[Show abstract][Hide abstract] ABSTRACT: Traits used in communication, such as colour signals, are expected to have positive consequences for reproductive success, but their associations with survival are little understood. Previous studies have mainly investigated linear relationships between signals and survival, but both hump-shaped and U-shaped relationships can also be predicted, depending on the main costs involved in trait expression. Furthermore, few studies have taken the plasticity of signals into account in viability selection analyses. The relationship between signal expression and survival is of particular interest in melanin-based traits, because their main costs are still debated. Here, we first determined the main factors explaining variability in a melanin-based trait linked to dominance: the bib size of a colonial bird, the sociable weaver Philetairus socius. We then used these analyses to obtain a measure representative of the individual mean expression of bib size. Finally, we used capture-recapture models to study how survival varied in relation to bib size. Variation in bib size was strongly affected by year and moderately affected by age, body condition and colony size. In addition, individuals bearing small and large bibs had higher survival than those with intermediate bibs, and this U-shaped relationship between survival and bib size appeared to be more pronounced in some years than others. These results constitute a rare example of disruptive viability selection, and point towards the potential importance of social costs incurred by the dominance signalling function of badges of status. This article is protected by copyright. All rights reserved.
This article is protected by copyright. All rights reserved.
Journal of Evolutionary Biology 08/2015; DOI:10.1111/jeb.12717 · 3.48 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: Dispersal is a critical driver of gene flow, with important consequences for population genetic structure, social interactions and other biological processes. Limited dispersal may result in kin-structured populations in which kin selection may operate, but it may also increase the risk of kin competition and inbreeding. Here, we use a combination of long-term field data and molecular genetics to examine dispersal patterns and their consequences for the population genetics of a highly social bird, the sociable weaver (Philetairus socius), which exhibits cooperation at various levels of sociality from nuclear family groups to its unique communal nests. Using 20 years of data, involving capture of 6508 birds and 3151 recaptures at 48 colonies, we found that both sexes exhibit philopatry and that any dispersal occurs over relatively short distances. Dispersal is female-biased, with females dispersing earlier, further, and to less closely related destination colonies than males. Genotyping data from 30 colonies showed that this pattern of dispersal is reflected by fine-scale genetic structure for both sexes, revealed by isolation by distance in terms of genetic relatedness and significant genetic variance among colonies. Both relationships were stronger among males than females. Crucially, significant relatedness extended beyond the level of the colony for both sexes. Such fine-scale population genetic structure may have played an important role in the evolution of cooperative behaviour in this species, but it may also result in a significant inbreeding risk, against which female-biased dispersal alone is unlikely to be an effective strategy. This article is protected by copyright. All rights reserved.
This article is protected by copyright. All rights reserved.
[Show abstract][Hide abstract] ABSTRACT: Hosts of brood-parasitic birds must distinguish their own eggs from parasitic mimics, or pay the cost of mistakenly raising a foreign chick. Egg discrimination is easier when different host females of the same species each lay visually distinctive eggs (egg 'signatures'), which helps to foil mimicry by parasites. Here, we ask whether brood parasitism is associated with lower levels of correlation between different egg traits in hosts, making individual host signatures more distinctive and informative. We used entropy as an index of the potential information content encoded by nine aspects of colour, pattern and luminance of eggs of different species in two African bird families (Cisticolidae parasitized by cuckoo finches Anomalospiza imberbis, and Ploceidae by diederik cuckoos Chrysococcyx caprius). Parasitized species showed consistently higher entropy in egg traits than did related, unparasitized species. Decomposing entropy into two variation components revealed that this was mainly driven by parasitized species having lower levels of correlation between different egg traits, rather than higher overall levels of variation in each individual egg trait. This suggests that irrespective of the constraints that might operate on individual egg traits, hosts can further improve their defensive 'signatures' by arranging suites of egg traits into unpredictable combinations.
