Conspicuous displays in cryptic males of a polytypic poison-dart frog

The evolution of aposematism is linked to increased opportunities for conspicuous sexual displays since detection by potential predators is no longer disadvantageous. Therefore, phenotypic divergence in aposematic species leading to relatively cryptic forms is expected to constrain such opportunities, by restoring the trade-off between natural and sexual selection on the boldness of sexual displays. We asked if and how a derived phenotype of the poison-dart frog Oophaga granulifera that appears relatively cryptic to potential predators exhibits conspicuous sexual displays for potential mates. We used visual modeling of frog contrasts against their natural backgrounds to test if for conspecifics green frogs appear less conspicuous than red frogs as they do for birds. We conducted behavioral observations of focal red and green males to determine if green frogs adjust their display behavior to the availability of potential mates. Dorsal brightness is known to influence female preferences in at least one poison frog species. We found that, despite being less visible under some measures, green frogs may appear as bright as red frogs for conspecifics but not birds, when viewed on dark backgrounds. Additionally, green males called more intermittently than red males when advertising to distant females, but they exhibited a dramatic increase in calling activity in proximity of a female and were as active as red males in this context. Together, our results suggest that green frogs retain context-dependent conspicuousness to conspecifics despite the evolution of relative crypsis to potential predators.     

Blue eggs do not reduce nest predation in the song thrush,Turdus philomelos

Summary Many passerine birds with open cup-shaped nests lay blue or blue-green eggs. In thrushes, blue eggs may be cryptic and provide camouflage by imitating spots of light on green leaves. Alternatively, egg coloration may be selectively neutral because nest predators detect nests and not eggs, or it may be maladaptive because organisms are not always well adapted to their present environment. I evaluated these hypotheses by studying predation on artificial song thrush (Turdus philomelos) nests with quail eggs, painted either white, blue, or spotted (cryptic to a human eye). Corvids were the major nest predators. For concealed as well as exposed nests, I found no differences in the predation rates of nests with white, blue, or spotted eggs. Predators apparently detected the nests, and not the eggs, first. In a second experiment, I placed egg groups without nests in trees to study the effect of color per se. The predation rate of the spotted egg groups was significantly lower than that of the white and blue egg groups, for concealed as well as exposed egg groups. These results suggest that blue eggs in the song thrush are not cryptic but may be selectively neutral or even maladaptive with regard to nest predation.     

Egg coloration in ring-billed gulls (<Emphasis Type="Italic">Larus delawarensis</Emphasis>): a test of the sexual signaling hypothesis

Although many avian eggs appear to be cryptically colored, many species also lay vibrant blue green eggs. This seemingly conspicuous coloration has puzzled biologists since Wallace, as natural selection should favor reduced egg visibility to minimize predation pressure. The sexual signaling hypothesis posits that blue green egg coloration serves as a signal of female quality and that males exert post-mating sexual selection on this trait by investing more in the nests of females laying more intensely blue green eggs. This hypothesis has received mixed support to date, and most previous studies have been conducted in cavity-nesting species where male evaluation of his partner’s egg coloration, relative to that of other females, may be somewhat limited. In this study, we test the sexual signaling hypothesis in colonially nesting ring-billed gulls (Larus delawarensis) where males have ample opportunity to assess their mate’s egg coloration relative to that of other females. We used correlational data and an experimental manipulation to test four assumptions and predictions of the sexual signaling hypothesis: (1) blue green pigmentation should be limiting to females; (2) extent of blue green egg coloration should relate to female quality; (3) extent of blue green egg coloration should relate to offspring quality; and (4) males should provide more care to clutches with higher blue green chroma. Our data provide little support for these predictions of the sexual signaling hypothesis in ring-billed gulls. In light of this and other empirical data, we encourage future studies to consider additional hypotheses for the evolution of blue green egg coloration.     

Bright plumage in the magpie: does it increase or reduce the risk of predation?

