Does spatial variation in predation pressure modulate selection for aposematism?

It is widely believed that aposematic signals should be conspicuous, but in nature, they vary from highly conspicuous to near cryptic. Current theory, including the honest signal or trade‐off hypotheses of the toxicity–conspicuousness relationship, cannot explain why adequately toxic species vary substantially in their conspicuousness. Through a study of similarly toxic Danainae (Nymphalidae) butterflies and their mimics that vary remarkably in their conspicuousness, we show that the benefits of conspicuousness vary along a gradient of predation pressure. Highly conspicuous butterflies experienced lower avian attack rates when background predation pressure was low, but attack rates increased rapidly as background predation pressure increased. Conversely, the least conspicuous butterflies experienced higher attack rates at low predation pressures, but at high predation pressures, they appeared to benefit from crypsis. Attack rates of intermediately conspicuous butterflies remained moderate and constant along the predation pressure gradient. Mimics had a similar pattern but higher attack rates than their models and mimics tended to imitate the signal of less attacked model species along the predation pressure gradient. Predation pressure modulated signal fitness provides a possible mechanism for the maintenance of variation in conspicuousness of aposematic signals, as well as the initial survival of conspicuous signals in cryptic populations in the process of aposematic signal evolution, and an alternative explanation for the evolutionary gain and loss of mimicry.     

Conservation implications of midden selection and use in an endangered gazelle (Gazella gazella)

This study examines midden selection and use by mountain gazelles Gazella gazella in central Saudia Arabia and the implications of middens (dung piles) for the conservation of this species. Specifically, we tested poachers' claims that mountain gazelles are easy to kill because midden use was predictable and conspicuous. We found that midden selection in mountain gazelles is non-random, with middens typically located at the base of the most conspicuous and largest trees, although the choice of midden sites is also influenced by tree species. Middens clearly have olfactory importance; the majority of mountain gazelles that visited middens were observed sniffing the ground. Mountain gazelles visit middens at night, which corroborates poachers' claims that the animals are easily hunted at night through spotlighting. Although the use of open habitats that are accessible to automobiles makes gazelles vulnerable to hunting and being run down, our data suggest that midden use is also an attribute that makes gazelles susceptible to hunting. Poachers can locate gazelles because of middens' predictable locations, as they are placed at prominent landmarks, are visually conspicuous as large dung piles, and are visited at night, which allows the animals to be found through spotlighting. Other taxa that use predictable and prominent landmarks, or communicate through visually conspicuous methods could also be susceptible to poaching.     

Mixed-phenotype grouping: the interaction between oddity and crypsis

Aggregations of different-looking animals are frequently seen in nature, despite well-documented selection pressures on individuals to maintain phenotypically homogenous groups. Two well-known theories, the ‘confusion effect’ (reduced ability of a predator to accurately target an individual in a group) and the ‘oddity effect’ (preferential targeting of phenotypically distinct, ‘odd’, individuals) act together to predict the evolution of behaviours in prey that lead to groups of animals that are homogeneous in appearance. In contrast, a recently proposed mechanism suggests that mixed groups could be maintained if one species in a mixed group is more conspicuous against the habitat than the other, as confusion effects generated by the conspicuous species impede predator targeting of the cryptic species; thus, cryptic species benefit from association with conspicuous ones. We test these contrasting predictions from the perspective of both predators and prey, and show that cryptic individual Daphnia are at reduced risk of predation from three-spine sticklebacks Gasterosteus aculeatus when in mixed-phenotype groups, a risk that is reduced further as the number of conspicuous individuals increases, supporting the hypothesis for the evolution of mixed groups. In contrast, while the preference for associating with colour-matched conspecifics by mollies (Poecilia sphenops) was reduced when they were cryptic, we found no evidence for active association with conspicuous conspecifics. We conclude that prey animals must balance the relative risks of oddity and conspicuousness in their social decisions, and that this could potentially lead to the evolution of mixed-phenotype grouping as a response to predation risk alone.     

Cryptic and conspicuous coloration in the pelagic environment

Despite the importance of cryptic and conspicuous coloration in pelagic ecosystems, few researchers have investigated the optimal reflectance spectra for either trait. In this study, the underwater radiance distribution in tropical oceanic water was modelled using measured inherent optical properties and radiative transfer calculations. The modelled light field was then used to predict the reflectance spectra that resulted in minimal or maximal object contrast as a function of depth, viewing angle, azimuth and solar elevation. The results matched commonly observed trends in the coloration of many pelagic organisms and showed that optimal coloration for either crypticity or conspicuity is a complex function of the parameters examined. The effects of viewing angle and depth were substantial and non-intuitive, showing that red coloration is most cryptic at depth. The effects of viewing azimuth were less significant and the effects of solar elevation were minor. White coloration and black coloration were equally cryptic/conspicuous when viewed from below. Although conspicuous objects viewed from below had the lowest contrast when viewed from a short distance, they had the longest sighting distances. The contrast of maximally conspicuous objects viewed from short distances was greatest at wavelengths displaced from the wavelength of maximum light penetration.     

