[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: 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.
Brain Behavior and Evolution 04/2013; · 2.89 Impact Factor
[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: 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. · 4.55 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. · 9.81 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: The most virulent avian brood parasites obligately kill host young soon after hatching, thus ensuring their monopoly of host parental care. While the host eviction behaviour of cuckoos (Cuculidae) is well documented, the host killing behaviour of honeyguide (Indicatoridae) chicks has been witnessed only once, 60 years ago, and never in situ in host nests. Here, we report from the Afrotropical greater honeyguide the first detailed observations of honeyguides killing host chicks with their specially adapted bill hooks, based on repeated video recordings (available in the electronic supplementary material). Adult greater honeyguides puncture host eggs when they lay their own, but in about half of host nests at least one host egg survived, precipitating chick killing by the honeyguide hatchling. Hosts always hatched after honeyguide chicks, and were killed within hours. Despite being blind and in total darkness, honeyguides attacked host young with sustained biting, grasping and shaking motions. Attack time of 1-5 min was sufficient to cause host death, which took from 9 min to over 7 h from first attack. Honeyguides also bit unhatched eggs and human hands, but only rarely bit the host parents feeding them.
[Show abstract][Hide abstract] ABSTRACT: Arms races between avian brood parasites and their hosts often result in parasitic mimicry of host eggs, to evade rejection. Once egg mimicry has evolved, host defences could escalate in two ways: (i) hosts could improve their level of egg discrimination; and (ii) negative frequency-dependent selection could generate increased variation in egg appearance (polymorphism) among individuals. Proficiency in one defence might reduce selection on the other, while a combination of the two should enable successful rejection of parasitic eggs. We compared three highly variable host species of the Afrotropical cuckoo finch Anomalospiza imberbis, using egg rejection experiments and modelling of avian colour and pattern vision. We show that each differed in their level of polymorphism, in the visual cues they used to reject foreign eggs, and in their degree of discrimination. The most polymorphic host had the crudest discrimination, whereas the least polymorphic was most discriminating. The third species, not currently parasitized, was intermediate for both defences. A model simulating parasitic laying and host rejection behaviour based on the field experiments showed that the two host strategies result in approximately the same fitness advantage to hosts. Thus, neither strategy is superior, but rather they reflect alternative potential evolutionary trajectories.
Proceedings of the Royal Society B: Biological Sciences 04/2011; 278(1724):3566-73. · 5.68 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: Understanding the evolution of cooperation requires determining the costs and benefits of cooperative behaviour. In cooperative breeders, where nonbreeding individuals assist in raising offspring, these ‘helpers’ are expected to increase the fitness of breeders and hence empirical research has focused on the effect they have on reproductive output and breeder survival. However, the effects of helpers during the postfledging period are poorly known because of the difficulty of tracking fledglings in the wild. Helper presence might be beneficial for fledglings, for example through continuous food delivery or increased predator vigilance, but potential competition between helpers and fledglings, or changes in investment of parents assisted by helpers, could counteract these positive effects and have a negative influence on postfledging survival probabilities or promote dispersal. We investigated the survival of juvenile sociable weavers, Philetairus socius, raised in pairs alone versus pairs with helpers by using capture–mark–recapture methods to control for individual detectability in survival estimation. We found that local survival in the first year was reduced in young raised by groups versus those raised by pairs. This may reflect either higher mortality or emigration of juveniles raised in groups. Hence, our study reveals significant postfledging effects of cooperative breeding that have not been reported previously and that need to be investigated in studies addressing the evolution of cooperative breeding.
[Show abstract][Hide abstract] ABSTRACT: Species are the fundamental units of biology, ecology and conservation, and progress in these fields is therefore hampered by widespread taxonomic bias and uncertainty. Numerous operational techniques based on molecular or phenotypic data have been designed to overcome this problem, yet existing procedures remain subjective or inconsistent, particularly when applying the biological species concept. We address this issue by developing quantitative methods for a classic technique in systematic zoology, namely the use of divergence between undisputed sympatric species as a yardstick for assessing the taxonomic status of allopatric forms. We calculated mean levels of differentiation in multiple phenotypic characters – including biometrics, plumage and voice – for 58 sympatric or parapatric species-pairs from 29 avian families. We then used estimates of mean divergence to develop criteria for species delimitation based on data-driven thresholds. Preliminary tests show that these criteria result in relatively few changes to avian taxonomy in Europe, yet are capable of extensive reassignment of species limits in poorly known tropical regions. While we recognize that species limits are in many cases inherently arbitrary, we argue that our system can be applied to the global avifauna to deliver taxonomic decisions with a high level of objectivity, consistency and transparency.
