Evidence for aggressive mimicry in an adult brood parasitic bird,and generalized defences in its host

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.     

EVOLUTION OF PLUMAGE COLOR IN MALE PIED FLYCATCHERS (FICEDULA HYPOLEUCA): EVIDENCE FOR FEMALE MIMICRY

We present the first evidence for sexual deception by female mimicry in birds. Using live, caged birds we show that territorial male pied flycatchers behave aggressively toward bright-colored males but display sexually toward female-like male intruders. We also show that the males that are fooled are those that lack recent sexual experience. All male pied flycatchers are dull-colored in winter. It is possible that young males are more constrained during the spring molt than older males since the former are more dull-colored in spring. According to the molt-constraints hypothesis a subadult plumage would be maladaptive in the breeding season. Analysis of male settling pattern at breeding sites in spring suggests that brownish males are allowed to settle closer to already-established males than dark-colored males. This result suggests an adaptive value of having a subadult plumage color, in particular for young males arriving late from spring migration. However, we also show that mimicry incurs a cost, that of increased aggression from females, which may explain why female-like males have reduced mating success.     

Evolution of Dendrocolaptes platyrostris (Aves: Furnariidae) between the South American open vegetation corridor and the Atlantic forest

The open vegetation corridor of South America is a region dominated by savanna biomes. It contains forests (i.e. riverine forests) that may act as corridors for rainforest specialists between the open vegetation corridor and its neighbouring biomes (i.e. the Amazonian and Atlantic forests). A prediction for this scenario is that populations of rainforest specialists in the open vegetation corridor and in the forested biomes show no significant genetic divergence. We addressed this hypothesis by studying plumage and genetic variation of the Planalto woodcreeper Dendrocolaptes platyrostris Spix (1824) (Aves: Furnariidae), a forest specialist that occurs in both open habitat and in the Atlantic forest. The study questions were: (1) is there any evidence of genetic continuity between populations of the open habitat and the Atlantic forest and (2) is plumage variation congruent with patterns of neutral genetic structure or with ecological factors related to habitat type? We used cytochrome b and mitochondrial DNA control region sequences to show that D. platyrostris is monophyletic and presents substantial intraspecific differentiation. We found two areas of plumage stability: one associated with Cerrado and the other associated with southern Atlantic Forest. Multiple Mantel tests showed that most of the plumage variation followed the transition of habitats but not phylogeographical gaps, suggesting that selection may be related to the evolution of the plumage of the species. The results were not compatible with the idea that forest specialists in the open vegetation corridor and in the Atlantic forest are linked at the population level because birds from each region were not part of the same genetic unit. Divergence in the presence of gene flow across the ecotone between both regions might explain our results. Also, our findings indicate that the southern Atlantic forest may have been significantly affected by Pleistocene climatic alteration, although such events did not cause local extinction of most taxa, as occurred in other regions of the globe where forests were significantly affected by global glaciations. Finally, our results neither support plumage stability areas, nor subspecies as full species. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 801–820.     

Evolutionary significance of geographic variation in a plumage-based foraging adaptation: an experimental test in the slate-throated redstart (Myioborus miniatus)

Geographic variation in the plumage pattern of birds is widespread but poorly understood, and in very few cases has its evolutionary significance been investigated experimentally. Neotropical warblers of the genus Myioborus use their contrasting black-and-white plumage to flush insect prey during animated foraging displays. Although previous experimental work has demonstrated that white plumage patches are critical to flush-pursuit foraging success, the amount of white in the plumage shows considerable interspecific and intraspecific geographic variation. We investigated the evolutionary significance of this geographic variation by experimentally decreasing or increasing the amount of white in the tail of slate-throated redstarts (Myioborus miniatus comptus) from Monteverde, Costa Rica, to mimic the natural extremes of tail pattern variation in this species. In addition to measuring the effects of plumage manipulation on foraging performance, we performed field experiments measuring the escape response of a common insect prey species (an asilid fly) using model redstarts representing four different Myioborus plumage patterns. Our experiments were designed to test four hypotheses that could explain geographic variation in plumage pattern. Compared to controls, experimental birds with reduced-white tails that mimic the plumage pattern of M. miniatus hellmayri of Guatemala showed significant reductions in flush-pursuit foraging performance. In contrast, the addition of white to the tail to mimic the plumage pattern of M. miniatus verticalis of Bolivia had no significant effect on foraging performance of Costa Rican redstarts. In field experiments with asilid flies, model redstarts simulating the plumage of M. miniatus comptus of Costa Rica and M. miniatus verticalis of Bolivia elicited greater responses than did models of other Myioborus taxa with either less or more white in the plumage. The results of our experiments with both birds and insects allow us to reject two hypotheses for geographic variation in plumage pattern: (1) that geographic variation is a nonadaptive result of genetic drift, and (2) that selection for enhanced flush-pursuit foraging performance generally favors increased white in the plumage, but evolutionary trade-offs constrain the evolution of extensive patches of white in some geographic regions. Instead, our results suggest that geographic variation in the plumage pattern of Myioborus redstarts reflects adaptation to regional habitat characteristics that enhances flush-pursuit foraging performance.     

