Extreme ultraviolet sexual dimorphism in jumping spiders (Araneae: Salticidae)

Jumping spiders (Salticidae) have acute vision with some cells in the retina that are sensitive to ultraviolet (UV) spectra (< 400 nm). However, no study has documented the use of UV signals in salticids. To appreciate the function of UV vision, it is necessary to characterize the UV colours of salticids. In the present study, the UV and human-visible wavelengths of a tropical ornate salticid spider, Cosmophasis umbratica , were analysed using reflectance spectrometry to obtain evidence of sex-specific UV colours. An absolute sexual dimorphism in the UV colours of this salticid species was found. All of the body parts of adult males that are displayed to conspecifics during intra-specific interactions reflected UV (300–400 nm) light, whereas the adult females and juveniles did not reflect UV light from any body part. A great deal of variation was also found in the UV wavebands among males. This is the first full UV characterization of a salticid spider and the first study to demonstrate an extreme sexual UV dimorphism in jumping spiders. The findings obtained provide evidence that UV reflectance may comprise important sexual signals in jumping spiders.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 89 , 397–406.     

Ultraviolet reflectance and cryptic sexual dichromatism in the ocellated lizard, Lacerta (Timon) lepida (Squamata: Lacertidae)

Ultraviolet (UV) colorations have garnered extensive theoretical and empirical treatment in recent years, although the majority of studies have concerned themselves with avian taxa. However, many lizards have acute visual systems with retinal photoreceptors that are sensitive to UV wavelengths, and also display UV-reflecting colour patches. In the present study, we used UV photography and full-spectrum reflectance spectrophotometry to describe intra- and intersexual colour variation in adult ocellated lizards Lacerta ( Timon ) lepida and to obtain evidence of UV-based ornamentation. We also investigated whether any colour traits correlate with morphological traits potentially related to individual quality. The results obtained show that the prominent eyespots and blue outer ventral scales (OVS) that ocellated lizards have on their flanks reflect strongly in the UV range and are best described as UV/blue in coloration. The eyespots of males are larger and cover a larger surface area than those of females. However, these differences can be entirely accounted for by sex differences in body size, with males being generally larger than females. We also found differences in the shape of reflectance curves from males and females, with the eyespots and blue OVS of males being more UV-shifted than those of females. Other body regions have extremely low UV reflectance and are not sexually dichromatic. Eyespot size and the total surface area covered by eyespots increases with body size in males but not in females, suggesting that they may be signalling an intrinsic individual characteristic such as body size or male fighting ability. We also discuss the alternative and non-exclusive hypothesis that eyespots may function in lizards of both sexes as protective markings against predators.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 766–780.     

Heterochrony and its role in sex determination of cryptically dioecious Consolea (Cactaceae) staminate flowers

Ovule development, megasporogenesis, and megagametogenesis were studied in six cryptically dioecious species of Consolea. All species showed uniform development typical for the Opuntioideae. Ovule development proceeds acropetally, but shows developmental asynchrony across floral morphs. At anthesis, female morph ovules are functional and available for fertilization, whereas staminate flower ovules are senescing and incapable of being fertilized. In occasional plants of some species, staminate flowers may reach anthesis with a few functional apical ovules capable of seed formation. Such plants are described as inconstant/leaky males. Ovule fertility differences across morphs are interpreted as resulting from heterochronic ovule development and senescence, although variation in embryo sac longevity cannot be ruled out. Significantly, ovule abortion follows a common pattern and timing in staminate flowers of both male morphs in all species. Thus, on the basis of this uniformity, a common origin for the cryptically dioecious breeding system in Consolea is hypothesized. Furthermore, staminate expression in Consolea appears to be controlled by a common, genetically determined heterochronic ovule developmental programme affecting the relative timing of ovule receptivity and flower opening. This is the first report of heterochrony as a mechanism of male sex determination.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 156 , 305–326.     

