Comparative evidence for costs of secondary sexual characters: adaptive vane emargination of ornamented feathers in birds

We used a comparative approach, by comparing bird species with tail ornamentation with sister taxa without ornamentation, to deduce the aerodynamic function of extravagant feather ornaments and the costs of such ornaments in birds. First, the aerodynamic function of tail feather ornaments in birds can be deduced from asymmetry in the width of tail feather vanes, since flightless birds have symmetrical vanes while flying birds without feather exaggeration by sexual selection have asymmetrical vanes. Distal inner vanes at the tip of tail feathers were more narrow in ornamented as compared to nonornamented birds, and vane asymmetry at the tip of the feather was therefore reduced in ornamented species, suggesting marginal aerodynamic function of the distal part of extravagant feather ornaments. Second, the cost of feather ornaments due to parasite drag is proportional to the area of feathers extending beyond the maximum continuous width of the tail, and aerodynamic costs of long tails could therefore be diminished by a reduction in feather width. Consistent with this prediction, the outermost tip of feather ornaments was narrower than the homologous character in nonornamented sister taxa, while the base of the feather had similar width in the two groups of birds. These results suggest that the costs of extravagant ornamentation have been diminished by a reduction in feather width, leading to a reduction in drag. Costs of feather ornaments, as demonstrated by their fine morphology, thus appear to have been extensive during the evolution of these characters.     

Sex-limited expression of ornamental feathers in birds

Extravagant secondary sexual characters show sexual size dimorphismin some species but are completely sex limited in others. Sexualornamentation has been hypothesized to benefit mainly malesthrough sexual selection, but the costs of secondary sexualcharacters initially would be experienced by both sexes. Theevolution of sexual size dimorphism of ornaments and, eventually,the complete sex-limited expression of these characters, willdepend on the effects of sexual and natural selection on thetwo sexes. A phylogenetic analysis controlling for similaritiesdue to common ancestry of 60 independent evolutionary originsof feather ornamentation in birds was used to investigate ecologicalfactors correlated with sexual size dimorphism and sex-limitedexpression of secondary sexual characters. When the size ofan ornament is large relative to body size, the trait willbe particularly costly for females, resulting in selectionfor increased sexual size dimorphism of the ornament. Indeed,sexual size dimorphism of ornaments was positively relatedto the relative size of male ornaments but was unrelated torelative size of female ornaments. Species with polygynousand lekking mating systems with little or no male parentalcare (in particular nest building and incubation) demonstratedsex-limited expression of ornaments as compared to monogamousspecies. Species with no food provisioning of offspring by themale showed a trend for increased sexual size dimorphism ofornaments. Therefore, large natural selection costs duringreproduction imposed by the expression of secondary sexualcharacters are related to the evolution of sexual size dimorphismof ornaments and eventually their complete loss from females.     

Tail length in birds in relation to tail shape,general flight ecology and sexual selection

Relative tail length (longtailedness) of Palearctic birds was assessed by the standardized residuals of log–log regressions of tail length on wing length and tarsus length. The mean degree of tail shortening was greater than mean degree of tail lengthening, but there was a greater frequency of extreme long-tailed than short-tailed species. Longtailedness was greater in ornamental pin, lyre, deep forked and graduated shaped tails. These shapes (except graduated, for which data were lacking) were also relatively long-tailed according to shortest-rectrix lengths, this extra length potentially contributing compensatory lift. In forked tails, tail ratio increased linearly with longtailedness to above the aerodynamic optimum, and thus the most elongated forked tails were also more deeply forked. Tail shortening was marked for rounded tails, a surprising result in view of their slightly ornamental shape. Phylogenetically independent contrasts showed significantly greater longtailedness in graduated than square-tailed species, confirming the species-wide analysis. In phylogenetically independent contrasts of longtailedness and ecological factors, short-tailed species had significantly greater flight distances than medium-tailed species, but long- and medium-tailed species did not differ in migratory distance, foraging distance, overall flight distance or importance of aerial foraging. The data suggest that ecological factors, i.e. natural selection, are more important in the evolution of short-tailedness than longtailedness in birds, and that an additional influence of sexual selection on tail length and shape is also widespread.     