Conspicuous color patterns in birds may lead to increased risk of predation. Alternatively, bright birds may be aposematic or unprofitable prey, which leads to decreased predation. During four autumns, I examined whether the black-and-white plumage of (stuffed) magpies Pica pica increases or reduces the risk of attack by migrating goshawks Accipiter gentilis. Attack risk was higher for normal magpies than for cryptic, brown magpies whether the mounts were placed near one another or far apart. However, the brown magpie may have been avoided because of its novelty. For magpies and cryptic jays Garrulus glandarius exposed together, the attack risk was similar. In 2 years, magpie and jay mounts were also exposed far apart. In 1994, with an invasion of migrating jays, attack risk was much higher for jays than for magpies. In 1995, with a normal density of jays, hawks attacked the magpies more often. The results demonstrate frequency-dependent prey selection by goshawks, which would influence any predation cost of bright plumage. The attacks on normally colored magpie mounts suggest that magpies are not aposematic. Trials with photographs and human observers indicated that normal magpies were somewhat easier to detect than jays. The plumage of the magpie possibly increases the risk of predation, but may be favoured by sexual or social selection. Received: 18 January 1996 / Accepted after revision: 6 October 1996     

Vivid birds respond more to acoustic signals of predators

Because conspicuous morphology such as colorful plumage may increase predation risk, we aimed to see if variation in plumage coloration could explain variation in avian anti-predator behavior. We included several measures of plumage coloration: human perception of vividness from images in field guides, total intensity from reflectance spectra of museum skins, contrasts calculated from physiological models of these spectra parameterized for both raptors and humans, chroma, and spectral saturation. We investigated how well these measurements predicted risk assessment in ten species of birds in St. John, U.S. Virgin Islands. We quantified how each species responded to playbacks of a predator’s calls and compared this response to that elicited by songs from a non-predatory, sympatric bird. We found that human-determined measures of vividness best predicted anti-predator responses of birds—more vividly colored species responded more to predators than duller species. No spectrophotometric variable explained variation in species reactions to a predator call. Our results suggest that vivid birds may compensate for their conspicuousness by being more responsive to the sound of predators and that more work is needed to better evaluate how animal coloration is quantified in comparative studies.     

Spectral reflectance and communication in the wolf spider, <Emphasis Type="Italic">Schizocosa ocreata</Emphasis> (Hentz): simultaneous crypsis and background contrast in visual signals

Animal color patterns often reflect a compromise between natural selection for crypsis or inconspicuousness to predators and sexual selection for conspicuousness to potential mates. In leaf litter-dwelling wolf spider species like Schizocosa ocreata, body coloration often closely matches the background coloration of a generally brown environment. However, body parts used in communication should exhibit high contrast against background coloration. We used spectral analysis to examine male and female S. ocreata for matching and contrasting coloration against leaf litter. Values were plotted in multivariate color space, based on reflectivity in different frequency ranges. When viewed from above, colors of both males and females overlap with values for dead brown leaf litter and soil, suggesting cryptic coloration when viewed by potential predators. However, when viewed from a lateral perspective, both males and females show color values that are polar opposites of litter backgrounds, suggesting higher contrast when viewed by other spiders. Moreover, male secondary characters used in visual signaling by S. ocreata (tibia brushes) show the highest level of background contrast. These findings suggest that S. ocreata wolf spiders have color patterns that provide both crypsis and background contrast at the same time, depending on receiver viewing perspective.     

Taste-rejection by predators and the evolution of unpalatability in prey

Aposematic species advertise their unpalatability to potential predators using conspicuous warning colouration. The initial evolution of aposematism is thought to occur by warningly coloured mutants emerging in an already unpalatable cryptic species. However, possessing defence chemicals is often costly, and it is difficult to understand what the selective benefits might be for a mutation causing its bearer to be defended in a population of otherwise palatable cryptic prey. One solution to this problem is that chemically defended individuals are tasted and rejected by predators, and are, therefore, more likely to survive predatory attacks than undefended individuals. Using naïve domestic chicks Gallus gallus domesticus as predators and cryptic green chick crumbs as prey, we asked whether the accuracy with which birds discriminated between palatable and unpalatable prey was affected by the palatability of the unpalatable prey (moderately or highly defended), or their frequency in the population (10 or 25%). Birds could discriminate between green prey on the basis of their defences, and showed better discrimination between palatable and unpalatable prey when defended crumbs were highly unpalatable, compared to when they were moderately unpalatable. Although there was no detectable effect of the frequency of unpalatable prey in the population on predator taste-rejection behaviour in our main analysis, frequency did appear to affect the strategies that birds used in their foraging decisions when prey were only moderately unpalatable. How birds used taste to reject prey also suggests that birds may be able to monitor and regulate their chemical intake according to the frequency and defence levels of the unpalatable prey. Taken together, these results show that avian predators can generate selection for unpalatability in cryptic prey by sampling and taste-rejecting prey, but that a relatively large chemical difference between palatable and unpalatable prey may be necessary before unpalatable prey can enjoy a selective advantage. The exact nature of this evolutionary dynamic will depend on other environmental factors, such as defence costs and prey availability, but it provides a mechanism by which defences can evolve in a cryptic population.     