Variation in ultraviolet reflectance by carotenoid-bearing feathers of tanagers (Thraupini: Emberizinae: Passeriformes)

Avian visual sensitivity encompasses both the human visible range (400–700 nm) and also near‐ultraviolet (UV) wavelengths (320–400 nm) invisible to normal humans. I used reflectance spectrophotometry to assess variation in UV reflectance for yellow, orange and red plumage in 67 species of tanager (Passeriformes). Previous chemical studies, and my analysis of reflectance minima, suggest that carotenoids are the dominant pigments in yellow, orange and red tanager plumage. Spectra recorded over the range of wavelengths to which birds are sensitive (320–700 nm) were invariably bimodal, with both a plateau of high reflectance at longer (> 500 nm) wavelengths and a distinct secondary peak at UV (< 400 nm) wavelengths. Within this overall framework, variation in UV reflectance was expressed within well‐defined quantitative limits: (1) peak reflectance was always lower than the corresponding plateau of reflectance at longer visible wavelengths; (2) the intensity of peak reflectance declined steadily below 350 nm; (3) wavelengths of peak reflectance clustered between 350 and 370 nm. Significant correlations were detected between various measures of total reflectance in the UV and visible wavebands, but not between various measures of spectral location of UV and visible reflectance. I propose that the strong absorption band at short visible wavelengths (～ 380–550 nm) responsible for bimodal spectra of carotenoids in vitro is also responsible for bimodal reflectance by carotenoid‐based plumage colours. The construction of the UV and visible reflectance bands from different sides of this same absorbance band provides a mechanism for the observed covariation between UV and visible wavelengths. Lack of an association between the spectral locations of the UV and visible reflectance bands may result from the limited variation in spectral location of the UV band. These patterns suggest that plumage colours are subject to constraints, just as are more traditional morphological characters. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 84 , 243–257.     

Inter- and intrasexual dimorphism in the diving beetle Hydroporus memnonius Nicolai (Coleoptera: Dytiscidae)

Sexual conflict can drive rapid intersexual arms races, and lead to pronounced sexual dimorphism. Such dimorphism is frequent in diving beetles, where males typically possess expanded front and middle tarsi, supplied with adhesive setae to grasp females during mating, and females often have rough dorsal surfaces which hinder male attachment. In a number of species, females are dimorphic, being either smooth and male-like, or heavily sculptured dorsally. Smooth and sculptured females often have distinct biogeographies, and may be expected to be associated with specific counter-adaptations in males. The European diving beetle, Hydroporus memnonius Nicolai, includes a smooth male-like female, and a matt morph, var. castaneus Aubé, which are largely allopatric in distribution. We show that the two morphs differ in the density and intensity of their surface microreticulation, and that matt females are associated with morphologically distinct males, which have developed specific countermeasures on their tarsi, including a greater number of large adhesive setae, individually larger in area. Such males are expected to be more successful in pairing with both matt and shining females, and it is suggested that a process of population replacement, partly driven by sexual interactions, may occur where the two forms overlap in range.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 685–697.     

Sympatric occurrence of diverged mtDNA lineages of Pityogenes chalcographus (Coleoptera, Scolytinae) in Europe

Differentiation among the European populations of the six-toothed spruce bark beetle Pityogenes chalcographus was observed for the first time in the 1970s as mating males from Northern Europe with females from Central Europe led to a significant decline in fecundity. Morphological examination revealed that P. chalcographus can be separated into two European races. Here, we investigated the genetic background of this separation by analysing 39 populations ( n  = 695), sequencing almost the complete cytochrome oxidase I gene (1543 bp) and applying single-strand conformation polymorphism. Phylogenetic analysis of 58 haplotypes yielded three major clades with a maximum sequence divergence of 2.33%, indicating that the demographic events took place in the late Pleistocene. These results support the hypothesis of allopatric divergence of the mtDNA lineages, which postglacially came into sympatric existence in Europe. However, as a result of partial crossing incompatibility the diverged lineages retained their genetic identity during postglacial times.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 331–340.     

It is all in the head: morphological basis for differences in bite force among colour morphs of the Dalmatian wall lizard

Males of the lizard Podarcis melisellensis occur in three distinct colours that differ in bite performance, with orange males biting harder than white or yellow ones. Differences in bite force among colour morphs are best explained by differences in head height, suggesting underlying variation in cranial shape and/or the size of the jaw adductors. To explore this issue further, we examined variation in cranial shape, using geometric morphometric techniques. Additionally, we quantified differences in jaw adductor muscle mass. No significant differences in size corrected head shape were found, although some shape trends could be detected between the colour morphs. Orange males have relatively larger jaw adductors than yellow males. Not only the mass of the external jaw adductors, but also that of the internal jaw adductors was greater for the orange morph. Data for other cranial muscles not related to biting suggest that this is not the consequence of an overall increase in robustness in orange individuals. These results suggest that differences in bite performance among morphs are caused specifically by an increase in the mass of the jaw adductor, which may be induced by differences in circulating hormone levels.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 13–22.     