Male barn swallows use different resource allocation rules to produce ornamental tail feathers

Sexual ornaments compete for resources with other functionaltraits. Such resource allocation trade-offs should ensure thehonesty of sexual ornaments according to the Zahavi's handicapprinciple. However, the existence of costly signals could notbe enough to guarantee honesty if different individuals investdifferent proportions of their limited resources in ornaments.Then, a certain level of sexual signaling would correspond toseveral levels of individual condition. Here, we explore whetherthere are different resource allocation rules in tail featherornaments between males within a barn swallow (Hirundo rustica)population and whether these different rules confer differentviability to males. We assessed the proportion of resourcesinvested in ornamental feathers compared with other functionalfeathers moulted and growing during the same period at expensesof the same resources. We found that 1) different males allocatea different proportion of resources to ornamental feathers inrelation to functional feathers and this proportion is repeatablebetween years and 2) male survival likelihood decreased as theproportion of resources allocated to ornamental feathers increased.Survival costs associated with increased investments in ornamentscan maintain the sexual signaling system honest at populationlevel but do not preclude the existence of an array of differentallocation rules between males. Thus, males with different viabilitycan produce ornamental feathers of the same length. These resultsshow that the relationship between male viability and ornamentexpression can be less straightforward than considered previously.     

Resonating feathers produce courtship song

Male Club-winged Manakins, Machaeropterus deliciosus (Aves: Pipridae), produce a sustained tonal sound with specialized wing feathers. The fundamental frequency of the sound produced in nature is approximately 1500 Hz and is hypothesized to result from excitation of resonance in the feathers'' hypertrophied shafts. We used laser Doppler vibrometry to determine the resonant properties of male Club-winged Manakin''s wing feathers, as well as those of two unspecialized manakin species. The modified wing feathers exhibit a response peak near 1500 Hz, and unusually high Q-values (a measure of resonant tuning) for biological objects (Q up to 27). The unmodified wing feathers of the Club-winged Manakin do not exhibit strong resonant properties when measured in isolation. However, when measured still attached to the modified feathers (nine feathers held adjacent by an intact ligament), they resonate together as a unit near 1500 Hz, and the wing produces a second harmonic of similar or greater amplitude than the fundamental. The feathers of the control species also exhibit resonant peaks around 1500 Hz, but these are significantly weaker, the wing does not resonate as a unit and no harmonics are produced. These results lend critical support to the resonant stridulation hypothesis of sound production in M. deliciosus.     

Effects of polyandry on male phenotypic diversity

Polyandry has the potential to affect the distribution of phenotypes and to shape the direction of sexual selection. Here, we explore this potential using Trinidadian guppies as a model system and ask whether polyandry leads to directional and/or diversifying selection of male phenotypic traits. In this study, we compare the phenotypic diversity of offspring from multiply and singly sired broods. To quantify phenotypic diversity, we first combine phenotypic traits using multivariate methods, and then take the dispersion of individuals in multivariate space as our measure of diversity. We show that, when each trait is examined separately, polyandry generates offspring with a higher proportion of bright coloration, indicating directional selection. However, our multivariate approach reveals that this directionality is accompanied by an increase in phenotypic diversity. These results suggest that polyandry (i) selects for the production of sons with the preferred brighter colour phenotypes whereas (ii) enhancing the diversity of male sexual traits. Promoting phenotypic diversity may be advantageous in coping with environmental and reproductive variability by increasing long‐term fitness.     

Patterns of variation in tail ornament size in birds

In recent years several different kinds of sexual selection models have been developed, and tail ornaments in birds have frequently been used as an example of a sexually selected character where the models might apply. However, very little is known about intra- and interpopulation variation in ornament size. We have studied the elongated tail ornaments in four species of whydahs Vidua , the forktailed flycatcher Tyrannus savana and the Asian paradise flycatcher Terpsiphone paradisi. Ornaments were relatively longer in males with the longest tarsi ('heterogony' with positive allometry). Also, tail lengths were remarkably variable within each geographical area, the coefficient of variation (average = 11%) being three times as high as for body size characters. Models, with female preference of ornaments bearing no relation to male viability, usually generate lines of neutral equilibria. Thus, they predict extraordinary variation in ornaments between populations. However, elongated tail ornaments did not show higher geographical variation than the body size characters, suggesting that there is no line of equilibria for these ornaments.     

Exposure to pesticides impairs the expression of fish ornaments reducing the availability of attractive males

Ornament magnitude often reflects a local balance between sexual selection and other sources of natural selection opposing their elaboration. Human activity may disrupt this balance if it modifies the costs of producing, maintaining or displaying the ornaments. When costs are increased, a shortage of acceptable partners may ensue, with consequences commensurate with how stringent (and effective) the process of mate choice is. Here, we show that the expression of ornaments in the viviparous amarillo fish (Girardinichthys multiradiatus) is influenced by embryonic exposure to low concentrations of an organophosphorus insecticide. Male ornamental fin size, dimorphic yellow coloration and display rates were all compromised in exposed fish, but unaffected in their paternal half-sibling controls and in their sisters (morphology and colour). Exposure resulted in smaller fish of both sexes, thus the differential effect by sex was restricted to attributes such as fin size only above the naturally selected magnitude shown by females. Father phenotype predicted offspring morphology of controls, but not of exposed males, which were discriminated against by both control and exposed females. Since stringent female mate choice can result in females refusing to mate with suboptimal mates, this sub-lethal developmental effect can reduce the effective population size of amarillo fish populations.     

