Multiple Ornaments in Male Northern Cardinals, Cardinalis cardinalis, as Indicators of Condition

Investigations of male ornaments in the context of sexual selection have tended to focus on single ornaments, although many species of birds possess multiple ornaments. Understanding the evolution of multiple ornaments requires knowledge of correlations among ornaments in the same individual and the extent to which ornament expression reflects individual condition and behavior. Variation in four male ornaments in socially monogamous, biparental northern cardinals (Cardinalis cardinalis) was related to body size, indices of condition, level of paternal care, and reproductive success. Redness of breast plumage positively predicted body size and negatively predicted nestling feeding rate. Bill color predicted current body condition, with birds with redder bills in better condition. Birds with smaller black face masks had greater reproductive success. These results are consistent with the hypothesis that different ornaments in male cardinals provide information on different aspects of condition and behavior.     

The evolution of female preferences for multiple indicators of quality

In a variety of species, females exhibit preferences for multiple male ornaments. Several hypotheses have been proposed to explain this phenomenon. Which, if any, of these hypotheses is the most plausible in general remains largely unresolved based on the available empirical data. Yet theoretical studies conclude that the evolution of preferences for multiple signals of male quality is unlikely, especially when the use of an additional cue in mate choice strongly increases the overall cost of choice. This would imply that most male courtship characters do not reflect the male's genetic quality but instead evolved through Fisherian sexual selection. However, the existing models focus on ornaments that signal overall genetic quality and do not address the possibility that different ornaments provide information about different aspects of quality. Therefore, we develop a model in which the ornaments act as signals for distinct quality components. When the ornaments provide overlapping information about these quality components, we retrieve the results of earlier models. However, when the ornaments provide independent information, preferences for multiple ornaments may evolve, even when exhibiting multiple preferences is costly. We discuss our results in relation to the multiple-message and redundant-signal hypotheses for ornament diversity and identify parallels between Fisherian and good-genes mechanisms for the evolution of multiple ornaments.     

Long-term persistence of exaggerated ornaments under Fisherian runaway despite costly mate search

Exaggerated ornaments often evolve due to the mating preferences of the opposite sex. Genetic correlations between preferences and ornaments can lead both traits to elaborate dramatically in tandem, in a process known as ‘Fisherian runaway’. However, in most previous models of Fisherian runaway, elaborate ornaments are not expected to persist when preferences are consistently costly to the choosing sex. In contrast, we show here that exaggerated male ornaments can be maintained long term even when females must pay a cost to choose their mates. Preferences per se are not costly in our model, but females can only act on their preferences by investing resources in mate search. We predict that mate search effort should decrease with the cost of sampling additional mates and increase with the number of possible ornaments that females can choose from. The potential for multiple exaggerated ornaments to coexist depends on subtleties of their cost structure: strict trade-offs (additive costs) favour sequential ornament evolution, whereas looser trade-offs (multiplicative costs) allow for coexistence. Lastly, we show that pleiotropy affecting both ornaments and preferences makes it difficult for Fisherian runaway to initiate, increasing the evolutionary time until ornamentation. Our model highlights the important but neglected role of mate search effort in sexual selection.     

Patterns of variation in ornaments of Crested Auklets Aethia cristatella

We investigated patterns of variation of feather and bill ornaments of Crested Auklets Aethia cristatella , a monogamous seabird, based on 963 individuals measured in the years 1990 to 1998. Three prominent ornaments were displayed: a forehead crest, composed of 11–31 curved feathers averaging about 40 mm in length, bilaterally symmetrical white auricular plumes on the sides of the head behind the eyes, averaging about 30 mm in length, and brightly coloured semi-circular rictal plates at the corners of the bill. As in other putative sexually selected traits, auklet ornaments were more variable across individuals than non-ornamental traits. Crest length and auricular plume length were positively correlated within individuals but not across years. Among the traits measured there was evidence for slight sexual dimorphism for the auricular plume and rictal plate ornaments and for culmen length and tarsus (males were slightly larger than females) but not for the crest ornament. Breeding adult females and males had greater crest and plume ornament expression than non-breeding adults. Paradoxically, females' crests and rictal plates were more variable than males' crests and rictal plates. Based on independent samples, the expression of feather ornaments and rictal plate varied among years between 1990 and 1998. Crested Auklet ornaments did not vary in concert with the ornaments of Whiskered Aethia pygmaea and Least Auklets Aethia pusilla during this period. Crested Auklet subadults had smaller ornaments than adults. Based on adults remeasured after an interval of one to seven years, the size of individuals' feather ornaments increased with age. We found no relationship between auricular plume length and asymmetry. Male auricular plumes and female crests were weakly correlated with body condition.     

