A DNA test to sex most birds.

Coevolutionary theory predicts that the most common long-term outcome of the relationships between brood parasites and their hosts should be coevolutionary cycles based on a dynamic change selecting the currently least-defended host species, given that when well-defended hosts are abandoned, hosts will be selected to decrease their defences as these are usually assumed to be costly. This is assumed to be the case also in brood parasite-host systems. Here I examine the frequency of the three potential long-term outcomes of brood parasite-host coevolution (coevolutionary cycles, lack of rejection, and successful resistance) in 182 host species. The results of simple exploratory comparisons show that coevolutionary cycles are very scarce while the lack of rejection and successful resistance, which are considered evolutionary enigmas, are much more frequent. I discuss these results considering (i) the importance of different host defences at all stages of the breeding cycle, (ii) the role of phenotypic plasticity in long-term coevolution, and (iii) the evolutionary history of host selection. I suggest that in purely antagonistic coevolutionary interactions, such as those involving brood parasites and their hosts, that although cycles will exist during an intermediate phase of the interactions, the arms race will end with the extinction of the host or with the host acquiring successful resistance. As evolutionary time passes, this resistance will force brood parasites to use previously less suitable host species. Furthermore, I present a model that represents the long-term trajectories and outcomes of coevolutionary interactions between brood parasites and their hosts with respect to the evolution of egg-rejection defence. This model suggests that as an increasing number of species acquire successful resistance, other unparasitized host species become more profitable and their parasitism rate and the costs imposed by brood parasitism at the population level will increase, selecting for the evolution of host defences. This means that although acceptance is adaptive when the parasitism rate and the costs of parasitism are very low, this cannot be considered to represent an evolutionary equilibrium, as conventional theory has done to date, because it is not stable.

Long-term coevolution between avian brood parasites and their hosts.

Avian parents and social insect colonies are victimized by interspecific brood parasites-cheats that procure costly care for their dependent offspring by leaving them in another species' nursery. Birds and insects defend themselves from attack by brood parasites; their defences in turn select counter-strategies in the parasite, thus setting in motion antagonistic co-evolution between the two parties. Despite their considerable taxonomic disparity, here we show striking parallels in the way that co-evolution between brood parasites and their hosts proceeds in insects and birds. First, we identify five types of co-evolutionary arms race from the empirical literature, which are common to both systems. These are: (a) directional co-evolution of weaponry and armoury; (b) furtiveness in the parasite countered by strategies in the host to expose the parasite; (c) specialist parasites mimicking hosts who escape by diversifying their genetic signatures; (d) generalist parasites mimicking hosts who escape by favouring signatures that force specialization in the parasite; and (e) parasites using crypsis to evade recognition by hosts who then simplify their signatures to make the parasite more detectable. Arms races a and c are well characterized in the theoretical literature on co-evolution, but the other types have received little or no formal theoretical attention. Empirical work suggests that hosts are doomed to lose arms races b and e to the parasite, in the sense that parasites typically evade host defences and successfully parasitize the nest. Nevertheless hosts may win when the co-evolutionary trajectory follows arms race a, c or d. Next, we show that there are four common outcomes of the co-evolutionary arms race for hosts. These are: (1) successful resistance; (2) the evolution of defence portfolios (or multiple lines of resistance); (3) acceptance of the parasite; and (4) tolerance of the parasite. The particular outcome is not determined by the type of preceding arms race but depends more on whether hosts or parasites control the co-evolutionary trajectory: tolerance is an outcome that parasites inflict on hosts, whereas the other three outcomes are more dependent on properties intrinsic to the host species. Finally, our review highlights considerable interspecific variation in the complexity and depth of host defence portfolios. Whether this variation is adaptive or merely reflects evolutionary lag is unclear. We propose an adaptive explanation, which centres on the relative strength of two opposing processes: strategy-facilitation, in which one line of host defence promotes the evolution of another form of resistance, and strategy-blocking, in which one line of defence may relax selection on another so completely that it causes it to decay. We suggest that when strategy-facilitation outweighs strategy-blocking, hosts will possess complex defence portfolios and we identify selective conditions in which this is likely to be the case.

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Cuckoos versus hosts in insects and birds: adaptations, counter‐adaptations and outcomes

中国鸟类图鉴 = Field guide to the birds of China

Although deforestation and forest degradation have long been considered the most significant threats to tropical biodiversity, across Southeast Asia (Northeast India, Indochina, Sundaland, Philippines) substantial areas of natural habitat have few wild animals (>1 kg), bar a few hunting-tolerant species. To document hunting impacts on vertebrate populations regionally, we conducted an extensive literature review, including papers in local journals and reports of governmental and nongovernmental agencies. Evidence from multiple sites indicated animal populations declined precipitously across the region since approximately 1980, and many species are now extirpated from substantial portions of their former ranges. Hunting is by far the greatest immediate threat to the survival of most of the region's endangered vertebrates. Causes of recent overhunting include improved access to forests and markets, improved hunting technology, and escalating demand for wild meat, wildlife-derived medicinal products, and wild animals as pets. Although hunters often take common species, such as pigs or rats, for their own consumption, they take rarer species opportunistically and sell surplus meat and commercially valuable products. There is also widespread targeted hunting of high-value species. Consequently, as currently practiced, hunting cannot be considered sustainable anywhere in the region, and in most places enforcement of protected-area and protected-species legislation is weak. The international community's focus on cross-border trade fails to address overexploitation of wildlife because hunting and the sale of wild meat is largely a local issue and most of the harvest is consumed in villages, rural towns, and nearby cities. In addition to improved enforcement, efforts to engage hunters and manage wildlife populations through sustainable hunting practices are urgently needed. Unless there is a step change in efforts to reduce wildlife exploitation to sustainable levels, the region will likely lose most of its iconic species, and many others besides, within the next few years.