Carotenoid-derived ornaments reflect parental quality in male and female yellow-eyed penguins (<Emphasis Type="Italic">Megadyptes antipodes</Emphasis>)

The handicap principle suggests that ornamental traits that function as honest signals in mate selection must be costly to be effective. We evaluated in the sexually monochromatic yellow-eyed penguin (Megadyptes antipodes) whether the carotenoid-derived plumage and eye coloration predicts parental quality and whether males and females within pairs mate assortatively in relation to these carotenoid-derived ornaments. In addition, we investigated whether age or body condition was related to the coloration of the ornamental traits. In yellow-eyed penguins, parental quality of males and females was predicted by eye and head plumage coloration. Even when we controlled for gender- and age-specific differences, eye and head plumage coloration reflected honestly parental quality. Males and females mated assortatively in relation to these ornamental traits. While age influenced coloration of both the eye and head plumage, body condition was related only to the saturation of plumage coloration. These results provide evidence that the carotenoid-derived ornaments in yellow-eyed penguins reflect the parental abilities of birds and, therefore, may be costly signals. Potentially, female and male yellow-eyed penguins could use eye and plumage coloration as an indirect cue in assessing age and quality of individual birds during mate choice. This is only the second study to examine plumage coloration in relation to sexual selection in penguins, while conspicuous ornamental traits in other species of penguin beg the question whether they also play a role in sexual selection.Electronic Supplementary Material Supplementary material is available in the online version of this article at Communicated by C.R. Brown     

Does egg colour affect predation rate on open passerine nests?

The breeding success of many passerines is strongly reduced by egg predation. The adaptive significance of egg crypsis in open nesters is often taken for granted, but visually searching predators may first detect the nest or adult bird and not the eggs. Götmark predicted that selection should favour egg crypsis in the absence of conspicuous nests, whereas birds with conspicuous nests should have non-cryptic eggs. I compared the effect of egg colour treatment (white, blue, brown-spotted) on nest survival (1) among species characterized by different egg coloration, nest size and nest placement, and (2) between relatively well and poorly concealed nests within species. I used artificial nests (n=1,296) and eggs mimicking (except in egg colour) those of the yellowhammer (Emberiza citrinella), blackcap (Sylvia atricapilla) and song thrush (Turdus philomelos). Concurrently, I monitored survival of real nests (n=1,106). Nest survival differed among species, increased with nest concealment and throughout the breeding season, but was not significantly related to egg colour in any species. Nevertheless, the data for the yellowhammer suggest a trend in survival rates across the colour treatments. Brown eggs survived better than white eggs by 11% and 4% in 2 years, but this study had insufficient power to detect effects of this size. The results thus suggest that egg coloration in the song thrush and blackcap (shrub nesters) may be a neutral trait with regard to nest predation, whereas egg crypsis may be an anti-predation feature for the yellowhammer (ground/near-ground nester). The role of predation in the evolution of eggshell colour may vary not only between cavity and open nesters, but also across nest sites within the latter group.     

Sexual selection and competitive mating success of males guppies (Poecilia reticulata) from four Trinidad populations

Summary Luyten and Liley (1985) obtained evidence that in populations of Trinidad guppies from two clear headwater streams, where Rivulus harth is the major aquatic predator, selection has favored visual aspects of male mating behavior — display activity and conspicuous coloration. Males in two lowland turbid stream populations, with numerous aquatic predators, were found to rely more heavily upon non-display components of courtship. In this study males of the four populations were placed in competition in clear water for mating with females of the same populations. Males of two populations were allowed to compete in turbid water. Mating success was determined on the basis of the frequency of insemination and the proportion of sperm attributable to males of the competing populations. The sperm of competing males were identified by radioisotope labelling and autoradiographic techniques. Males from headwater populations were more successful than lowland males in mating with headwater females in clear water. There was no difference in mating success of headwater and lowland males in competition for lowland females in clear water, but in turbid water lowland males were more successful in inseminating lowland females. Males were more successful in mating with females of their own population when the two headwater population were in competition. The results are consistent with the hypothesis that conspicuous coloration and high frequency and duration of display are adaptations to clear water conditions, and that non-display tactics are adapted to turbid water. Evidence of intrapopulation preferences suggest that male traits and female sexual responses in the guppy have evolved in parallel as predicted by sexual selection theory.     