Carotenoid and melanin-based ornaments signal similar aspects of male quality in two populations of the common yellowthroat

1.  Female preferences for particular male ornaments may shift between populations as a consequence of ecological differences that change the reliability and detectability of the ornament, but few studies have examined how ornaments function in different populations.
2.  We examined the signalling function of male plumage ornaments in a warbler, the common yellowthroat ( Geothlypis trichas ), breeding in New York (NY) and Wisconsin (WI), USA. Males have two prominent ornaments: a black facial mask pigmented with melanin and a yellow bib pigmented by carotenoids. Previous studies in WI indicate that the size of the mask, and not the bib, is primarily related to female choice and male reproductive success. In NY, however, the pattern is reversed and attributes of the bib (size and colour), and not the mask, are the target of sexual selection.
3.  We found that brightness of the yellow bib was the best signal of humoral immunity (immunoglobulin G) in NY and mask size was the best signal in WI, after controlling for breeding experience and capture date. Thus, similar aspects of male quality appeared to be signalled by different ornaments in different populations.
4.  There was no difference between populations in the level of plasma carotenoids or the prevalence of malarial parasites, which may affect the costs and benefits of choosing males with particular ornaments in each location.
5.  Even though females in different populations prefer different ornaments produced by different types of pigments, these ornaments appear to be signalling similar aspects of male quality. Our results caution against inferring the function of particular ornaments based simply on their type of pigment.     

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.     

Why are female birds ornamented?

Sexual selection is now widely accepted as the main evolutionary explanation of extravagant male ornaments. By contrast, ornaments occurring in females have received little attention and often have been considered as nonadaptive, correlated effects of selection on males. However, recent comparative evidence suggests that female ornaments have evolved quite independently of male showiness. Also, new theoretical models predict that both male mate choice and female contest competition will occur under certain circumstances. This is supported by recent experimental studies. Thus, selection acting on females might be a widespread cause of female ornaments.     

Gravettian ivory ornaments in Central Europe,Moravia (Czech Republic)

In the sense of the gravettian technocomplex of Central Europe in Moravia it is mammoth ivory which was mainly used to manufacture and create personal ornaments. The hunters and gatherers selected and stored mammoth tusks as the raw material for the manufacture of tools, weapons, furniture and ornaments. In Moravia, personal ornaments were found in a great variability of forms and sizes – mostly beads like breast-shaped beads, bilobated beads, bilobated flat beads, tear drop shaped beads and round ornaments – rings, bracelets, discs as well as flat elements – diadems and other pendants (zoomorphs, antropomorphs, geometric forms). Some types of personal ornaments show equivalent forms at other gravettian sites and some are unique in Moravia. This article illustrates the variability of these personal ornaments in the context of the Gravettian in Moravia.     

SIGNALING EFFICACY DRIVES THE EVOLUTION OF LARGER SEXUAL ORNAMENTS BY SEXUAL SELECTION

Why are there so few small secondary sexual characters? Theoretical models predict that sexual selection should lead to reduction as often as exaggeration, and yet we mainly associate secondary sexual ornaments with exaggerated features such as the peacock's tail. We review the literature on mate choice experiments for evidence of reduced sexual traits. This shows that reduced ornamentation is effectively impossible in certain types of ornamental traits (behavioral, pheromonal, or color‐based traits, and morphological ornaments for which the natural selection optimum is no trait), but that there are many examples of morphological traits that would permit reduction. Yet small sexual traits are very rarely seen. We analyze a simple mathematical model of Fisher's runaway process (the null model for sexual selection). Our analysis shows that the imbalance cannot be wholly explained by larger ornaments being less costly than smaller ornaments, nor by preferences for larger ornaments being less costly than preferences for smaller ornaments. Instead, we suggest that asymmetry in signaling efficacy limits runaway to trait exaggeration.     

Experimental evidence that female ornamentation increases the acquisition of sperm and signals fecundity

Mate choice can lead to the evolution of sexual ornamentation. This idea rests on the assumption that individuals with more elaborate ornaments than competitors have higher reproductive success due to gaining greater control over mating decisions and resources provided by partners. Nevertheless, how the resources and quality of sexual partners that individuals gain access to are influenced by the ornamentation of rival individuals remains unclear. By experimentally concealing and subsequently revealing female ornaments to males, we confirm in the fowl, Gallus gallus, that female ornamentation influences male mating decisions. We further show, by manipulating the relative ornament size of females, that when females had larger ornaments than competitors they were more often preferred by males and obtained more sperm, especially from higher quality males, as measured by social status. Males may benefit by investing more sperm in females with larger ornaments as they were in better condition and produced heavier eggs. Female ornament size also decreased during incubation, providing a cue for males to avoid sexually unreceptive females. This study reveals how inter-sexual selection can lead to the evolution of female ornaments and highlights how the reproductive benefits gained from mate choice and bearing ornaments can be dependent upon social context.     