A new genus and species of passerine from the eastern rain forest of Madagascar

Cryptosylvicola randrianasoloi is described as a new genus and species in the family Sylviidae. This bird is endemic to the rain forests of eastern Madagascar, where it is restricted to elevations between 900 and 2100 m a.s.l. Cryptosylvicola randrianasoloi is distinguished from other Malagasy, African and Asian taxa by plumage colouration, external morphological characters and voice. The discovery of this new taxon in several well-known areas of Madagascar, and the fact that it has a relatively broad distribution in that country, indicates that much still remains to be learned about the biota of the island.     

Unmelanized plumage patterns in Old World leaf warblers do not correspond to sequence variation at the melanocortin-1 receptor locus (MC1R)

Evolutionary changes in patterns and coloration of plumage are likely to represent a major mechanism for speciation among birds, yet the molecular basis for such changes remains poorly understood. Recently much attention has focused on the melanocortin-1 receptor (MC1R) as a candidate locus for determining the level and extent of epidermal melanin deposition. We tested the hypothesis that MC1R sequence variation is associated with interspecific variation in unmelanized plumage pattern elements in Old World leaf warblers (genus Phylloscopus). This genus is characterized by a variety of plumage patterns that nonetheless vary along similar lines. Species vary in the presence or absence of pale (unmelanized) pattern elements against a dark background, and these patterns are used in species recognition and courtship. We sequenced most of the MC1R coding region for eight Phylloscopus species, representing the full range of plumage patterns found in this genus. Although MC1R sequence varied among species, this variation was not related to melanin-based plumage variation. Rather, evolution of this locus in these birds appears to be conservative. Ratios of nonsynonymous to synonymous substitutions (dN/dS) were consistently low, suggesting that strong purifying selection has operated at this locus, and likelihood ratio testing revealed no evidence of variable selective pressures among lineages or across codons. Adaptive evolution at MC1R may be constrained by the adaptive importance of plumage pattern elements in this genus.     

Sympatry drives colour and song evolution in wood-warblers (Parulidae)

Closely related species often exhibit similarities in appearance and behaviour, yet when related species exist in sympatry, signals may diverge to enhance species recognition. Prior comparative studies provided mixed support for this hypothesis, but the relationship between sympatry and signal divergence is likely nonlinear. Constraints on signal diversity may limit signal divergence, especially when large numbers of species are sympatric. We tested the effect of sympatric overlap on plumage colour and song divergence in wood-warblers (Parulidae), a speciose group with diverse visual and vocal signals. We also tested how number of sympatric species influences signal divergence. Allopatric species pairs had overall greater plumage and song divergence compared to sympatric species pairs. However, among sympatric species pairs, plumage divergence positively related to the degree of sympatric overlap in males and females, while male song bandwidth and syllable rate divergence negatively related to sympatric overlap. In addition, as the number of species in sympatry increased, average signal divergence among sympatric species decreased, which is likely due to constraints on warbler perceptual space and signal diversity. Our findings reveal that sympatry influences signal evolution in warblers, though not always as predicted, and that number of sympatric species can limit sympatry''s influence on signal evolution.     

