Freezing behaviour facilitates bioelectric crypsis in cuttlefish faced with predation risk.

Cephalopods, and in particular the cuttlefish Sepia officinalis, are common models for studies of camouflage and predator avoidance behaviour. Preventing detection by predators is especially important to this group of animals, most of which are soft-bodied, lack physical defences, and are subject to both visually and non-visually mediated detection. Here, we report a novel cryptic mechanism in S. officinalis in which bioelectric cues are reduced via a behavioural freeze response to a predator stimulus. The reduction of bioelectric fields created by the freeze-simulating stimulus resulted in a possible decrease in shark predation risk by reducing detectability. The freeze response may also facilitate other non-visual cryptic mechanisms to lower predation risk from a wide range of predator types.

Fruit and Seed Volatiles: Multiple Stage Settings, Actors and Props in an Evolutionary Play

Plants emit volatile organic compounds (VOCs) from most parts of their anatomy. Conventionally, the volatiles of leaves, flowers, fruits and seeds have been investigated separately. This review presents an integrated perspective of volatiles produced by fruits and seeds in the context of selection on the whole plant. It suggests that fruit and seed volatiles may only be understood in the light of the chemistry of the whole plant. Fleshy fruit may be viewed as an ecological arena within which several evolutionary games are being played involving fruit VOCs. Fruit odour and colour may be correlated and interact via multimodal signalling in influencing visits by frugivores. The hypothesis of volatile crypsis in the evolution of hard seeds as protection against volatile diffusion and perception by seed predators is reviewed. Current views on the role of volatiles in ant dispersal of seeds or myrmecochory are summarised, especially the suggestion that ants are being manipulated by plants in the form of a sensory trap while providing this service. Plant VOC production is presented as an emergent phenotype that could result from multiple selection pressures acting on various plant parts; the ``plant'' phenotype and VOC profile may receive significant contributions from symbionts within the plant. Viewing the plant as a holobiont would benefit an understanding of the emergent plant phenotype.

Estimating the abundance of clustered and cryptic marine macro-invertebrates in the Galápagos with particular reference to sea cucumbers

Estimating the abundance of marine macro-invertebrates is complicated by a variety of factors: 1) human factors, such as diver efficiency and diver error; and 2) biological factors, such as aggregation of organisms, crypsis, and nocturnal emergence behavior. Diver efficiency varied according to the detectability of an organism causing under-estimation of density by up to 50% in some species. All common species were aggregated at scales from 10-50 m. Transects need to be long enough to transcend the scale of patchiness to improve accuracy. Some species of sea urchins and sea cucumbers (pepinos) which are cryptic by day emerged at night so that daytime censuses underestimated their abundance by up to 10 times. In the sea cucumber fishery, estimates of abundance need to be made at the scale of the population, i.e. at hundreds of km. A strategy for this is proposed.