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.     

The allometric pattern of sexually size dimorphic feather ornaments and factors affecting allometry

The static allometry of secondary sexual characters is currently subject to debate. While some studies suggest an almost universal positive allometry for such traits, but isometry or negative allometry for nonornamental traits, other studies maintain that any kind of allometric pattern is possible. Therefore, we investigated the allometry of sexually size dimorphic feather ornaments in 67 species of birds. We also studied the allometry of female feathers homologous to male ornaments (female ornaments in the following) and ordinary nonsexual traits. Allometries were estimated as reduced major axis slopes of trait length on tarsus length. Ornamental feathers showed positive allometric slopes in both sexes, although that was not a peculiarity for ornamental feathers, because nonsexual tail feathers also showed positive allometry. Migration distance (in males) and relative size of the tail ornament (in females) tended to be negatively related to the allometric slope of tail feather ornaments, although these results were not conclusive. Finally, we found an association between mating system and allometry of tail feather ornaments, with species with more intense sexual selection showing a smaller degree of allometry of tail ornaments. This study is consistent with theoretical models that predict no specific kind of allometric pattern for sexual and nonsexual characters.     

A novel test of the phenotype-linked fertility hypothesis reveals independent components of fertility

The phenotype-linked fertility hypothesis predicts that male sexual ornaments signal fertilizing efficiency and that the coevolution of male ornaments and female preference for such ornaments is driven by female pursuit of fertility benefits. In addition, directional testicular asymmetry frequently observed in birds has been suggested to reflect fertilizing efficiency and to covary with ornament expression. However, the idea of a phenotypic relationship between male ornaments and fertilizing efficiency is often tested in populations where environmental effects mask the underlying genetic associations between ornaments and fertilizing efficiency implied by this idea. Here, we adopt a novel design, which increases genetic diversity through the crossing of two divergent populations while controlling for environmental effects, to test: (i) the phenotypic relationship between male ornaments and both, gonadal (testicular mass) and gametic (sperm quality) components of fertilizing efficiency; and (ii) the extent to which these components are phenotypically integrated in the fowl, Gallus gallus. We show that consistent with theory, the testosterone-dependent expression of a male ornament, the comb, predicted testicular mass. However, despite their functional inter-dependence, testicular mass and sperm quality were not phenotypically integrated. Consistent with this result, males of one parental population invested more in testicular and comb mass, whereas males of the other parental population had higher sperm quality. We found no evidence that directional testicular asymmetry covaried with ornament expression. These results shed new light on the evolutionary relationship between male fertilizing efficiency and ornaments. Although testosterone-dependent ornaments may covary with testicular mass and thus reflect sperm production rate, the lack of phenotypic integration between gonadal and gametic traits reveals that the expression of an ornament is unlikely to reflect the overall fertilizing efficiency of a male.     

山西柿子滩旧石器遗址蚌饰品制作工艺研究

本文是对柿子滩遗址蚌饰品制作工艺的研究。该遗址分布于山西省吉县清水河流域, 是一处含有细石器遗存的旧石器时代晚期遗址, 年代为25000BP—10000BP。蚌质的出土遗物17件, 发现于不同地点的不同层位中,其中有穿孔的5件, 有磨制痕迹的3件, 分别发现于S9、S12A和S29三个地点。研究表明, 蚌饰品穿孔有"钻孔"和"磨孔"两种形式, 存在三种制作程序, 早期到晚期的制作和穿孔技术表现出越来越进步的特点。     

The direction of mothers' and daughters' preferences and the heritability of male ornaments in red jungle fowl (Gallus gallus)

We used a breeding design involving 18 sires and 108 dams tostudy the heritabilities of male ornaments in red jungle fowl(Gallus gallus). Ornaments used by females to choose mates showedlow heritabilities, with the exception of comb and wattle measures.The general absence of heritability suggests that a geneticcovariance did not exist at the time of this study between mostmale ornaments and female preferences for those ornaments. Thisresult is contrary to a key prediction of the arbitrary or Fisherianhypothesis of sexual selection. Comb size and color are condition-dependenttraits that reflect short-term changes in health, and comb sizeof males was positively correlated with offspring weight. Ourresults are consistent with the expectation of good-genes hypothesesthat male ornaments reflect the ability of males to withstandenvironmental stresses.     