Impacts of hunting on tropical forests in Southeast Asia

© 2010 Yang et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

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Coevolution in Action: Disruptive Selection on Egg Colour in an Avian Brood Parasite and Its Host

In this expose we provide the first review of host use by brood parasitic cuckoos in a multiple-cuckoo system in China, based on our own long-term field data and a compilation of observations obtained from the literature. In total, we found that 11 species of cuckoos utilized altogether 55 host species. These hosts belong to 15 families, in which Sylviidae, Turdidae and Timaliidae account for 22.6%, 20.8% and 17.0% of parasitism records, respectively. The Common Cuckoo (Cuculus canorus) had the widest range of host species, accounting for 45.5% of the total number of parasitized species (25 in 10 families) of all parasitism records and is the most frequent brood parasite in the country. Cuckoo species differed in their egg coloration and the extent of egg polymorphism with most of them, e.g. the Common Cuckoo, the Lesser Cuckoo (C. poliocephalus) and the Plaintive Cuckoo (Cacomantis merulinus) laying well mimetic eggs with respect to their hosts based on human being’s visual observations, while others such as the Large Hawk-cuckoo (C. sparverioides), the Himalayan Cuckoo (C. saturatus) and the Asian Emerald Cuckoo (Chrysococcyx maculatus) usually laid non-mimetic eggs. The use of cuckoo hosts and egg color variation in China are compared with those in other parts of their ranges in Asia.

Diversity of parasitic cuckoos and their hosts in China

In China, many bird species are generally thought to be threatened mainly, or at least partly, by hunting. However, there have been few studies of bird hunting at a local scale. Bird hunting and trade in Nanmao, a remote mountainous village of Hainan Island, China, was investigated during March–July 2003 and September–October 2005. In total, 86 households were visited, of which 43% reported that they engaged in hunting of birds while 91% of households were seen to have hunted birds or hunting tools. This indicated that hunting by village people was widespread. Most hunters were male, and were between 12 and 68 years old. A total of 78 bird species were hunted, including 2 First Class and 19 Second Class national protected species. This extreme level of hunting has changed from a more moderate subsistence hunting tradition since about 1980, when local urban markets for wild meat started to develop. We outline a strategic plan designed to conserve birds, other wildlife and their forest habitats, whilst improving the livelihoods and preserving the minority tribal traditions of the people of Nanmao forest.

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Extreme levels of hunting of birds in a remote village of Hainan Island, China.

How do potential hosts escape detrimental interactions with brood parasites? Current consensus is that hole-nesting and granivorous birds avoid brood parasites, like common cuckoos Cuculus canorus, by their inaccessible nest-sites and food unsuitable for parasites, respectively. Any open-nesting insectivorous hosts are believed to remain open to brood parasite exploitation which leads to the evolution of costly host defences like egg or chick discrimination. In contrast to this coevolutionary scenario, we show for the first time that a previously not studied but seemingly suitable host species escapes brood parasites. The Asian verditer flycatcher Eumyias thalassinus, feed newly hatched chicks entirely with beetles and grasshoppers. These are poor quality and hard to digest diet items that are rarely fed to own or cuckoo chicks by regular hosts. Indeed, chick cross-fostering experiments showed that these food items remained undigested by either cuckoos or other sympatric passerines causing them to die quickly. Egg discrimination experiments showed that the flycatcher accepts any foreign eggs. Although most but not all other potential explanations can be safely excluded at present, the most parsimonious historical explanation for these patterns is that the flycatcher exploits a trophic niche that no other sympatric bird can exploit, and that any cuckoo lineages that switch from their original hosts to the flycatcher have no possibilities for establishing viable populations. Thus, the current classification of host suitability based on diet composition may need revision, raising an important cautionary tale for comparative studies and the interpretation of apparent host rejection of parasitic chicks.

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Host selection in parasitic birds: are open-cup nesting insectivorous passerines always suitable cuckoo hosts?

Department of Biology, Norwegian University of Scienceand Technology (NTNU), NO-7491 Trondheim, NorwayWe documented brood parasitism by the poorly studiedLarge Hawk-Cuckoo on a previously unknown host spe-cies, the Chinese Babax. Furthermore, we describe anew egg colour for the Large Hawk-Cuckoo. The parasit-ism rate of Chinese Babax nests over 4 years was 6.9%(11 of 159 nests), with signiﬁcant temporal variation.The Large Hawk-Cuckoo laid immaculate white eggsthat appeared non-mimetic to the blue Babax eggs, animpression that was conﬁrmed by avian visual modelling.Nevertheless, most Cuckoo eggs were accepted by thehost, suggesting that this host–parasite system may beevolutionarily recent.

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Yan Cai

Papers

Coevolution in Action: Disruptive Selection on Egg Colour in an Avian Brood Parasite and Its Host

Diversity of parasitic cuckoos and their hosts in China

Extreme levels of hunting of birds in a remote village of Hainan Island, China.

Host selection in parasitic birds: are open-cup nesting insectivorous passerines always suitable cuckoo hosts?

Large Hawk‐Cuckoo Hierococcyx sparverioides parasitism on the Chinese Babax Babax lanceolatus may be an evolutionarily recent host–parasite system