Gorgonians of the South of Portugal: biology, ecology and conservation

O papel ecológico das gorgónias (Octocorallia: Alcyonacea) nos fundos marinhos rochosos é mundialmente reconhecido. Contudo, a informação acerca da ecologia e biologia das espécies de gorgónias nas zonas temperadas do NE Atlântico é manifestamente escassa, especialmente tendo em consideração as actuais perturbações globais, regionais e locais. Nos fundos rochosos da costa algarvia até aos 30 m, verificouse que várias espécies de gorgónias são abundantes e frequentes, nomeadamente Eunicella labiata, Eunicella gazella, Eunicella verrucosa, Leptogorgia lusitanica e Leptogorgia sarmentosa. As populações de gorgónias são co-dominadas por diferentes espécies que apresentaram elevados índices de associação, indicando reduzidos níveis de competição entre elas. Em todo o caso, a estrutura dos povoamentos diferiu com as condições locais. Todas as espécies evidenciaram padrões de distribuição semelhantes ao longo do gradiente de profundidade, i.e. a abundância aumenta significamente com a profundidade após os 15 m. A profundidades mais baixas (até aos 15 m), a distribuição das gorgónias parece ser condicionada por factores abióticos e pela competição com algas. Com efeito, os padrões de distribuição espacial das espécies de gorgónias na costa algarvia são determinados pela interacção de pressões naturais e antropogénicas (ex. pesca). Ainda que as colónias de maior tamanho não tenham sido restritas a áreas menos pescadas, em áreas mais perturbadas pela pesca, a distribuição dos tamanhos das colónias estava maioritariamente desviada para tamanhos mais pequenos. Os efeitos das perturbações naturais nas populações de gorgónias foram evidenciados pela ocorrência de padrões demográficos distintos em áreas vizinhas sujeitas a níveis semelhantes de pressões antropogénicas. Estes estudos demonstraram, ainda, que os efeitos na distribuição de frequências de tamanho das colónias são dependentes das espécies de gorgónias em causa: Eunicella labiata não parece ser afectada; Leptogorgia sarmentosa é tendencialmente afectada por pressões antropogénicas; Eunicella gazella e Leptogorgia lusitanica aparentam ser afectadas, quer por pressões naturais, quer por pressões antropogénicas. Os efeitos verificados nos padrões da distribuição de frequências de tamanho, particularmente a tendência para o desvio destas frequências para tamanhos mais pequenos em áreas sujeitas a perturbações, poderão ter consequências para a biodiversidade dos fundos sublitorais rochosos na costa algarvia. Com efeito, o presente estudo apoia o paradigma geral de que os corais são habitats que suportam comunidades de elevada biodiversidade e abundância. Num dos poucos estudos que examinam a relação entre as gorgónias e as suas comunidades de invertebrados epibentónicos, foi verificado que as gorgónias (Eunicella gazella e Leptogorgia lusitanica) sustentam comunidades ricas (11 phyla, 181 taxa) e abundantes (7284 indivíduos). Estas comunidades são dominadas por anfípodes, mas os poliquetas tiveram um grande contributo para os níveis elevados de biodiversidade. Verificou-se, igualmente, que o tamanho da colónia desempenha um papel fundamental na biodiversidade, na medida em que as colónias de menor tamanho apresentaram um contributo mais baixo, comparativamente às médias e grandes. Ainda que ambas as gorgónias partilhem a maioria das espécies amostradas, 11 e 18 taxa foram exclusivos de Eunicella gazella e Leptogorgia lusitanica, respectivamente (excluindo indivíduos com presenças únicas). No entanto, a maioria destes taxa eram ou pouco abundantes ou pouco frequentes. A excepção foi a presença de planárias (Turbellaria) de coloração branca nas colónias de Eunicella gazella, provavelmente beneficiando do efeito de camuflagem proporcionado pelos ramos com a mesma coloração. Com efeito, a complementaridade entre as comunidades epibentónicas associadas a ambas as gorgónias diminuiu quando usados os dados de presença/ausência, sugerindo que os padrões de biodiversidade são mais afectados pelas alterações na abundância relativa das espécies dominantes do que pela composição faunística. As comunidades de epifauna bentónica associadas a estas gorgónias não só apresentaram valores elevados de ®-diversidade, como de ¯- diversidade, resultantes de padrões intrincados de variabilidade na sua composição e estrutura. Ainda que o conjunto de espécies disponíveis para colonização seja, na generalidade, o mesmo para ambos os locais, cada colónia apresenta uma parte deste conjunto. Na sua totalidade, as colónias de gorgónias poderão funcionar