Impact of Invasive Cane Toads on Australian Birds

Abstract: The cane toad (Bufo marinus), a large, toxic, American anuran, was introduced to Australia in 1935. Populations of many of Australia's reptiles (snakes, varanid lizards, crocodiles) and carnivorous mammals (dasyurid marsupials) have declined because these predators are killed by the toad's powerful toxins. In contrast to these well‐studied species, little is known about the cane toads impacts on Australian birds. We reviewed published and unpublished data on behavioral interactions between Australian avian predators and cane toads and collated distributional and dietary information to identify avian taxa potentially at risk from cane toad invasion. Cane toads are sympatric with 172 frog‐eating bird species in Australia, and an additional 8 bird species overlap with the predicted future range of the toad. Although many bird species thus are potentially at risk, behavioral observations suggest the risk level is generally low. Despite occasional reports of Australian birds being killed when they ingest cane toads, most birds either ignore toads or survive the predation event. The apparently higher tolerance of Australian birds to toad toxins, compared with Australian reptiles and marsupials, may reflect genetic exchange between Australian birds and Asian populations that encounter other bufonid species regularly and hence have evolved the capacity to recognize or tolerate this toxic prey.     

An experimental test of female choice relative to male structural coloration in eastern bluebirds

Several experimental studies have shown that female birds use ornamental melanin and carotenoid plumage coloration as criteria in mate choice. Whether females choose mates based on natural variation in structural coloration, however, has not been well established. Male eastern bluebirds (Sialia sialis) display brilliant ultraviolet (UV)-blue plumage coloration on their head, back, wings, and tail, which is positively correlated with condition, reproductive effort, and reproductive success. We experimentally tested the hypothesis that female eastern bluebirds prefer as mates males that display brighter structural coloration by presenting breeding-condition females with males of variable coloration. We conducted two types of mate-choice experiments. First, females chose between males whose coloration was manipulated within the natural range of variation in the population; feathers were either brightened with violet marker or dulled with black marker. Second, females chose between males with naturally dull or bright plumage coloration. In both manipulated and unmanipulated coloration trials, female choice did not differ significantly from random with respect to structural coloration. We found no support for the hypothesis that the UV–blue coloration of male eastern bluebirds functions as a criterion in female mate choice.     

Voices in the dark: predation risk by owls influences dusk singing in a diurnal passerine

Predation is an important cost of communication in animals and thus a potent selection pressure on the evolution of signaling behavior. Heterospecific eavesdropping by predators may increase the vulnerability of vocalizing prey, particularly during low light, such as at dusk when nocturnal predators are actively hunting. Despite the risk it entails, dawn and dusk chorusing is common in passerines. However, the dusk chorus has not been studied much, neglecting the opportunity for understanding how eavesdropping between predators and prey may shape communication in birds. Here, we report the first demonstration of simulated predation risk (playback of owl vocalizations) altering the dusk chorus of a diurnal passerine, the veery (Catharus fuscescens). Veeries have a pronounced dusk chorus, singing well after sunset and potentially exposing themselves to predation by owls. In response to brief playbacks of owl calls (~30 s of calls presented three times over 25 min), veeries sang fewer songs post-sunset and stopped singing earlier relative to control trials. These changes in singing remained evident 30 min after the last owl stimulus. Although the avian dusk chorus has received relatively little attention to date, our results suggest that the dusk chorus may pose a higher predation risk to singing males that may influence the evolution of singing behavior in diurnal birds.     