[Show abstract][Hide abstract] ABSTRACT: The offspring of brood parasitic birds benefit from hatching earlier than host young. A proposed but little-known strategy to achieve this is 'internal incubation', by retaining the egg in the oviduct for an additional 24 h. To test this, we quantified the stage of embryo development at laying in four brood parasitic birds (European cuckoo, Cuculus canorus; African cuckoo, Cuculus gularis; greater honeyguide, Indicator indicator; and the cuckoo finch, Anomalospiza imberbis). For the two cuckoos and the honeyguide, all of which lay at 48 h intervals, embryos were at a relatively advanced stage at laying; but for the cuckoo finch (laying interval: 24 h) embryo stage was similar to all other passerines laying at 24 h intervals. The stage of embryo development in the two cuckoos and honeyguide was similar to that of a non-parasitic species that lay at an interval of 44-46 h, but also to the eggs of the zebra finch Taeniopygia guttata incubated artificially at body temperature immediately after laying, for a further 24 h. Comparison with the zebra finch shows that internal incubation in the two cuckoos and honeyguide advances hatching by 31 h, a figure consistent with the difference between the expected and the observed duration of incubation in the European cuckoo predicted from egg mass. Rather than being a specific adaptation to brood parasitism, internal incubation is a direct consequence of a protracted interval between ovulation (and fertilization) and laying, but because it results in early hatching may have predisposed certain species to become brood parasitic.
Proceedings of the Royal Society B: Biological Sciences 09/2010; 278(1708):1019-24. · 5.68 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: Cuckoo eggs are renowned for their mimicry of different host species, leading to the evolution of host-specific races (or 'gentes') defined by egg colour and pattern. This study aims to test the prediction that another property of parasitic eggs, namely shell strength, might also have experienced divergent selection within cuckoo species. Host races of the common cuckoo Cuculus canorus encountering stronger host rejection have thicker-shelled eggs than those parasitising less discriminating species, as expected if egg strengthening discourages host rejection. Moreover, in the diederik cuckoo Chrysococcyx caprius, eggshell thickness was correlated across cuckoo gentes and host species, as expected if eggshell strength has been involved in coevolutionary interactions. This is the first report of host-specific differences in cuckoo egg properties other than colour and pattern and lends correlational support to the hypothesis that the strong eggshells of brood parasites are an adaptation to reduce host rejection.
Journal of Evolutionary Biology 08/2010; 23(8):1792-9. · 3.48 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: One of the most striking outcomes of coevolution between species is egg mimicry by brood parasitic birds, resulting from rejection behavior by discriminating host parents. Yet, how exactly does a host detect a parasitic egg? Brood parasitism and egg rejection behavior provide a model system for exploring the relative importance of different visual cues used in a behavioral task. Although hosts are discriminating, we do not know exactly what cues they use, and to answer this it is crucial to account for the receiver's visual perception. Color, luminance ("perceived lightness") and pattern information have never been simultaneously quantified and experimentally tested through a bird's eye. The cuckoo finch Anomalospiza imberbis and its hosts show spectacular polymorphisms in egg appearance, providing a good opportunity for investigating visual discrimination owing to the large range of patterns and colors involved. Here we combine field experiments in Africa with modeling of avian color vision and pattern discrimination to identify the specific visual cues used by hosts in making rejection decisions. We found that disparity between host and foreign eggs in both color and several aspects of pattern (dispersion, principal marking size, and variability in marking size) were important predictors of rejection, especially color. These cues correspond exactly to the principal differences between host and parasitic eggs, showing that hosts use the most reliable available cues in making rejection decisions, and select for parasitic eggs that are increasingly mimetic in a range of visual attributes.