Proceedings of the Royal Society B: Biological Sciences 07/2015; 282(1810). DOI:10.1098/rspb.2015.0598 · 5.29 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: Mimicry of a harmless model (aggressive mimicry) is used by egg, chick and fledgling brood parasites that resemble the host's own eggs, chicks and fledglings. However, aggressive mimicry may also evolve in adult brood parasites, to avoid attack from hosts and/or manipulate their perception of parasitism risk. We tested the hypothesis that female cuckoo finches (Anomalospiza imberbis) are aggressive mimics of female Euplectes weavers, such as the harmless, abundant and sympatric southern red bishop (Euplectes orix). We show that female cuckoo finch plumage colour and pattern more closely resembled those of Euplectes weavers (putative models) than Vidua finches (closest relatives); that their tawny-flanked prinia (Prinia subflava) hosts were equally aggressive towards female cuckoo finches and southern red bishops, and more aggressive to both than to their male counterparts; and that prinias were equally likely to reject an egg after seeing a female cuckoo finch or bishop, and more likely to do so than after seeing a male bishop near their nest. This is, to our knowledge, the first quantitative evidence for aggressive mimicry in an adult bird, and suggests that host-parasite coevolution can select for aggressive mimicry by avian brood parasites, and counter-defences by hosts, at all stages of the reproductive cycle.
Proceedings of the Royal Society B: Biological Sciences 07/2015; 282(1810). DOI:10.1098/rspb.2015.0795 · 5.29 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: The Critically Endangered Archer's Lark (now Liben Lark) Heteromirafra archeri was formerly considered to be endemic to north-western Somalia and known only from the Tog Wajaale Plain, where 18 specimens were collected between 1918 and 1922. Fifteen visits between 1970 and 2008 failed to relocate the species there, although populations are now known from adjacent Ethiopia. We conducted three days of intensive surveys on the Tog Wajaale Plain in May 2010. Despite the three other lark species present being in full display, and H. archeri being recorded to have bred in early June, no Liben Larks were found. Vegetation structure surveys indicated that the plain has a taller and denser growth of grass than either of the other known localities for Liben Lark (the Liben and Jijiga Plains) making Tog Wajaale Plain seem superficially more suitable for the species, which prefers areas of taller grass elsewhere. However, previous large-scale agricultural activities may have altered the composition of grass species and precipitated the observed invasion of exotic weeds, notably Parthenium hysterophorus. Importantly, the Tog Wajaale Plain has a greater density of bushes than either the Liben or Jijiga Plains, possibly making ground-nesting birds more susceptible to predation by perch hunters.
[Show abstract][Hide abstract] ABSTRACT: 1.Cooperatively breeding species are typically long-lived and hence, according to theory, are expected to maximise their lifetime reproductive success through maximising survival. Under these circumstances, the presence of helpers could be used to lighten the effort of current reproduction for parents to achieve higher survival. 2.In addition, individuals of different sexes and ages may follow different strategies, but whether male and female breeders and individuals of different ages benefit differently from the presence of helpers has often been overlooked. Moreover only one study that investigated the relationship between parental survival and the presence of helpers used Capture-Mark-Recaptures analyses (CMR). These methods are important since they allow us to account for the non-detection of individuals that are alive in the population but not detected, and thus the effects on survival and recapture probability to be disentangled. 3.Here we used multi-event CMR methods to investigate whether the number of helpers was associated with an increase in survival probability for male and female breeders of different ages in the sociable weaver Philetairus socius. In this species, both sexes reduce their feeding rate in presence of helpers. We therefore predicted that the presence of helpers should increase the breeders' survival in both sexes, especially early in life when individuals potentially have more future breeding opportunities. In addition, sociable weaver females reduce their investment in eggs in the presence of helpers, so we predicted a stronger effect of helpers on female than male survival. 4.As expected we found that females had a higher survival probability when breeding with more helpers. Unexpectedly, however, male survival probability decreased with increasing number of helpers. This antagonistic effect diminished as the breeders grew older. 5.These results illustrate the complexity of fitness costs and benefits underlying cooperative behaviours and how these may vary with the individuals' sex and age. They also highlight the need for further studies on the sex-specific effects of helpers on survival. This article is protected by copyright. All rights reserved.
This article is protected by copyright. All rights reserved.