Delayed plumage maturation and delayed reproductive investment in birds

Delayed plumage maturation is the delayed acquisition of a definitive colour and pattern of plumage until after the first potential breeding period in birds. Here we provide a comprehensive overview of the numerous studies of delayed plumage maturation and a revised theoretical framework for understanding the function of delayed plumage maturation in all birds. We first distinguish between hypotheses that delayed plumage maturation is attributable to a moult constraint with no adaptive function and hypotheses that propose that delayed plumage maturation is a component of an adaptive life‐history strategy associated with delayed reproductive investment. We then recognize three potential benefits of delayed plumage maturation: crypsis, mimicry and status signaling. Evidence suggests that delayed plumage maturation is not a consequence of developmental constraints and instead represents a strategy to maximize reproductive success in circumstances where young adults cannot effectively compete with older adults for limited resources, particularly breeding opportunities. A multi‐factorial explanation that takes into account lifespan and the degree of competition for limited breeding resources and that combines the benefits of an inconspicuous appearance with the benefits of honest signaling of reduced competitiveness provides a general explanation for the function of delayed plumage maturation in most bird species. Delayed plumage maturation should be viewed as a component of alternative reproductive strategies that can include delay in both plumage and sexual development. Such strategies are frequently facultative, with individuals breeding prior to the acquisition of definitive plumages when conditions are favourable. Presumably, the benefits of delayed plumage maturation ultimately enhance lifetime reproductive success, and studying delayed plumage maturation within the context of lifetime reproductive success should be a goal of future studies.     

Interspecific social dominance mimicry in birds

Interspecific social dominance mimicry (ISDM) is a proposed form of social parasitism in which a subordinate species evolves to mimic and deceive a dominant ecological competitor in order to avoid attack by the dominant, model species. The evolutionary plausibility of ISDM has been established previously by the Hairy‐Downy game (Prum & Samuelson). Psychophysical models of avian visual acuity support the plausibility of visual ISDM at distances ～>2–3 m for non‐raptorial birds, and ～>20 m for raptors. Fifty phylogenetically independent examples of avian ISDM involving 60 model and 93 mimic species, subspecies, and morphs from 30 families are proposed and reviewed. Patterns of size differences, phylogeny, and coevolutionary radiation generally support the predictions of ISDM. Mimics average 56–58% of the body mass of the proposed model species. Mimics may achieve a large potential deceptive social advantage with <20% reduction in linear body size, which is well within the range of plausible, visual size confusion. Several, multispecies mimicry complexes are proposed (e.g. kiskadee‐type flycatchers) which may coevolve through hierarchical variation in the deceptive benefits, similar to Müllerian mimicry. ISDM in birds should be tested further with phylogenetic, ecological, and experimental investigations of convergent similarity in appearance, ecological competition, and aggressive social interactions between sympatric species. Evolutionary explanations of mimicry must consider the possibility that mimics evolve to deceive model species themselves. © 2014 The Linnean Society of London     

Why are there so many mimicry rings? Correlations between habitat,behaviour and mimicry in Heliconius butterflies

In the new world tropics there is an extravagant array of sympatric butterfly mimicry rings. This is puzzling under strictly coevolutionary (Müllerian) mimicry: all unpalatable species should converge as ‘co-mimics' to the same pattern. If mimicry has usually evolved in unpalatable species by one-sided (Batesian) evolution, however, it is easy to see that mimicry rings centred on different models could remain distinct. If mimicry rings were also segregated by habitat, a diversity of mimicry rings could be stabilized. In this paper we report correlations between behaviour and mimicry of nine unpalatable Heliconius species. It is already known that co-mimics fly in similar habitats, and non-mimics fly in different habitats, although there is much overlap. Contrary to a previous report, we find little difference in flight heights of heliconiine mimicry rings; all species fly from ground level to the canopy. However, co-mimics roost at night in similar habitats and at similar heights above the ground, but in different habitats and at different heights from species in other mimicry rings. Heliconius (especially the erato taxonomic group) are renowned for roosting gregariously; and co-mimics roost gregariously with each other more often than with non-mimics. Gregarious roosting is therefore common between species, as well as within species. There are thus strong links between mimicry and behavioural ecology in Heliconius. The paradoxical correlation between nocturnal roosting and visual mimicry is presumably explained by bird predation at dusk when roosts are forming, or at dawn before they have disbanded. Direct evidence of predation is lacking, but there are high rates of disturbance by birds at these times. These results, together with knowledge of the phylogeny of Heliconius, suggest that species from the melpomene-group of Heliconius have radiated to occupy mimetic niches protected by model species in the Ithomiinae and the erato-group of Heliconius. A variety of sympatric mimicry rings is apparently maintained because key models fail to converge, while more rapidly-evolving unpalatable mimics evolve towards the colour patterns of the models. The maintenance of mimetic diversity would be aided by the habitat and behavioural differences between mimicry rings revealed here, provided that different predators are found in different habitats. This explanation for the maintenance of multiple mimicry rings is more plausible for Heliconius mimicry than alternatives based on visual mating constraints, thermal ecology, or camouflage.     