Ultraviolet and carotenoid‐based coloration in the viviparous lizard Zootoca vivipara (Squamata: Lacertidae) in relation to age,sex, and morphology

Lizards display structural and pigment‐based colorations, and their visual system is sensitive to wavelengths of 300–700 nm. However, few studies in squamate reptiles have quantified interindividual colour variation that includes the structural ultraviolet (UV) component (300–400 nm). In the present study, we investigated variability of a ventral UV/yellow–red ornamentation in the common lizard Zootoca vivipara, including an analysis of spatial distribution, as well as sex and age differences. We also investigated whether the expression of coloration is related to body size and condition. Our analyses revealed two distinct patches: a gular patch with a strong UV reflectance and a belly patch with a dominant yellow–red reflectance. Males displayed a less saturated throat coloration with higher UV chroma and UV hue, and had a redder but duller belly coloration than females. Yearlings had less elaborate ornaments than adults, although they already displayed a yellow–red sexual dichromatism on the belly. UV sexual dichromatism was only apparent in adults as a result of a weaker UV reflectance in females, suggesting potential fitness costs of a bright UV coloration in that sex. Different colour traits were related to body size in both sexes, as well as to body condition in males. We discuss the potential evolutionary scenarios leading to the maintenance of this ornament in common lizards. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 128–141.     

Enhanced colour polymorphisms in island populations of the land snail Euhadra peliomphala

Variations in visible genetic polymorphisms are assumed to decrease in populations on small islands because of intense founder effects, genetic drift and inbreeding. However, we have found evidence of a marked enhancement of colour polymorphisms within populations on small oceanic islands that were colonized from the mainland. The source populations on the mainland of the land snail Euhadra peliomphala in four oceanic islands were estimated by phylogenetic analysis of mitochondrial DNA sequences. Diversity of shell colour was higher in the island populations than in the source populations on the mainland. In addition, the shell colour morphs differed not only among populations from different islands but also between the island populations and the source populations on the mainland. By contrast, no mtDNA variations were found in any of the island populations, even though the source populations possessed high mtDNA diversity. Thus, components of colour morphs changed in the island populations after their colonization, and colour polymorphisms are enhanced in these islands despite the loss of genetic variation. The above findings suggest that ecological mechanisms such as morphological release owing to a release from competition may overcome the tendency toward reduced genetic variation in islands to enhance the colour polymorphism.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 81 , 417–425.     

Seasonal changes in a ultraviolet structural colour signal in blue tits, Parus caeruleus

Recent studies of blue tits, Parus caeruleus , have found sexual selection and a viability-indicating function of the structural ultraviolet and blue crown plumage, but the reasons for this signal variation are not understood. Furthermore, studies in England and Sweden have yielded somewhat different results (particularly with regard to the spectral position of the reflectance peak). Here we investigate whether the blue tit UV/blue ornament varies with time of year since such variation might be relevant to the signalling function as well as the apparent difference between populations. From 400 blue tits captured at two different localities in Sweden, we found that objective measures of 'hue' (spectral location), 'chroma' (spectral purity) and 'brightness' (spectral intensity), varied substantially with season. Just after moult (October), crown 'hue' is maximally UV-shifted (359 nm for males and 373 nm for females). Thereafter the peak drifts upwards and by the time of nestling feeding (June) male reflectance peaks at 404 nm and female at 413 nm. This change is probably due to feather wear as well as fat and dirt accumulation, which might constitute an additional male quality cue. Our results suggest that it is important to consider plumage age when exploring variation in structural plumage coloration, and that it can largely explain the difference between the British and Swedish studies. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 76 , 237–245.     

