a beneficial acoustic trait intercepted by a heterospecific and used against the emitter.

Multiple parasitism reduces egg rejection in the host (Acrocephalus arundinaceus) of a mimetic avian brood parasite (Cuculus canorus).

A host that has been targeted by an avian brood parasite can recover most of its potential fitness loss by ejecting the foreign egg(s) from its nest. The propensity for some hosts to engage in egg rejection behavior has put selective pressure on their parasites to evolve mimetic eggshells resembling the host's own shell colors and maculation. In turn, hosts have counterevolved increasingly more sophisticated detection methods such as narrowing visual egg acceptance thresholds or using social cues to recognize parasitism. However, multiple cognitive mechanisms acting simultaneously could theoretically interfere with one another and ultimately decrease egg rejection accuracy, especially if these heuristics yield differing targets for rejection. By painting hosts own eggs, we studied a host species of the common cuckoo Cuculus canorus, the great reed warbler Acrocephalus arundinaceus, and tested its responses to the presence of "foreign" eggs of varying quantity, colors, and uniformity. Using reflectance spectra of egg background coloration and avian perceptual modeling, we then estimated the sensory thresholds triggering egg rejection by this host for each treatment. As previously reported, rejection rates were positively related to the perceptual distance between own and foreign eggs in the nests in all treatments. However, rejection thresholds were more permissive (error prone) both with greater proportions of foreign eggs per clutch and/or when the suite of foreign eggs was perceptually more variable within the nest. These results suggest that parasites, through multiple parasitism, can partially overcome the evolution of hosts' recognition of mimetic parasite eggs. (PsycINFO Database Record (c) 2019 APA, all rights reserved).

Are both notes of the common cuckoo's call necessary for familiarity recognition?

Common cuckoos (Cuculus canorus) are best known for their simple two-note calls ("cu-coo"), which are uttered only by males during the breeding season. A previous playback study revealed that territorial males were more tolerant toward playbacks of the calls of familiar, neighbouring individuals than toward unfamiliar, stranger simulated intruders, exhibiting the classical "dear-enemy" phenomenon. Here we experimentally assessed whether the acoustic cues for familiarity recognition are encoded in the first and/or second note of these simple calls. To do so, we played mixed sound files to radio-tagged cuckoos, where the first part of the two-note calls was taken from strangers and the second part from neighbours, or vice versa. As controls, we used behavioural data from two-note neighbour and two-note stranger call playbacks. Cuckoos responded consistently to the two types of mixed sound files. When either the first or second note of the call was taken from a stranger and the other note from a neighbour, they responded to these sound files similarly to two-note playbacks of strangers: they approached the speaker of the playbacks more closely and the calling response-latency to playbacks was longer than to familiar controls. These findings point to the importance of both notes in familiarity recognition. We conclude that familiarity recognition in male common cuckoos needs the complete structure of the two-note cuckoo call, which is characteristic for this species.

Competition with a host nestling for parental provisioning imposes recoverable costs on parasitic cuckoo chick's growth.

Chicks of the brood parasitic common cuckoo (Cuculus canorus) typically monopolize host parental care by evicting all eggs and nestmates from the nest. To assess the benefits of parasitic eviction behaviour throughout the full nestling period, we generated mixed broods of one cuckoo and one great reed warbler (Acrocephalus arundinaceus) to study how hosts divide care between own and parasitic young. We also recorded parental provisioning behaviour at nests of singleton host nestlings or singleton cuckoo chicks. Host parents fed the three types of broods with similar-sized food items. The mass of the cuckoo chicks was significantly reduced in mixed broods relative to singleton cuckoos. Yet, after the host chick fledged from mixed broods, at about 10-12 days, cuckoo chicks in mixed broods grew faster and appeared to have compensated for the growth costs of prior cohabitation by fledging at similar weights and ages compared to singleton cuckoo chicks. These results are contrary to suggestions that chick competition in mixed broods of cuckoos and hosts causes an irrecoverable cost for the developing brood parasite. Flexibility in cuckoos' growth dynamics may provide a general benefit to ecological uncertainty regarding the realized successes, failures, and costs of nestmate eviction strategies of brood parasites.

Egg eviction imposes a recoverable cost of virulence in chicks of a brood parasite.

BACKGROUND: Chicks of virulent brood parasitic birds eliminate their nestmates and avoid costly competition for foster parental care. Yet, efforts to evict nest contents by the blind and naked common cuckoo Cuculus canorus hatchling are counterintuitive as both adult parasites and large older cuckoo chicks appear to be better suited to tossing the eggs and young of the foster parents. METHODOLOGY/PRINCIPAL FINDINGS: Here we show experimentally that egg tossing imposed a recoverable growth cost of mass gain in common cuckoo chicks during the nestling period in nests of great reed warbler Acrocephalus arundinaceus hosts. Growth rates of skeletal traits and morphological variables involved in the solicitation of foster parental care remained similar between evictor and non-evictor chicks throughout development. We also detected no increase in predation rates for evicting nests, suggesting that egg tossing behavior by common cuckoo hatchlings does not increase the conspicuousness of nests. CONCLUSION: The temporary growth cost of egg eviction by common cuckoo hatchlings is the result of constraints imposed by rejecter host adults and competitive nestmates on the timing and mechanism of parasite virulence.

Experimental shift in hosts' acceptance threshold of inaccurate-mimic brood parasite eggs.

Hosts are expected to evolve resistance strategies that efficiently detect and resist exposure to virulent parasites and pathogens. When recognition is not error-proof, the acceptance threshold used by hosts to recognize parasites should be context dependent and become more restrictive with increasing predictability of parasitism. Here, we demonstrate that decisions of great reed warblers Acrocephalus arundinaceus to reject parasitism by the common cuckoo Cuculus canorus vary adaptively within a single egg-laying bout. Hosts typically accept one of their own eggs with experimentally added spots and the background colour left visible. In contrast, hosts reject such spotted eggs when individuals had been previously exposed to and rejected one of their own eggs whose background colour had been entirely masked. These results support patterns of adaptive modulation of antiparasitic strategies through shifts in the acceptance threshold of hosts and suggest a critical role for experience in the discrimination decisions between inaccurate-mimic parasite eggs and hosts' own eggs.