Different modes of evolution in males and females generate dichromatism in fairy‐wrens (Maluridae)

Sexual dichromatism in birds is often attributed to selection for elaboration in males. However, evolutionary changes in either sex can result in plumage differences between them, and such changes can result in either gains or losses of dimorphism. We reconstructed the evolution of plumage colors in both males and females of species in Maluridae, a family comprising the fairy‐wrens (Malurus, Clytomias, Sipodotus), emu‐wrens (Stipiturus), and grasswrens (Amytornis). Our results show that, across species, males and females differ in their patterns of color evolution. Male plumage has diverged at relatively steady rates, whereas female coloration has changed dramatically in some lineages and little in others. Accordingly, in comparisons against evolutionary models, plumage changes in males best fit a Brownian motion (BM) model, whereas plumage changes in females fit an Ornstein Uhlenbeck (OU) multioptimum model, with different adaptive peaks corresponding to distributions in either Australia or New Guinea. Levels of dichromatism were significantly associated with latitude, with greater dichromatism in more southerly taxa. Our results suggest that current patterns of plumage diversity in fairy‐wrens are a product of evolutionary changes in both sexes, driven in part by environmental differences across the distribution of the family.     

Song mimicry of Black-bellied Firefinch Lagonosticta rara and other finches by the brood-parasitic Cameroon Indigobird Vidua camerunensis in West Africa

Brood-parasitic finches Vidua spp. mimic songs of their foster species, with most Vidua species both mimicking songs and parasitizing nests of a single estrildid finch species. We describe a behavioural radiation in the Cameroon Indigobird Vidua camerunensis . Local populations are polymorphic in behaviour, each male mimicking songs of a single species, with certain males mimicking songs of one species and other males mimicking songs of another host species. The species most often mimicked in song are Black-bellied Firefinch Lagonosticta rara and African Firefinch L. rubricata ; other species mimicked in song are Brown Twinspot Clytospiza monteiri and Dybowski's Twinspot Euschistospiza dybowskii . Indigobirds in the different mimicry song populations do not differ morphologically in plumage colour or size. The lack of morphological differences between male indigobirds with different mimicry songs is consistent with a recent behavioural radiation through host shifts, perhaps facilitated by environmental change associated with prehistoric cultivation of grain. The mimicry song populations of indigobirds, behaviourally imprinted upon different host species, support the idea of a process of speciation driven by a shift to new host species.     

Reconstructing plumage evolution in orioles (Icterus): repeated convergence and reversal in patterns

Several empirical studies suggest that sexually selected characters, including bird plumage, may evolve rapidly and show high levels of convergence and other forms of homoplasy. However, the processes that might generate such convergence have not been explored theoretically. Furthermore, no studies have rigorously addressed this issue using a robust phylogeny and a large number of signal characters. We scored the appearance of 44 adult male plumage characters that varied across New World orioles (Icterus). We mapped the plumage characters onto a molecular phylogeny based on two mitochondrial genes. Reconstructing the evolution of these characters revealed evidence of convergence or reversal in 42 of the 44 plumage characters. No plumage character states are restricted to any groups of species higher than superspecies in the oriole phylogeny. The high frequency of convergence and reversal is reflected in the low overall retention index (RI = 0.66) and the low overall consistency index (CI = 0.28). We found similar results when we mapped plumage changes onto a total evidence tree. Our findings reveal that plumage patterns and colors are highly labile between species of orioles, but highly conserved within the oriole genus. Furthermore, there are at least two overall plumage types that have convergently evolved repeatedly in the three oriole clades. This overall convergence leads to significant conflict between the molecular and plumage data. It is not clear what evolutionary processes lead to this homoplasy in individual characters or convergence in overall pattern. However, evolutionary constraints such as developmental limitations and genetic correlations between characters are likely to play a role. Our results are consistent with the belief that avian plumage and other sexually selected characters may evolve rapidly and may exhibit high homoplasy. The overall convergence in oriole plumage patterns is an interesting evolutionary phenomenon, but it cautions against heavy reliance on plumage characters for constructing phylogenies.     