Proceedings of the National Academy of Sciences 05/2010; 107(19):8672-6. · 9.81 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: The Critically Endangered Liben Lark (formerly Sidamo Lark) is known only from the Liben Plain of southern Ethiopia, where rapid grassland deterioration is driving the species towards extinction. Fieldwork on the Liben Plain in May 2009 to assess changes in habitat and population since June 2007 recorded a significant deterioration in habitat and decline in numbers. In both 2007 and 2009, birds were associated with areas with greater than average grass cover, and in 2007 with areas of higher grass. However, between 2007 and 2009 there was a significant decline in grass cover and height, a 40% decline in number of birds recorded along repeated transects, and a contraction of 38% in the occupied area of the Liben Plain. Moreover, the cover of bare ground increased more in areas where the species was recorded in 2007 than at random points, suggesting a more rapid degradation of the best sites. There was also a loss to arable agriculture of 8% of the grassland present in 2007. Invading fennel plants increased in number and area on the plain but did not appear to influence the distribution of the lark. An analysis of NDVI showed that grassland deterioration could not be explained by drought, and the most likely explanation is that grassland quality is suffering from overgrazing. Predictive modelling suggests that, apart from a smaller and politically insecure area some 500 km to the north-east near Somalia, there is no suitable habitat for this species elsewhere in the Horn of Africa. As a matter of extreme urgency, cattle exclosures need to be established on the Liben Plain to allow grassland regeneration. This may require the ploughing of land to reduce soil compaction and re-sowing with local grass species. In the longer term, further degradation of the plain should be prevented by, for example, clearing encroaching scrub to increase grassland area and reduce grazing pressure, and by developing sustainable rangeland management practices. These actions have the full and active support of local pastoralists.
[Show abstract][Hide abstract] ABSTRACT: The presence of more than two adults attending the nest is a commonly observed phenomenon in raptors and has been reported in 18% of species of the Falconidae (Kimball et al. 2003). Hypotheses to explain its unusually frequent evolu-tion among raptors have been suggested to differ from those more generally considered likely to apply in birds (Kimball et al. 2003). Hence, it may be useful to document further exam-ples from additional species. In this note we report from two sites in the Northern Cape Province, South Africa, the first observations of helping at the nest in breeding Pygmy Falcons Polihierax semitorquatus, which imply that this species is a hitherto overlooked co-operative breeder. We then assess the evidence for four routes to helping which seem possible in this species: (i) co-operative polyandry (Faaborg et al. 1995, Malan et al. 1997), (ii) delayed disper-sal of grown offspring due to territory limitation or life-history considerations (reviewed by Hatchwell and Komdeur 2000), (iii) thermoregulatory benefits of group formation (Kimball et al. 2003), and (iv) group defence. In the arid zone of southern Africa, Pygmy Falcons breed almost exclusively in nest chambers of the communally-breeding Sociable Weaver Philetairus socius (Mendelsohn 1997), as well as infrequently using nests of Red-billed Buffalo Weavers Bubalornis niger and possibly Glossy Starlings Lamprotornis nitens (Marx and Adam 1986). On Benfontein Game Farm (28°52'S, 24°51'E) near Kimberley, South Africa, about 30 colonies of Sociable Weavers breed in a c. 1 000 ha tract of Kalahari sandveld with Camelthorn Acacia erioloba woodland. At least one pair of Pygmy Falcons has bred here regularly since at least 1993 (MD Anderson pers. comm.). During summer 2002, a male falcon was seen actively calling in the vicinity of one colony from 24 October, and from 25 October a female sat intermittently in a specific chamber, before any eggs were laid. During early November three eggs were laid and they hatched between 29 November and 5 December. On 24 December blood samples were taken from the nestlings. On 10 January 2003 CS and EH captured the attending adults using mist-nets, and caught two provisioning males within minutes of each other, followed by one adult female, which is clear evidence of helping at the nest. The adults respectively carried a cicada, a grasshopper and a gecko. Blood samples were also taken from the adults. This could be either an instance of co-operative polyandry or true monogamy with helper(s), which in this instance happened to be male. If the former, then genetic analyses ought to reveal shared