Journal of Animal Ecology 04/2015; DOI:10.1111/1365-2656.12377 · 4.73 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: Hosts of brood-parasitic birds typically evolve anti-parasitism defences, including mobbing of parasitic intruders at the nest and the ability to recognise and reject foreign eggs from their clutches. The Greater Honeyguide Indicator indicator is virulent brood parasites that punctures host eggs and kills host young, and accordingly a common host, the Little Bee-eater Merops pusillus, frequently rejects entire clutches that have been parasitised. We predicted that given the high costs of accidentally rejecting an entire clutch, and that the experimental addition of a foreign egg is insufficient to induce this defence, bee-eaters require the sight of an adult parasite near the nest as an additional cue of parasitism before they reject a clutch. We found that many Little Bee-eater parents mobbed Greater Honeyguide dummies while ignoring barbet control dummies, showing that they recognised them as a threat. Surprisingly, however, neither a dummy honeyguide nor the presence of a foreign egg, either separately or in combination, was sufficient to stimulate egg rejection.This article is protected by copyright. All rights reserved.
[Show abstract][Hide abstract] ABSTRACT: The montane inselbergs of northern Mozambique have been comparatively little-studied, yet recent surveys have shown they have a rich biodiversity with numerous endemic species. Here we present the main findings from a series of scientific expeditions to one of these inselbergs, Mt Mabu, and discuss the conservation implications. Comprehensive species lists of plants, birds, mammals and butterflies are presented. The most significant result was the discovery of a c. 7,880 ha block of undisturbed rainforest, most of it at medium altitude (900–1,400 m), a forest type that is not well represented elsewhere. It is possibly the largest continuous block of this forest type in southern Africa. To date, 10 new species (plants, mammals, reptiles and butterflies) have been confirmed from Mt Mabu, even though sampling effort for most taxonomic groups has been low. The species assemblages indicate a relatively long period of isolation and many species found are at the southern limit of their range. Conservationists are now faced with the challenge of how best to protect Mt Mabu and similar mountains in northern Mozambique, and various ways that this could be done are discussed.
[Show abstract][Hide abstract] ABSTRACT: Population trends are determined by gains through reproduction and immigration, and losses through mortality and emigration. These demographic quantities and resulting population dynamics are affected by different external and internal drivers. We examined how these demographic quantities were affected by weather, research-induced disturbance, local density, colony site and year in a metapopulation of 17 sociable weaver (Philetairus socius) colonies over 17 years of study (4 years for reproduction). Most colonies declined, but at different rates. The four demographic quantities were related to different drivers. Survival strongly varied among years and colonies and was positively related to rainfall and negatively related to extreme temperature (together explaining 30 % of variation) and disturbance (measured as number of captures conducted at a colony; 7 %). There was a trend for a positive relationship between reproduction and rainfall (50 %). Movement was mainly related to local density: individuals were more likely to emigrate from small to large colonies and from colonies that were either well below or above their long-term mean. They were more likely to immigrate into colonies that were nearby, and below their mean size. We then quantified the effects of these relationships on metapopulation dynamics using a multi-site matrix projection model. Rainfall was potentially a strong driver of metapopulation dynamics. In addition, field-work disturbance might have contributed to the decline of this metapopulation but could not explain its full magnitude. Hence, through a combination of analytical methods we were able to obtain information on the main drivers affecting dynamics in a declining metapopulation.
[Show abstract][Hide abstract] ABSTRACT: Cuckoo eggs famously mimic those of their foster parents to evade rejection from discriminating hosts. Here we test whether parasites benefit by repeatedly parasitizing the same host nest. This should make accurate rejection decisions harder, regardless of the mechanism that hosts use to identify foreign eggs. Here we find strong support for this prediction in the African tawny-flanked prinia (Prinia subflava), the most common host of the cuckoo finch (Anomalospiza imberbis). We show experimentally that hosts reject eggs that differ from an internal template, but crucially, as the proportion of foreign eggs increases, hosts are less likely to reject them and require greater differences in appearance to do so. Repeated parasitism by the same cuckoo finch female is common in host nests and likely to be an adaptation to increase the probability of host acceptance. Thus, repeated parasitism interacts with egg mimicry to exploit cognitive and sensory limitations in host defences.
[Show abstract][Hide abstract] ABSTRACT: Many brood parasitic birds lay eggs that mimic their hosts' eggs in appearance. This typically arises from selection from discriminating hosts that reject eggs which differ from their own. However, selection on parasitic eggs may also arise from parasites themselves, because it should pay a laying parasitic female to detect and destroy another parasitic egg previously laid in the same host nest by a different female. In this study, I experimentally test the source of selection on greater honeyguide (Indicator indicator) egg size and shape, which is correlated with that of its several host species, all of which breed in dark holes. Its commonest host species did not discriminate against experimental eggs that differed from their own in size and shape, but laying female honeyguides preferentially punctured experimental eggs more than host or control eggs. This should improve offspring survival given that multiple parasitism by this species is common, and that honeyguide chicks kill all other nest occupants. Hence, selection on egg size in greater honeyguides parasitizing bee-eaters appears to be imposed not by host defences but by interference competition among parasites themselves.