Evolution of an avian pigmentation gene correlates with a measure of sexual selection

The extravagant plumage traits of male birds are a favourite example of sexual selection. However, to date the units that selection is acting upon, the genes themselves have been a 'black box'. Here, we report evidence of change driven by sexual selection at a pigmentation gene locus in the galliform birds. Across species, we find a correlation between the rate of amino acid change (dN/dS) at this locus (MC1R) and the degree of sexual dichromatism, which we use as a measure of the strength of sexual selection. There is no evidence for a similar pattern in any of five other loci (four candidate and one control locus). This is consistent with previous work on colour polymorphisms and suggests that MC1R may be a key target for selection acting on plumage colour. The pattern of selection at MC1R seems to be consistent with the continuous or cyclical evolution of traits and preferences that is the outcome of several Fisherian and good-genes models of sexual selection. In contrast, we found no support for models of sexual selection that predict an increase in purifying selection as a result of purging of deleterious mutations or for models that predict an increased rate of mutation in association with stronger sexual selection.     

Evolutionary innovation and diversification of carotenoid‐based pigmentation in finches

The ornaments used by animals to mediate social interactions are diverse, and by reconstructing their evolutionary pathways we can gain new insights into the mechanisms underlying ornamental innovation and variability. Here, we examine variation in plumage carotenoids among the true finches (Aves: Fringillidae) using biochemical and comparative phylogenetic analyses to reconstruct the evolutionary history of carotenoid states and evaluate competing models of carotenoid evolution. Our comparative analyses reveal that the most likely ancestor of finches used dietary carotenoids as yellow plumage colorants, and that the ability to metabolically modify dietary carotenoids into more complex pigments arose secondarily once finches began to use modified carotenoids to create red plumage. Following the evolutionary “innovation” that enabled modified red carotenoid pigments to be deposited as plumage colorants, many finch species subsequently modified carotenoid biochemical pathways to create yellow plumage. However, no reversions to dietary carotenoids were observed. The finding that ornaments and their underlying mechanisms may be operating under different selection regimes—where ornamental trait colors undergo frequent reversions (e.g., between red and yellow plumage) while carotenoid metabolization mechanisms are more conserved—supports a growing empirical framework suggesting different evolutionary patterns for ornaments and the mechanistic innovations that facilitate their diversification.     

Multiple-brooding in birds of prey: South African Black Sparrowhawks Accipiter melanoleucus extend the boundaries

Multiple-brooding (raising more than one brood of young in quick succession) occurs infrequently in raptors and is generally restricted to either smaller species with shorter nesting periods, co-operative breeders or species capable of capitalizing on conditions of prolonged food abundance whenever they occur. This paper presents the first recorded cases of multiple-brooding in the Black Sparrowhawk Accipiter melanoleucus from two distinct locales in South Africa. In the Western Cape (Cape Peninsula), four attempts to multiple-brood were recorded in four different years, involving three distinct pairs of birds, and in KwaZulu–Natal (Eshowe), three distinct pairs of Sparrowhawks successfully multiple-brooded on several occasions over a 5-year study period. These results establish the Black Sparrowhawk as one of only two relatively large, monogamous raptor species, and the only specialist bird-eating raptor, in which multiple-brooding has been recorded with any frequency . The species' capacity to thrive in human-modified environments (i.e. alien plantations) and particularly to exploit associated foraging opportunities (e.g. high densities of doves and pigeons in suburban areas) may, at least partly, account for the instances of multiple-brooding reported here. We suggest that biologists be more vigilant for cases of multiple-brooding in raptors, as it is possible that this trait is more common than originally thought and has previously been overlooked.     

Are simple plumage traits sufficient for species discrimination by harrier males?

I investigated whether plumage features are sufficient for species recognition in two sibling sympatric raptor species: the Montagu's harrier Circus pygargus and the hen harrier C. cyaneus . These species are similar in body size and differ slightly in plumage. During prelaying and laying periods, I placed artificial models close to nests of both species, simulating a conspecific male harrier (male of the same species), a heterospecific male harrier (male of the other species) or a wood pigeon Columba palumbus (acting as a control of the same size and colour). All of them represented birds perched in a neutral posture, so that they could only be recognised by their plumage differences. The use of plumage traits during this period would be particularly relevant in discriminating between their own and other species, offering the opportunity to avoid unnecessary fights, and therefore maximising the benefits of defence. I measured male responsiveness by recording aggressive behaviour towards each model (aggressive rate and time spent close to the female). Male presence time was significantly longer during conspecific decoy presentation than during heterospecific decoy presentation. Furthermore, males of both species were more aggressive towards the conspecific decoy than towards the heterospecific one, and no attacks towards the control were recorded. These results provide evidence that these birds are able to recognise plumage features of their own and the sibling species.     