como uma metacomunidade, mas a estrutura das comunidades associadas a cada colónia (ex. número total de espécies e abundância) parecem depender dos atributos da colónia, nomeadamente superfície disponível para colonização (altura, largura e área), complexidade e heterogeneidade (dimensão fractal e lacunaridade, respectivamente) e cobertura epibentónica “colonial” (ex. fauna colonial e algas macroscópicas; CEC). Numa primeira tentativa para quantificar a relação entre as gorgónias e os invertebrados epibentónicos a elas associados (em termos de abundância e riqueza específica), verificou-se que a natureza e a intensidade destas relações dependem da espécie hospedeira e variam para os grupos taxonómicos principais. No entanto, independentemente do grupo taxonómico, a riqueza específica e a abundância estão significativamente correlacionadas com a CEC. Com efeito, a CEC provavelmente devido a um efeito trófico (aumento da disponibilidade alimentar directo ou indirecto), combinado com a superfície disponível para colonização (efeito espécies-área) foram as variáveis mais relacionadas com os padrões de abundância e riqueza específica. Por outro lado, ainda que a complexidade estrutural seja frequentemente indicada como um dos factores responsáveis pela elevada diversidade e abundância das comunidades bentónicas associadas a corais, a dimensão fractal e a lacunaridade apenas foram relevantes nas comunidades associadas a Leptogorgia lusitanica. A validade do paradigma que defende que a complexidade estrutural promove a biodiversidade poderá ser, então, dependente da escala a que se realizam os estudos. No caso das gorgónias, o efeito da complexidade ao nível dos agregados de gorgónias poderá ser muito mais relevante do que ao nível da colónia individual, reforçando a importância da sua conservação como um todo, por forma a preservar a diversidade de espécies hospedeiras, o seu tamanho e estrutura. Actividades antropogénicas como a pesca, podem, ainda, ter efeitos negativos ao nível da reprodução de espécies marinhas. Analogamente ao verificado para os padrões de distribuição espacial das populações de gorgónias na costa algarvia, a informação relativa à sua reprodução é igualmente escassa. Os estudos realizados em populações de Eunicella gazella a 16m de profundidade, demonstraram que o desenvolvimento anual das estruturas reprodutivas é altamente sincronizado entre os sexos. A razão entre sexos na população foi de 1.09 (F:M), encontrando-se perto da paridade. A espermatogénese estende-se por 6 a 8 meses, enquanto que a oogénese é mais demorada, levando mais de um ano para que os oócitos se desenvolvam até estarem maduros. Antes da libertação dos gâmetas, foi observada uma elevada fecundidade nas fêmeas (27.30§13.24 oócitos pólipo−1) e nos machos (49.30§31.14 sacos espermáticos pólipo−1). Estes valores encontram-se entre os mais elevados reportados à data para zonas temperadas. A libertação dos gâmetas (não há evidência de desenvolvimento larvar, nem à superfície da colónia, nem no seu interior) occorre em Setembro/ Outubro, após um período de elevada temperatura da água do mar. As fêmeas emitem oócitos maduros de elevadas dimensões, retendo, todavia, os oócitos imaturos que se desenvolvem apenas na época seguinte. Ainda que o efeito da pesca nas populações de gorgónias da costa do Algarve seja perceptível, às taxas actuais, o mergulho recreativo não aparenta afectar seriamente estas populações. Contudo, sendo uma indústria em expansão e conhecendo-se a preferência de mergulhadores por áreas rochosas naturais ricas em espécies bentónicas, futuramente poderá vir a afectar estes habitats. A monitorização de mergulhadores na costa algarvia mostrou que a sua maioria (88.6 %) apresenta comportamentos que podem impactar o habitat, com uma taxa média de contactos de 0.340§0.028 contactos min−1. Esta taxa foi mais elevada em mergulhadores com moderada experiência e na fase inicial do mergulho (0–10 min). Os contactos com as barbatanas e mãos foram comuns, resultando, maioritariamente, na resuspensão do sedimento, mas geralmente apresentando um impacto reduzido. Todavia, a fauna também foi afectada, quer por danos físicos, quer pela interacção com os mergulhadores, e num cenário de expansão significativa desta actividade, os impactos na fauna local poderão aumentar, com consequências para os ecossistemas de fundos rochosos da costa sul de Portugal. Na sua globalidade, a informação recolhida nos estudos que contemplam esta tese, por ser em grande parte totalmente nova para a região, espera-se que contribua para a gestão da zona costeira do Algarve.