[Show abstract][Hide abstract] ABSTRACT: The African lark genus Heteromirafra is thought to consist of three threatened species inhabiting mid-altitude grasslands, one in South Africa and two in the Horn of Africa. One of the latter, Archer’s Lark H. archeri of Somaliland, has not been seen with certainty since 1922. We surveyed its type locality as well as a nearby area of grassland east of Jijiga in adjacent north-eastern Ethiopia, where sightings of Heteromirafra larks have recently been made. First, we used a combination of morphological and molecular evidence to show that these recent sightings refer to the same taxon as Archer’s Lark. Second, we used a combination of morphological, molecular and vocal evidence to show that these populations are conspecific with the Liben (Sidamo) Lark H. sidamoensis of southern Ethiopia, but that the Horn of Africa populations are highly distinct from Rudd’s Lark H. ruddi of South Africa. Third, we suggest that the extent and quality of their habitat in north-eastern Ethiopia is small and poor, and that the type locality of Archer’s Lark in Somaliland has been completely transformed. Taken together, these results imply that there is a single species of Heteromirafra in the Horn of Africa (for which the scientific name H. archeri has priority, and which we suggest retains the English name Liben Lark), consisting of two tiny populations separated by 590 km of apparently unsuitable habitats. Environmental niche models suggest that there are no environmentally similar locations elsewhere within the region. Despite the discovery of a second population, the Liben Lark remains a highly threatened species in urgent need of conservation intervention to avert the extinction of both of its populations.
[Show abstract][Hide abstract] ABSTRACT: Honeyguides (Indicatoridae, Piciformes) are unique among birds in several respects. All subsist primarily on wax, are obligatory brood parasites and one species engages in 'guiding' behavior in which it leads human honey hunters to bees' nests. This unique life history has likely shaped the evolution of their brain size and morphology. Here, we test that hypothesis using comparative data on relative brain and brain region size of honeyguides and their relatives: woodpeckers, barbets and toucans. Honeyguides have significantly smaller relative brain volumes than all other piciform taxa. Volumetric measurements of the brain indicate that honeyguides have a significantly larger cerebellum and hippocampal formation (HF) than woodpeckers, the sister clade of the honeyguides, although the HF enlargement was not significant across all of our analyses. Cluster analyses also revealed that the overall composition of the brain and telencephalon differs greatly between honeyguides and woodpeckers. The relatively smaller brains of the honeyguides may be a consequence of brood parasitism and cerophagy ('wax eating'), both of which place energetic constraints on brain development and maintenance. The inconclusive results of our analyses of relative HF volume highlight some of the problems associated with comparative studies of the HF that require further study.
[Show abstract][Hide abstract] ABSTRACT: We attempt to describe and explain the peculiarly restricted distribution of the globally threatened Ethiopian Bush-crow Zavattariornis stresemanni. At a regional scale, models containing only correlates of land cover suggested a far wider distribution of suitable habitat in north-east Africa than the area actually occupied. However, models including only climate variables predicted the known distribution almost perfectly, and suggested that the species’ area of occupancy is delimited by a pocket of climate that is cooler, dryer and more seasonal than surrounding areas. The predicted probability of occurrence was low outside a narrow range of mean annual temperatures of 17.5–20°C. Within the area predicted to be climatically most suitable, records of Bush-crows were concentrated in 1-km cells of marginally but significantly lower normalised difference vegetation index (NDVI), indicating a preference for areas of lower photosynthetic activity. At a finer spatial scale within a 10-km2 intensive study site in the core of the species’ range, nests were located in 30-m cells of higher NDVI but always close to areas of lower NDVI. These areas of lower NDVI comprise open grassland, which standardised observations of individual birds showed to be the main foraging habitat. However, taller vegetation is also necessary for nesting and roosting; the average height of nests from the ground was nearly 5 m. Therefore, the species’ range appears to be defined primarily by a unique climate pocket within which it shows a preference for park-like habitats of grassland interspersed with taller vegetation, largely the result of clearance of vegetation by people and their associated grazers. The diet appeared unspecialised and a wide range of feeding methods was observed. Models estimate the species’ optimal climatic range to cover around 6,000 km2, of which perhaps 4,500 km2 has suitable land cover. We tentatively estimate the global population to be at least 9,000 breeding pairs, with a potentially larger additional population of non-breeding birds, particularly nest-helpers. Several climate models predict increases in both temperature and precipitation in the southern highlands of Ethiopia. The species’ narrow climatic range suggests that global climate change may therefore pose a serious threat to its long-term survival.