Delayed plumage maturation in green‐backed flycatchers (Ficedula elisae): An evidence of female mimicry

Delayed plumage maturation (DPM) is the delayed acquisition of an adult color and pattern of plumage until after the first potential breeding period. Among the hypotheses proposed to explain DPM, the female mimicry hypothesis (FMH) has received considerable attention. FMH predicts that after‐second‐calendar‐year (ASY) males should attack ASY males more than second‐calendar‐year (SY) males and females, while no difference between the two latter. Few studies have been thought as support for FMH, while in fact most of them give no conclusive evidence to the assumption of FMH that ASY males are unable to distinguish SY males from females. Thus, other support besides behavioral experiments, such as it is physiologically impossible to discriminate SY males and females from avian perspective, is important in testing FMH. We studied color differences in six plumage patches between ASY male, SY male and female by analyzing reflectance spectra in the avian visual system, and tested the prediction of FMH by conducting intrusion experiments on territorial males with three kinds of conspecifics (i.e., ASY males, SY males and females) as territory “intruders” in the green‐backed flycatcher (Ficedula elisae). It is physiologically impossible for ASY males to distinguish SY males and females through the plumage color of crown, mantle, rump, and throat patches, moreover difficult to distinguish through breast and belly patches even under reasonable viewing conditions. ASY males were more aggressive toward ASY males than SY males and females with no differences between the two latter. SY males showed a slightly but not significantly higher attack intensity on ASY males than SY males and females. Our results suggest that female mimicry is more likely to be the explanation for DPM in the green‐backed flycatcher. To our knowledge, this is the first study combining avian visual system and behavioral experiments in testing hypotheses of DPM.     

Frugivory in four sympatric lemurs: implications for the future of Madagascar's forests

Although some conservationists accept that not all species can be saved, we illustrate the difficulty in deciding which species are dispensable. In this article, we examine the possibility that the integrity of a forest relies on its entire faunal assemblage. In Madagascar, one faunal group, the lemurs, accounts for the greatest biomass and species richness among frugivores. For example, 7 of the 13 sympatric lemur species in Madagascar's eastern rainforests consume primarily fruit. Because of this, we suggest that some tree species may rely heavily on particular lemur taxa for both seed dispersal and germination. In Ranomafana National Park, the diets for four of the day-active lemur frugivores have been documented during annual cycles over a 5-year period. We predicted that, although the fruit of some plant taxa would be exploited by multiple lemur species, the fruit of others would be eaten by one lemur species alone. Analyses reveal that while lemurs overlap in a number of fruit taxa exploited, 46% (16/35) of families and 56% (29/52) of genera are eaten exclusively by one lemur species. We, therefore, predict local changes in forest composition and structure if certain of these lemur species are eliminated from a forest owing to hunting, disease, or habitat disturbance. We also suggest that this result may be of global significance because carbon sequestration by the tropical forests in Madagascar may be reduced as a result of this predicted change in forest composition.     

Sexual dimorphism and dichromatism in Steere's Liocichla (Liocichla steerii)

ABSTRACT.   Although sexual differences in birds can be extreme, differences between males and females in body size and plumage color are more subtle in many species. We used a genetic-based approach to determine the sex of male and female Steere's Liocichla ( Liocichla steerii ) and examine the degree of size dimorphism and plumage dichromatism in this apparently monomorphic species. We found that males were significantly larger than females. In addition, Steere's Liocichla have a prominent yellow plumage patch on the lores that was significantly larger in males than females for both live birds and museum specimens. We also used reflectance spectrometry to quantify the color of the yellow-green breast feathers of Steere's Liocichla and found no significant differences between males and females in brightness, intensity, saturation, or hue. However, females tended to have brighter breast plumage, particularly at long wavelengths. Collectively, these color variables were useful in discriminating birds according to sex when used in a discriminant function analysis. Our study suggests that sexual selection may be more widespread than once assumed, even among birds considered monomorphic, and emphasizes the need for additional data from tropical and subtropical species.