Iridescent ultraviolet signal in the orange sulphur butterfly (Colias eurytheme): spatial, temporal and spectral properties

Many of nature's most striking animal colours are iridescent, exhibiting a high degree of spectral purity and strong angular dependence of intensity and hue. Although a growing number of studies have detailed the intricate mechanisms responsible for producing iridescent colours, few attempts have been made to describe their dynamic appearance in ecologically and behaviourally realistic contexts. We suggest that the optical properties unique to iridescent structural colours are important for understanding how they function as signals during behavioural interactions. Using males of the orange sulphur butterfly, Colias eurytheme , which exhibit an iridescent ultraviolet (UV) reflectance on their dorsal wing surfaces, we develop a holistic framework for inferring the appearance of this signal to conspecifics under field conditions that incorporate data on their spectral sensitivity. We show that, during flight, the UV signal is brightest within a wing beat cycle when viewed from directly above the male. Spectral properties of the signal under natural lighting indicate that male wing colour should be readily perceived and distinguished from that of females and from the dark green visual background of UV-absorbing vegetation. Finally, our analyses permit predictions regarding how signal senders and receivers should orientate themselves for maximal transmission and reception of this ultraviolet iridescent signal.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 349–364.     

Temporal shift of diet in alternative cannibalistic morphs of the tiger salamander

Evolutionary theory predicts that alternative trophic morphologies are adaptive because they allow a broad use of resources in heterogeneous environments. The development of a cannibal morphology is expected to result in cannibalism and high individual fitness, but conflicting results show that the situation is more complex. The goal of the present study was to increase our understanding of the ultimate benefits of a cannibalistic polyphenism by determining temporal changes in the feeding habits and biomass intake in a population of tiger salamanders ( Ambystoma tigrinum nebulosum ). Cannibals in this species develop a larger head than typicals and have prominent teeth, both useful for consuming large prey. Although cannibalism was only detected in cannibal morphs, large temporal variation in resource partitioning was found between morphs. The two morphs always differed in their foraging habits, but cannibalism mainly occurred immediately after the ontogenetic divergence between morphs. Cannibals shifted their foraging later to a more planktivorous diet (i.e. the primarily prey of the typical morph). Cannibals also obtained more prey biomass than typicals. These results indicate that the cannibalistic morph is advantageous over the typical development, but that these advantages vary ontogenetically. Although the results obtained are consistent with models predicting the maintenance of cannibalism polyphenism in natural populations, they show that the foraging tactics utilized by cannibal morphs, and the fitness consequences accrued by such tactics, are likely to be more complex and dynamic than previous studies have suggested.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 89 , 373–382.     

Testing the role of coadapted genes versus bet-hedging for mating strategies in colour polymorphic pygmy grasshoppers

Recent investigations of mate choice indicate that the genetic effect of sires on offspring fitness may depend on the interaction between maternal and paternal genotypes and the environmental conditions experienced by the offspring. Alternative colour morphs of the pygmy grasshopper, Tetrix subulata , represent ecological strategies that differ in body size, life history, thermoregulatory behaviour, and habitat selection. The hypothesis that selection promotes behaviours maintaining coadapted gene complexes predicts individuals to mate assortatively with respect to colour morph. On the other hand, the bet-hedging hypothesis predicts that the temporal variability of the environment inhabited by these animals may select for disassortative mating behaviour resulting in heterogeneous offspring. To distinguish between these competing hypotheses, we investigated mating behaviours using dual-choice experiments. Our results were not in agreement with the prediction of assortative mating but suggest instead that matings were random with regard to colour morph. Polyandry was common, and females mated with the second male regardless of whether the first mating was assortative or disassortative. Polyandry also was equally frequent among females in triads in which the two males belonged to different colour morphs as in triads where both males belonged to the same colour morph. A field experiment confirmed that polyandry occurred also among free-ranging individuals, and uncovered variation in mating success among male colour morphs, probably due to indirect effects of coloration on activity or habitat use. The consequences of this random and polyandrous mating strategy for the evolutionary dynamics of the colour polymorphism remain to be explored.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 491–499.     

A new species of Betula section Betulaster (Betulaceae) from China

A new species of Betulaceae, Betula fujianensis , is described and illustrated from Luoboyan Reserve, Fujian Province, south-eastern China. Betula fujianensis is distributed in subtropical evergreen and deciduous broad-leaved mixed forest at elevations above 500 m, mostly as scattered individuals and occasionally as populations alone or with B. luminifera . Betula fujianensis belongs to section Betulaster and differs from other species in general morphology and phenology. A key to all species currently recognized in this section is presented, and the relationships between the new species and its closely related taxa are discussed.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 156 , 523–528.     