Predation in Ground-Nesting Birds: an Experimental Study Using Natural Egg-Color Variation

We tested the hypothesis that cryptically colored eggs would suffer less predation than conspicuous eggs in the ground-nesting red-legged partridge, Alectoris rufa. We used A. rufa as a model species because it has a wide range of natural egg colors, the eggs are widely available from breeding farms, and nests are easily mimicked because they are scrapes containing no vegetation. The study was conducted in the spring of 2001 in forest and fallow fields of central Spain in Castilla La Mancha, Ciudad Real. We used 384 clutches of natural eggs that were white, white spotted, brown, or brown spotted. Within clutches, eggs were consistent in color and size; among clutches, color differences were distributed across habitats. Clutches were checked once after 2 wk of exposure. Cryptic coloration had a survival advantage that was dependent on the local suite of predators. Rodent predation was nonselective with respect to clutch color; however, avian predation was significantly higher for conspicuous clutches. In addition, there was an interaction of landscape and egg color for avian predation. In forest landscapes, the clutches with highest survival were brown spotted, whereas in fallow landscapes, brown and brown spotted clutches had higher survival than white and white potted clutches. Thus, both the predator suite and the landscape had significant effects on the value of cryptic egg coloration. Our study is relevant for conservationists and managers in charge of restocking programs in hunting areas. The release of other partridge species or their hybrids could result in hybridization with wild partridges, potentially leading to nonoptimal clutch pigmentation and reduced survival of the native species. We therefore recommend that local authorities, managers, and conservationists be cautious with the use of alien species and hybrids and release only autochthonous species of partridges within their natural ranges.

Ultraviolet cues affect the foraging behaviour of blue tits

The function of avian ultraviolet (UV) vision is only just beginning to be understood. One plausible hypothesis is that UV vision enhances the foraging ability of birds. To test this, we carried out behavioural experiments using wild-caught blue tits foraging for cabbage moth and winter moth caterpillars on natural and artificial backgrounds. The light environment in our experiments was manipulated using either UV-blocking or UV-transmitting filters. We found that the blue tits tended to find the first prey item (out of four) more quickly when UV cues were present. This suggests that UV vision offers benefits to birds when searching for cryptic prey despite the prey and backgrounds reflecting relatively little UV Although there was no direct effect of UV on the time taken to find all four prey items in a trial, search performance in the absence of UV wavelengths tended to increase over the course of an experiment. This may reflect changes in the search tactics of the birds. To our knowledge, these are the first data to suggest that birds use UV cues to detect cryptic insect prey and have implications for our understanding of protective coloration.

Predator mixes and the conspicuousness of aposematic signals

Conspicuous warning signals of unprofitable prey are a defense against visually hunting predators. They work because predators learn to associate unprofitability with bright coloration and because strong signals are detectable and memorable. However, many species that can be considered defended are not very conspicuous; they have weak warning signals. This phenomenon has previously been ignored in models and experiments. In addition, there is significant within- and among-species variation among predators in their search behavior, in their visual, cognitive, and learning abilities, and in their resistance to defenses. In this article we explore the effects of variable predators on models that combine positive frequency-dependent, frequency-independent, and negative frequency-dependent predation and show that weak signaling of aposematic species can evolve if predators vary in their tendency to attack defended prey.

Comparing entire colour patterns as birds see them

Colour patterns and their visual backgrounds consist of a mosaic of patches that vary in colour, brightness, size, shape and position. Most studies of crypsis, aposematism, sexual selection, or other forms of signalling concentrate on one or two patch classes (colours), either ignoring the rest of the colour pattern, or analysing the patches separately. We summarize methods of comparing colour patterns making use of known properties of bird eyes. The methods are easily modifiable for other animal visual systems. We present a new statistical method to compare entire colour patterns rather than comparing multiple pairs of patches. Unlike previous methods, the new method detects differences in the relationships among the colours, not just differences in colours. We present tests of the method's ability to detect a variety of kinds of differences between natural colour patterns and provide suggestions for analysis.