Delayed plumage maturation in Lazuli buntings: tests of the female mimicry and status signalling hypotheses

The evolutionary importance of delayed plumage maturation (DPM) in passerines, the condition when more than 1 year is required to achieve adult-like coloration, remains highly contentious. Adaptive hypotheses propose that aggression from after 2nd-year (ASY) males or predation favors DPM in 2nd-year (SY) males, thereby increasing SY male survivorship or reproductive success. However, each hypothesis suggests a distinct selective mechanism explaining “how” this is accomplished. Alternatively, DPM may be a consequence of a nonadaptive molt constraint. We tested the female mimicry and status signalling hypotheses in territorial ASY male lazuli buntings (Passerinaamoena) using three sets of model presentation experiments. The female mimicry hypothesis proposes that dull SY male plumage deceptively mimics female plumage, and predicts that ASY males can not distinguish SY male from female plumage. The status signalling hypothesis proposes that dull SY male plumage honestly signals low competitive threat, and predicts that ASY males respond less aggressively to dull versus bright, ASY-like plumage. Contrary to the female mimicry hypothesis, ASY males distinguished between SY male and female plumage, as they were aggressive to SY male models exclusively and attempted to copulate with female models. Supporting the status signalling hypothesis, ASY males were significantly less aggressive to SY versus ASY male plumage. While DPM may result from a physiological constraint on bright SY male plumage, our results support the idea that dull plumage in an SY male's first breeding season may be maintained by selection to reduce aggression from ASY males, serving as a signal of competitive status. Received: 21 February 1997 / Accepted after revision: 16 June 1997     

Satellites and sneakers: opportunistic male breeding tactics in pupfish (Cyprinodon pecosensis)

Summary Manipulations of population density, availability of oviposition substrate, and size of breeding habitat affected the proportion of pupfish (Cyprinodon pecosensis) males adopting territorial and satellite breeding tactics. Satellites occupied stations above breeding substrates defended by territorial males. Both territorial and satellite males developed a bright blue breeding coloration. A third breeding tactic, sneak-spawning, was occasionally observed. Sneak-spawning males retained the cryptic female coloration and occasionally spawned on territories. The lower reproductive success of satellites and sneak-spawning males suggested that both are conditional breeding tactics adopted by competitively inferior males. Satellites were present in all treatments that favored a territorial breeding system, but were absent when the breeding system was a dominance hierarchy. Satellites were associated with territorial males that had higher reproductive success and larger territories than males without satellites. Satellites functioned as parasites rather than as mutualists, since they disrupted spawnings and stole copulations from territorial males. The occurrence and frequency of the conditional breeding behaviours in pupfish represent facultative responses of males to changes in the intensity of competition for breeding sites and females.     

Species-specific antipredatory behaviours: effects on prey choice in different habitats

Summary Prey species may use many different behaviours to avoid predation. In this study, the antipredator behaviours of juvenile roach (Rutilus rutilus) and juvenile perch (Perca fluviatilis) were studied in wading pools with three kinds of structural complexity: no structure, structure simulating vegetation and structure simulating bottom crevices. Predation experiments with piscivorous perch and habitat choice experiments with the prey were performed, and the foraging success and prey choice of the predators were related to the type of structure. Predator foraging success was lower in the vegetation than in the other treatments. In the absence of structure and with vegetation structure, predators preferred perch over roach, while the preference was reversed in the crevice treatment. Roach and perch differed in their antipredatory behaviours. Roach responded to the presence of predators by schooling, moving fast and remaining at the surface, and escaped from attacks by jumping out of the water. In contrast, perch moved more slowly, dispersed after attacks and tried to hide at the bottom. Perch always preferred the vegetation structure to the non-structured part of the pool, while roach showed preference for the vegetation structure only when predators were present. Roach never occurred in crevices, whereas perch used crevices when predators where present. Predator pursuit speed was lower in the vegetation structure than in the non-structured treatment, but prey escape speed was unaffected. The results suggest that both the quantity and quality of structural complexity interacting with species-specific antipredator behaviours are important for predator-prey dynamics. It is also suggested that the presence of structure can have substantial effects on the structure of North Eurasian fish communities, by affecting relative and absolute predation pressures from piscivorous perch on prey species. Correspondence to: B. Christensen     

Experimental evidence that bright coloration is not important for territory defense in purple martins

Summary I tested the hypothesis that bright breeding plumage in territorial males acts as a badge of fighting ability or aggressive motivation to intimidate intruders. Territorial male purple martins (Progne subis) whose iridescent blue plumage was lightened to mimic the appearance of subadult males did not suffer an increase in intruder pressure or loss of territory compared with control males. Bright plumage color itself did not deter intruders and was not important for successful territory defense. Furthermore, a bright coloration of owners was not associated with an increased level of aggression toward intruders. Results from parallel studies on this species suggest that bright coloration is important in territory acquisition. The effectiveness of badges of fighting ability and aggressive motivation in territory defense is limited by whether intruders benefit from assessing these traits in owners. Differences in signaling systems between species are due in part to differences in floater tactics and the mode of territory acquisition.     