A test of the Darwin–Fisher theory for the evolution of male secondary sexual traits in monogamous birds

The Darwin–Fisher theory proposes that the presence of male secondary sexual traits in monogamous birds is selected for by early season breeding of females that are in good condition. These early breeding females have high fecundity because of their good condition, and they select mates based on secondary sex traits. We tested whether this hypothesis may be responsible for the presence of male sexual ornaments in the great frigatebird, a socially and genetically monogamous seabird. Consistent with the Darwin–Fisher theory, we found that reproductive success declined over the season. However, males with more exaggerated ornaments were not chosen as mates earlier in the season than males with less exaggerated ornaments, and selection gradients on these ornaments were not significantly different from zero.     

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-limited expression 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 related to 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 monogamous species. 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.     

Sexual Ornaments,Body Morphology,and Swimming Performance in Naturally Hybridizing Swordtails (Teleostei: Xiphophorus)

Determining the costs of sexual ornaments is complicated by the fact that ornaments are often integrated with other, non-sexual traits, making it difficult to dissect the effect of ornaments independent of other aspects of the phenotype. Hybridization can produce reduced phenotypic integration, allowing one to evaluate performance across a broad range of multivariate trait values. Here we assess the relationship between morphology and performance in the swordtails Xiphophorus malinche and X. birchmanni, two naturally-hybridizing fish species that differ extensively in non-sexual as well as sexual traits. We took advantage of novel trait variation in hybrids to determine if sexual ornaments incur a cost in terms of locomotor ability. For both fast-start and endurance swimming, hybrids performed at least as well as the two parental species. The sexually-dimorphic sword did not impair swimming performance per se. Rather, the sword negatively affected performance only when paired with a sub-optimal body shape. Studies seeking to quantify the costs of ornaments should consider that covariance with non-sexual traits may create the spurious appearance of costs.     

Sexual dimorphism, extrapair fertilizations, and operational sex ratio in great frigatebirds (Fregata minor)

Across taxa, the presence of sexual ornaments in one sex isusually correlated with disproportionately great parental effortby the other. Frigatebirds (Fregatidae) are sexually dimorphic,with males exhibiting morphological and behavioral ornaments,but males and females share in all aspects of parental effort.All other taxa in a clade of 237 species exhibit biparentalcare, but only frigatebirds exhibit pronounced sexual dimorphism. We tested for the presence of two factors that could contributeto the evolution of male ornaments in great frigatebirds: ahigh frequency of extrapair fertilizations and a male-biasedoperational sex ratio. In 92 families sampled over two breedingseasons, there was only one extrapair fertilization. However,in both seasons, there were more males than females availablefor mating, and the sex ratio among individuals actively engagedin mate-acquisition behavior was strongly male biased, withtypically five or six males available per female. Our resultssuggest that extrapair fertilizations are not responsible forthe exaggeration of sexual ornaments in male frigatebirds,and that operational sex ratio may be related to sexual dimorphismin this species. Further work is needed to determine whetherthe male-biased operational sex ratio creates the variancein male reproductive success that would be needed to drivethe evolution of male ornaments.     

Variable sexual ornaments in scarlet-tufted malachite sunbirds (Nectarinia johnstoni) on Mount Kenya

In order to be elaborated by sexual selection, sexual ornaments must vary perceptibly and genetically among individuals in natural populations. Rather little is known about ornament variation in monogamous species, in which sexual selection should act more weakly than in polygynous species. We report phenotypic variation in feather ornament size (elongated tails and pectoral tufts) and body size in the scarlet-tufted malachite sunbird Nectarinia johnstoni , a monogamous, sexually dimorphic nectarivore of East African alpine zones. Fully-expressed male ornaments are highly significantly more variable (CVs = 12–29%) than are skeletal and wing measures primarily affected by natural selection (CVs = 2 4%). Female sunbirds have pectoral tufts which are significantly (22–25%) smaller than those of adult males, but more variable (CV_s= 21–22%, CV_s= 12–15%), and more variable than body size. Among males with fully-grown ornaments, those with longer tails tend to have longer wings and wider tufts. The high variation in fully-grown ornaments in malachite sunbirds is consistent with the view that the ornaments are condition-dependent sexual signals. Finally, we review studies of feather ornament variation to date, and show that ornaments are much more variable in monogamous than non-monogamous species, apparently due to the relatively weak pressure of sexual selection.