Why cross the web: decoration spectral properties and prey capture in an orb spider (Argiope keyserlingi) web

An effective visual signal elicits a strong receiver response. The visual receptors of most insects are sensitive to ultraviolet (UV), blue and green light. The decorations of certain orb web spiders may be described as a type of visual signal, a sensory trap, as they exploit visual biases in insects. We filtered UV and blue light from the decorations of Argiope keyserlingi , under field conditions, using plastic sheets to test if the UV and blue light reflected affects the type of prey caught. We found that houseflies, blowflies, stingless bees, honeybees and vespid wasps were caught more frequently in webs with decorations than webs without, while ichneumonid wasps were caught less frequently. Blowflies, stingless bees, honeybees and vespid wasps were caught more often in unfiltered decorated webs. These insects also have receptor sensitivities in the blue and UV. We showed that exploiting visual sensory biases plays an integral role in attracting insects to orb web decorations. Whether UV light, blue light, or both, are the most important cue, however, requires further study.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 221–229.     

Reproductive ecology of 21 coexisting Psychotria species (Rubiaceae): when is heterostyly lost?

Heterostyly is a genetic polymorphism in which plant populations are composed of two or more morphs that differ in stigma and anther heights. The polymorphism promotes intermorph pollen transfer, thus outcrossing. Heterostyly has been reported in 28 angiosperm families and is frequently lost in heterostylous lineages. To assess ecological factors related to shifts from heterostyly to monomorphy, we examined the reproductive ecology of heterostylous tropical shrubs of Psychotria . Among 21 species at Barro Colorado Island and the nearby Parque Nacional Soberania, Panama, 14 species were heterostylous while seven were monomorphic. A molecular phylogeny and the existence elsewhere of heterostylous populations indicated that the breakdown of heterostyly had occurred independently. Heterostylous and monomorphic species were visited by the same bee species, although visit frequencies were lower in monomorphic species. Monomorphic species had significantly lower population density and greater fruit set than did heterostylous species. Autonomous autogamy made a large contribution to fruit set in monomorphic species and was only rarely observed in heterostylous species. The results indicate monomorphic and heterostylous species produce more seeds through selfing and outcrossing, respectively. The limitation of outcrossing as a result of low population density may be related to the breakdown of heterostyly and the evolution of selfing.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 125–134.     

Head shape allometry and proximate causes of head sexual dimorphism in Podarcis lizards: joining linear and geometric morphometrics

Podarcis bocagei and Podarcis carbonelli are two lacertid species endemic to the western Iberian Peninsula, and both show head size and shape sexual dimorphism. We studied immature and adult head sexual dimorphism and analysed ontogenetic trajectories of head traits with body and head size, aiming to shed light on the proximate mechanisms involved. Immatures were much less dimorphic than adults, but geometric morphometric techniques revealed that head shape sexual differences are already present at this stage. Males and females differed in allometry of all head characters with body size, with males showing a disproportionate increase of head size and dimensions. On the other hand, head dimensions and head shape changed with increasing head size following similar trends in both sexes, possibly indicating developmental restrictions. Consequently, adult sexual dimorphism for head characters in these species is the result of both shape differences in the immature stage and hypermetric growth of the head in relation to body size in males.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 111–124.     

Females show greater changes in wing colour with latitude than males in the green‐veined white butterfly,Pieris napi (Lepidoptera: Pieridae)

In butterflies, wing colour may simultaneously be under sexual selection in the context of mating selection and natural selection in the context of thermoregulation. In the present study, we collected mated females of the green‐veined white butterfly (Pieris napi) from locations spanning 960 km of latitude across Fennoscandia, and investigated sex‐specific latitudinal wing colour variation in their offspring raised under identical conditions. We measured wing colour characteristics, including reflectance at wavelengths 300–700 nm and the degree of wing melanization. At all latitudes, females reflected more light in the short wavelengths (< 400 nm) and less in the long wavelengths (> 450 nm), and they were more melanized than males. However, female wing colour varied more with latitude than that of males. Among females, long wavelength reflectance decreased, whereas short wavelength reflectance and melanization increased, towards the north. By contrast, among males, latitudinal variation was found only in the ventral hindwing melanization. These results are consistent with the idea that the balance between natural and sexual selection acting on wing colour changes with latitude differently in males than females. The dark wing colour of females in the north may be a thermoregulatory adaptation, although males may be constrained from evolving the dark dorsal wing colour favoured by natural selection because of constant sexual selection across latitudes. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ?? , ??–??.