Natural variation in the sexually selected feather ornaments of crested auklets (Aethia cristatella) does not predict future survival

We evaluated whether sexually selected crest and auricular plumefeather ornaments of crested auklet (Aethia cristatella) adultscovaried with individual local survival over 11 years (1991–2001).Crested auklets (n = 364 total) were captured near breedingsites, marked with color rings, and local survival estimateswere based on color ring resightings at a breeding colony. Survivalestimates and relationships among local survival and crest length,auricular plume length, mass and tarsus were evaluated usingthe program MARK. The best models included four groups, definedby sex and ease of resighting, that differed in resighting rate(p) but not local survival rate (). This model structure effectivelyexplained sources of variation in local survival and resightabilityamong individuals. The best fitting model showed local survivalrate varying annually, while accounting for differences in resightabilityof marked individuals between the sexes and groups ([t], p[sex*easeof resighting]). Annual local survival varied from 0.940 ±0.029 (SE) in 1993–1994 to 0.767 ± 0.034 in 1997–1998and averaged 0.859 ± 0.019. We found no evidence thatcrested auklet local survival covaried with continuous variationin individuals' ornaments. Simulations indicated that our dataset was sufficient to detect a relationship between local survivaland a covariate that equaled or exceeded a range of 8%. Theimplications for competing sexual selection mechanisms of empiricallymeasured survival–ornament relationships are controversial,but our study emphasizes that survival estimates for such investigationsmust control for confounding factors such as resighting rateas well as have sufficient statistical power and time scaleto be biologically meaningful. Our results are most consistentwith the idea that the conspicuous variation in crested auklet'sshowy ornaments is arbitrary with respect to individual viabilityas quantified by their long-term survival.     

The allometry of fluctuating asymmetry in southern African plants: flowers and leaves

Several studies of fluctuating asymmetry (FA) in animals show that secondary sexual characters used in signalling have a negative relationship between size and asymmetry. Larger sexual traits are presumably more costly to produce, which should lead to greater developmental stress and corresponding increases in asymmetry. In the absence of among individual variation in the ability to handle these costs, the relationship between size and asymmetry should thus be positive. A negative relationship therefore suggests that expression of these traits is condition-dependent. In plants, flowers act as signals for pollinators and may show similar trends to animal signals. Leaves which are uninvolved in signalling should not. Moller & Eriksson (1994) found that 89% of species ( n = 16 of 18) with insect-pollinated flowers showed a negative relationship between petal size and asymmetry, while 79% of species ( n = 15 of 19) showed a positive relationship between leaf size and asymmetry. I carried out a similar study of 18 plant species. The average relationship between petal size and asymmetry did not differ significantly from zero in those species showing measurable FA in flowers ( n = 12). The relationship was significantly negative in one species, and significandy positive in another. On average, leaves in species with FA did not show a significant positive relationship between size and asymmetry ( n = 7). There was no significant difference in the slopes of the relationship between size and asymmetry for leaves and flowers. Levels of floral asymmetry for species with FA were significandy repeatable on individual plants in 33% ( n = 4 of 12) of species, but leaf asymmetry was not significantly repeatable in any species. It is argued that condition-dependence of traits need not result in a negative relationship between size and asymmetry.     

硬叶兜兰居群表型变异研究

为了揭示硬叶兜兰表型变异的规律,以滇东南核心分布地区7个硬叶兜兰居群113个体的18个表型数量性状进行研究,通过变异系数、巢式方差分析、相关性和聚类分析,结果显示:除了叶长、叶宽、花梗粗、苞片长和苞片宽5个数量性状不存在显著差异外,其余数量性状在居群水平上都存在显著差异。各性状变异系数在0.088~0.189之间,花梗长、叶长、叶长宽比、附毛长的变异系数较大,而中萼高、花囊高和宽、花梗粗、合萼高、花瓣宽和长相对变异不大;18个性状表型分化系数(Vst)在2.05%~50.62%之间,平均值达到22.02%,暗示有77.98%表型变异存在于居群内。除雄蕊长和宽表现居群内和居群间的变异相当外,果蒴长、花瓣长和宽、中萼高和宽、合萼高和宽的分化系数分别为36.03%、23.74%、24.55%、28.13%、18.00%、20.56%和18.12%,叶长最小,为2.05%,居群间的变异明显小于居群内。性状关联性研究表明:花瓣、合萼、中萼高、花梗长和叶长宽比高度相关,而叶宽、苞片长、雄蕊性状和中萼高与其余性状关联性不密切,这些性状相对独立,稳定性较高;基于数量指标的欧氏距离聚类表明:7个居群可分为2大类:杨柳井居群和马固居群为近缘支;其余5个居群(斗咀、小坝子、夹寒箐、古林箐和田坝)为一类。     