Evolution of Nesting Height in an Endangered Hawaiian Forest Bird in Response to a Non‐Native Predator

Abstract: The majority of bird extinctions since 1800 have occurred on islands, and non‐native predators have been the greatest threat to the persistence of island birds. Island endemic species often lack life‐history traits and behaviors that reduce the probability of predation and they can become evolutionarily trapped if they are unable to adapt, but few studies have examined the ability of island species to respond to novel predators. The greatest threat to the persistence of the Oahu Elepaio (Chasiempis ibidis), an endangered Hawaiian forest bird, is nest predation by non‐native black rats (Rattus rattus). I examined whether Oahu Elepaio nest placement has changed at the individual and population levels in response to rat predation by measuring nest height and determining whether each nest produced offspring from 1996 to 2011. Average height of Oahu Elepaio nests increased 50% over this 16‐year period, from 7.9 m (SE 1.7) to 12.0 m (SE 1.1). There was no net change in height of sequential nests made by individual birds, which means individual elepaios have not learned to place nests higher. Nests ≤3 m off the ground produced offspring less often, and the proportion of such nests declined over time, which suggests that nest‐building behavior has evolved through natural selection by predation. Nest success increased over time, which may increase the probability of long‐term persistence of the species. Rat control may facilitate the evolution of nesting height by slowing the rate of population decline and providing time for this adaptive response to spread through the population.     

Factors predicting male fertilization success in an external fertilizer

In postcopulatory sexual selection both sperm competition and cryptic female choice are considered to be important selective agents, but their relative importance for male fertilization success has received little attention. We tested whether sperm quality, male spawning coloration, male heterozygosity, and genetic overlap with the female explained a male’s fertilization success in controlled in vitro fertilization competition trials between equal numbers of sperm from pairs of male Arctic charr (Salvelinus alpinus), an external fertilizer. Offspring were genotyped to determine each males’ share of paternity. The velocity of a male’s sperm relative to the velocity of the competing male’s sperm was the best predictor of male fertilization success. Yet, sperm velocity was not related to spawning coloration or male heterozygosity. In fact, the most brightly colored male in a pair had the lowest fertilization probability. This could result from cryptic female choice for pale males, but might rather be a result of paler males producing more competitive sperm than more colored males. Furthermore, the more microsatellite alleles a male shared with the female relative to the competing male, the higher fertilization success he had. We argue that this latter may be an effect of assortative cryptic female choice, which might prevent hybridization with sympatric Arctic charr morphs or one form of kin selection.     

Chemosensory cues allow male lizards<Emphasis Type="Italic"> Psammodromus algirus</Emphasis> to override visual concealment of sexual identity by satellite males

Males of many species of lizards show conspicuous breeding colors but, in some species, young competitively inferior males conceal their sexual identity by a female-like dull coloration that allows them to evade aggression from dominant males and to adopt an alternative satellite-sneaking mating tactic. However, large males of the lizard Psammodromus algirus reacted aggressively to young intruder males despite their female-like coloration, suggesting that they might have the ability to recognize competitor males by chemosensory cues. We experimentally manipulated the head coloration (brown vs orange) and scents (male vs female) of small young males. For staged agonistic encounters, we compared the response of resident unmanipulated large males to the different manipulated small males. When we manipulated only the color of small males, the response of resident large males was independent of the paint manipulation; brown and orange males elicited a similar aggressive response. However, when we also manipulated the scent, small males painted orange or brown, but bearing the scent of males, received a significantly higher number of aggressive responses than small males painted orange or brown, but bearing the scent of females. The results showed that, at close range, the reaction of large males to manipulated individuals was dependent on the scent, whereas color seemed to be less important. Coloration may be, however, more important in long-distance communication as shown by the outcome of the first encounters. Also, orange coloration may increase the intensity of the aggressive response. Effective sex recognition by territorial large males is important in natural situations to avoid sneak matings by young male competitors. Thus, even if small males visually conceal their sexual identity, chemosensory cues allow large males to identify them at close range.