paternity between the two attending males, according to the definition of polyandry. At the second site, south of Twee Rivieren (26°45'S, 20°37'E), OAER made observations at two active chambers of breeding Pygmy Falcons during October–November 2003. One chamber was attended by two males and a female, and the other by one male and two females. The for-mer instance provides further evidence of helping at the nest by more than one male Pygmy Falcon. However, in the lat-ter instance, one female was subsequently chased away by the other female before egg-laying began, which suggests that the second female was not a helper but rather disputing the breeding site. It would also be interesting to reveal the possible patterns of helping and paternity in situations where more than one pair of falcons breed in a single nest mass, one incident of which has been published (Braine and Braine 1968) and a second observed by OAER at Twee Rivieren in November 2003. At a third site in the Kgalagadi Transfrontier Park, during 1984–1992, CWS studied c. 25 Pygmy Falcon territories (not all active every year) along a 90km stretch of the Nossob Riverbed, centered on Nossob Camp (25°25'S, 20°35'E). A third individual was observed feeding nestlings on three occasions, and in at least two cases the helper was male. In each case the helper was a ringed member of a previous brood of the same pairs. Hence, it seems probable that the second male did not gain a share of paternity, though there was one observation of attempted incestuous polyandry, in which a female was observed copulating with one of her offspring from the previous breeding season. Why do Pygmy Falcons sometimes breed co-operative-ly? In the absence of molecular genetic information on pater-nity and relatedness, it is difficult to distinguish co-operative polyandry from alternatives but we nonetheless discuss four applicable hypotheses in the light of what is known of the Pygmy Falcon's natural history.
[Show abstract][Hide abstract] ABSTRACT: Bright yellow to red signals used in mate choice or intrasexual competition are based on carotenoid pigments that are hypothesized to be traded between physiological functions and coloration. These signals have recently been shown to be influenced by maternal effects. Indeed, yolk-derived carotenoids are essential for embryos to develop efficient carotenoid metabolism in posthatching life. Maternal effects facilitate adaptation to environmental variability and influence the evolution of phenotypic traits such as secondary sexual signals. Here we propose that maternal investment in yolk carotenoids promotes the evolution of carotenoid-based ornaments. We conducted a comparative analysis of lipid-soluble antioxidants (carotenoids and vitamins A and E) in the eggs of 112 species of bird. Species with large clutch sizes deposited higher yolk concentrations of the three antioxidants. There was a significant positive relationship between yolk carotenoids and the expression of male carotenoid-based signals, but not between yolk carotenoids and sexual dichromatism in these signals. These relationships were specific to carotenoids, as they were not found for vitamins A and E. This provides evidence consistent with the hypothesis that maternal effects mediated by yolk carotenoids play a role in the evolution of carotenoid-based signals as a response to sexual selection, likely based on organizational effects of carotenoids during embryo development.
The American Naturalist 09/2009; 174(5):696-708. · 4.55 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: Rangeland degradation by livestock threatens several restricted-range species, but is largely overlooked by conservation biologists. The Sidamo lark Heteromirafra sidamoensis, confined to the Liben Plain grassland in southern Ethiopia, is critically endangered by bush encroachment, permanent settlement and agricultural conversion. Its global range was previously estimated at 760 km2, but in 2007–2008 available habitat covered<35 km2. Density estimates from multi-model inference analysis of distance transect data provided a global population estimate of 90–256 adults (possibly with a serious sex-ratio bias towards males). Logistic regression models of habitat selection showed that males preferentially occurred in areas of grassland with greater cover of medium-length grass (5–15 cm), less cover of bare ground and fewer bushes. Habitat transects extending outward from its core range revealed massive and rapid bush encroachment, corroborating information from semi-structured interviews. The survival of both local Borana pastoralism and this species – mainland Africa's likeliest first avian extinction – depends on restoring seasonal patterns of grazing, resisting agricultural conversion of grasslands, reversing fire suppression policies and clearing bush.