Sexual selection and temporal phenotypic variation in a damselfly population

Temporal variation in selection can be generated by temporal variation in either the fitness surface or phenotypic distributions around a static fitness surface, or both concurrently. Here, we use within- and between-generation sampling of fitness surfaces and phenotypic distributions over 2 years to investigate the causes of temporal variation in the form of sexual selection on body size in the damselfly Enallagma aspersum. Within a year, when the average female body size differed substantially from the average male body size, male body size experienced directional selection. In contrast, when male and female size distributions overlapped, male body size experienced stabilizing selection when variances in body size were large, but no appreciable selection when the variances in body size were small. The causes of temporal variation in the form of selection can only be inferred by accounting for changes in both the fitness surface and changes in the distribution of phenotypes.     

Sexual selection and condition‐dependence

The handicap theory of sexual selection suggests that females prefer mates who display extravagant ornaments that advertise their quality or condition. It is often assumed that as such ornamental traits undergo sexually‐selected exaggeration, they must inevitably become more sensitive to condition, and thus more informative. Here, we show that this is not necessarily the case. Depending on the precise form of the relationship between trait size and cost, expression may become more or less condition‐dependent as the trait undergoes exaggeration, or may remain unchanged. This leads us to question how much of the information content of sexual signals can be attributed to sexual selection, and how much to pre‐existing, naturally‐selected condition‐dependence.     

Heterochrony, maternal effects, and phenotypic variation among sympatric pupfishes

Variation in ontogeny can produce phenotypic variation both within and among species. I investigated whether changes in timing and rate of growth were a source of phenotypic variation in a putative incipient species group of pupfish (Cyprinodon spp.). On San Salvador Island, Bahamas, sympatric forms of pupfish differ in morphology but show only partial reproductive isolation in the laboratory. Offspring from two forms and two geographical areas and their hybrids were bred in the laboratory, and ontogenetic trajectories of their feeding morphology were followed until maturity. In the Bahamian pupfish the two forms grow along similar size but not shape trajectories. Two heterochronic parameters, onset and rate of growth, alter shape trajectories in the Bahamian pupfish. Similar forms from different geographical areas (Florida and the Bahamas) grow along parallel shape trajectories, differing only in one heterochronic parameter, the onset shape. Hybrids within and between the pupfish forms produced intermediate feeding morphologies that were influenced by their maternal phenotype, suggesting that maternal effects may be a source of phenotypic variation in shape that can persist to maturity. In Cyprinodon, small changes in multiple heterochronic parameters translate into large phenotypic differences in feeding morphology.     

转基因植物的表型变异、分子检测与遗传分析

本讨论了转化方法、体细胞克隆和选育过程等影响转基因植物表型变异的因素,并对转基因植物不同群体的表型变异组成和效应进行了比较分析,提出了转基因植物分子检测和遗传分析的技术策略。多数情况下,分析转基因植物回交后代(BClF1)比分析T1代能获得更可靠和有价值的结论。     

Biological affinity, phenotypic variation and palaeoecology of Tetradium Dana, 1846

Comparisons of morphologies and modes of life of the Ordovician fossil Tetradium Dana, 1846, with chaetetid sponges, tabulate corals, and Recent rhodophytes show that several defining Tetradium characteristics are incompatible with its chaetetid and tabulate classifications. All Tetradium characteristics fit a rhodophyte identification, however, as a calcified uniaxial corticated florideophyte. It is argued from functional morphology that the fundamental subsquare cross-section of the Tetradium tube is an adaptation for close packing, which implies that the basic growth form is compact and should have developed where conditions were optimal. More open forms are derived from it and probably occupied less-favourable environments.
Palaeoecological studies from the Ottawa Valley, Canada, show that the Tetradium growth form is correlated with environmental stress and became more open as salinity increased: i.e. the compact T. fibratum form lived in normal marine conditions, radiating bundles of T. cellulosum tubes in low to middle hypersalinity and single T. syringoporoides tubes in high hypersalinity. Different Tetradium growth forms from the study area are phenotypic variants of a single species, rather than different species and genera. Therefore, classifications that divide Tetradium into different species and genera based only on growth form have no biological basis. There is little evidence of interspecific interactions with Tetradium .