[Show abstract][Hide abstract] ABSTRACT: Maternal effects can influence offspring phenotype with short- and long-term consequences. Yet, how the social environment may influence egg composition is not well understood. Here, we investigate how laying order and social environment predict maternal effects in the sociable weaver, Philetairus socius, a species that lives in massive communal nests which may be occupied by only a few to 100+ individuals in a single nest. This range of social environments is associated with variation in a number of phenotypic and life-history traits. We investigate whether maternal effects are adjusted accordingly. We found no evidence for the prediction that females might benefit from modifying brood hierarchies through an increased deposition of androgens with laying order. Instead, females appear to exacerbate brood reduction by decreasing the costly production of yolk mass and antioxidants with laying order. Additionally, we found that this effect did not depend on colony size. Finally, in accordance with an expected increased intensity of environmental stress with increasing colony size, we found that yolk androgen concentration increased with colony size. This result suggests that females may enhance the competitive ability of offspring raised in larger colonies, possibly preparing the offspring for a competitive social environment.
[Show abstract][Hide abstract] ABSTRACT: Coevolutionary arms races are a powerful force driving evolution, adaptation, and diversification. They can generate phenotypic polymorphisms that render it harder for a coevolving parasite or predator to exploit any one individual of a given species. In birds, egg polymorphisms should be an effective defense against mimetic brood parasites and are extreme in the African tawny-flanked prinia (Prinia subflava) and its parasite, the cuckoo finch (Anomalospiza imberbis). Here we use models of avian visual perception to analyze the appearance of prinia and cuckoo finch eggs from the same location over 40 years. We show that the two interacting populations have experienced rapid changes in egg traits. Egg colors of both species have diversified over time, expanding into avian color space as expected under negative frequency-dependent selection. Egg pattern showed signatures of both frequency-dependent and directional selection in different traits, which appeared to be evolving independently of one another. Host and parasite appear to be closely tracking one another's evolution, since parasites showed closer color mimicry of contemporaneous hosts. This correlational evidence suggests that hosts and parasites are locked in an ongoing arms race in egg appearance, driven by constant change in the selective advantage of different phenotypes, and that coevolutionary arms races can generate remarkably rapid phenotypic change.
The American Naturalist 05/2012; 179(5):633-48. DOI:10.1086/665031 · 4.45 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: Parasites that exploit multiple hosts often experience diversifying selection for host-specific adaptations. This can result in multiple strains of host specialists coexisting within a single parasitic species. A long-standing conundrum is how such sympatric host races can be maintained within a single parasitic species in the face of interbreeding among conspecifics specializing on different hosts. Striking examples are seen in certain avian brood parasites such as cuckoos, many of which show host-specific differentiation in traits such as host egg mimicry. Exploiting a Zambian egg collection amassed over several decades and supplemented by recent fieldwork, we show that the brood parasitic Greater Honeyguide Indicator indicator exhibits host-specific differentiation in both egg size and egg shape. Genetic analysis of honeyguide eggs and chicks show that two highly divergent mitochondrial DNA lineages are associated with ground- and tree-nesting hosts, respectively, indicating perfect fidelity to two mutually exclusive sets of host species for millions of years. Despite their age and apparent adaptive diversification, however, these ancient lineages are not cryptic species; a complete lack of differentiation in nuclear genes shows that mating between individuals reared by different hosts is sufficiently frequent to prevent speciation. These results indicate that host specificity is maternally inherited, that host-specific adaptation among conspecifics can be maintained without reproductive isolation, and that host specificity can be remarkably ancient in evolutionary terms.
Proceedings of the National Academy of Sciences 09/2011; 108(43):17738-42. DOI:10.1073/pnas.1109630108 · 9.81 Impact Factor