Animal visual systems and the evolution of color patterns; sensory processing illuminates signal evolution

Animal color pattern phenotypes evolve rapidly. What influences their evolution? Because color patterns are used in communication, selection for signal efficacy, relative to the intended receiver's visual system, may explain and predict the direction of evolution. We investigated this in bowerbirds, whose color patterns consist of plumage, bower structure, and ornaments and whose visual displays are presented under predictable visual conditions. We used data on avian vision, environmental conditions, color pattern properties, and an estimate of the bowerbird phylogeny to test hypotheses about evolutionary effects of visual processing. Different components of the color pattern evolve differently. Plumage sexual dimorphism increased and then decreased, while overall (plumage plus bower) visual contrast increased. The use of bowers allows relative crypsis of the bird but increased efficacy of the signal as a whole. Ornaments do not elaborate existing plumage features but instead are innovations (new color schemes) that increase signal efficacy. Isolation between species could be facilitated by plumage but not ornaments, because we observed character displacement only in plumage. Bowerbird color pattern evolution is at least partially predictable from the function of the visual system and from knowledge of different functions of different components of the color patterns. This provides clues to how more constrained visual signaling systems may evolve.

The bright incubate at night: sexual dichromatism and adaptive incubation division in an open-nesting shorebird

Ornamentation of parents poses a high risk for offspring because it reduces cryptic nest defence. Over a century ago, Wallace proposed that sexual dichromatism enhances crypsis of open-nesting females although subsequent studies found that dichromatism per se is not necessarily adaptive. We tested whether reduced female ornamentation in a sexually dichromatic species reduces the risk of clutch depredation and leads to adaptive parental roles in the red-capped plover Charadrius ruficapillus, a species with biparental incubation. Males had significantly brighter and redder head coloration than females. During daytime, when visually foraging predators are active, colour-matched model males incurred a higher risk of clutch depredation than females, whereas at night there was no difference in depredation risk between sexes. In turn, red-capped plovers maintained a strongly diurnal/nocturnal division of parental care during incubation, with males attending the nest largely at night when visual predators were inactive and females incubating during the day. We found support for Wallace's conclusion that reduced female ornamentation provides a selective advantage when reproductive success is threatened by visually foraging predators. We conclude that predators may alter their prey's parental care patterns and therefore may affect parental cooperation during care.

Ventilatory responses to skin extract in catfish

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Tribute to R.G. Boutilier: Skin colour and body temperature changes in basking Bokermannohyla alvarengai (Bokermann 1956)

In amphibians solar basking far from water sources is relatively uncommon since the highly permeable amphibian skin does not represent a significant barrier to the accompanying risk of losing water by evaporation. A South American frog, Bokermannohyla alvarengai (Bokermann 1956), however, spends a significant amount of the day exposed to full sun and relatively high temperatures. The means by which this frog copes with potentially high rates of evaporative water loss and high body temperatures are unknown. Thus, in this study, skin colour changes, body surface temperature, and evaporative water loss rates were examined under a mixture of field and laboratory conditions to ascertain whether changes in skin reflectivity play an important role in this animal's thermal and hydric balance. Field data demonstrated a tight correlation between the lightness of skin colour and frog temperature, with lighter frogs being captured possessing higher body temperatures. Laboratory experiments supported this relationship, revealing that frogs kept in the dark or at lower temperatures (20 degrees C) had darker skin colours, whereas frogs kept in the light or higher temperatures (30 degrees C) had skin colours of a lighter hue. Light exhibited a stronger influence on skin colour than temperature alone, suggesting that colour change is triggered by the increase in incident solar energy and in anticipation of changes in body temperature. This conclusion is corroborated by the observation that cold, darkly coloured frogs placed in the sun rapidly became lighter in colour during the initial warming up period (over the first 5 min), after which they warmed up more slowly and underwent a further, albeit slower, lightening of skin colour. Surprisingly, despite its natural disposition to bask in the sun, this species does not possess a 'waterproof' skin, since its rates of evaporative water loss were not dissimilar from many hylid species that live in arboreal or semi-aquatic environments. The natural history of B. alvarengai is largely unknown and, therefore, it is likely that the herein reported colour change and basking behaviour represent a complex interaction between thermoregulation and water balance with other ecologically relevant functions, such as crypsis.