The evolution of warning signals as reliable indicators of prey defense

It is widely argued that defended prey have tended to evolve conspicuous traits because predators more readily learn to avoid defended prey when they are conspicuous. However, a rival theory proposes that defended prey have evolved such characters because it allows them to be distinguished from undefended prey. Here we investigated how the attributes of defended (unprofitable) and undefended (profitable) computer-generated prey species tended to evolve when they were subject to selection by foraging humans. When cryptic forms of defended and undefended species were similar in appearance but their conspicuous forms were not, defended prey became conspicuous while undefended prey remained cryptic. Indeed, in all of our experiments, defended prey invariably evolved any trait that enabled them to be distinguished from undefended prey, even if such traits were cryptic. When conspicuous mutants of defended prey were extremely rare, they frequently overcame their initial disadvantage by chance. When Batesian mimicry of defended species was possible, defended prey evolved unique traits or characteristics that would make undefended prey vulnerable. Overall, our work supports the contention that warning signals are selected for their reliability as indicators of defense rather than to capitalize on any inherent educational biases of predators.     

Can the dietary conservatism of predators compensate for positive frequency dependent selection against rare,conspicuous prey?

The evolution of conspicuous colouration in prey is puzzling because such coloration attracts the attention of predators. Anti-apostatic selection, in which rare prey forms are predated disproportionately often, is a second potential obstacle to the evolution of conspicuous colouration in prey, as bright novel prey forms are likely to be very rare when they first appear in populations. It has recently been postulated that dietary conservatism in predators, an extended feeding avoidance of novel prey, would allow novel conspicuous prey to survive and multiply despite anti-apostatic and conspicuousness effects. We tested this hypothesis for a novel prey type arising in an otherwise cryptic population, providing a direct test of whether anti-apostatic selection or the predators’ wariness to attack the novel prey type is the more important force acting on the novel conspicuous prey. We conducted our experiment in the “Novel World”; an experimental system designed to test predators’ foraging decisions in a large landscape. We found that the conspicuous, novel prey suffered high initial costs of conspicuousness compared with cryptic prey, since most of these prey were attacked during the first “generation”, with no opportunity to “reproduce”. However, a subset of the 17 birds (24%) were following a dietary conservative foraging strategy and they were reluctant to eat the novel prey. Interestingly these birds were not more neophobic or less explorative. Our data demonstrate how difficult it is for the novel conspicuous prey to survive in cryptic populations, but they also highlight the importance of the predator’s foraging strategies in helping to promote the evolution and maintenance of aposematism.     

Dangerous liaisons: the predation risks of receiving social signals

Individuals are at risk when communicating because conspicuous signals attract both conspecifics and eavesdropping predators. This predation cost of communicating has typically been attributed to signalling individuals because of their conspicuous role, and is a core concept within sexual selection and communication ecology. But, if predators are attracted to signals, then receivers, both intended or otherwise, may also find themselves at risk of predation. Here, we review the theoretical basis and empirical evidence that receiving also carries a risk of predation. We distinguish between the risks of receiving and responding to signals, and we argue that receivers of signals that are long lived, are highly predictable in time or place and/or cannot be received quickly are likely to be at greater risk of predation compared to receivers of signals without these properties. We review recent empirical evidence from a variety of taxa that supports the hypothesis that receivers (including heterospecific prey) are aware of these risks and that they modify their behaviour to balance the risks against the benefits of receiving under predation threat. We also discuss the wider implications of risky receiving for receiving and signalling behaviour in prey, as well as for the prey's predators.     

Histological study on the oocyte filaments of the silverside Odonthestes bonariensis

Highly developed adhesive structures are characteristic of the eggs of Odonthestes bonariensis . At spawning each egg is connected to the other ones by four to six filaments. These are extroverted only in mature eggs, about 4–5 mm long and 10μm thick, starting from coneshaped bases at the egg's surface. Histologically, no filaments are detectable during the primary growth phase of the oocytes. At the lipidic vesicle stage, rounded eosinophilic filaments appear at the cell's surface. During the following stages they increase in volume and length, running along the whole oocyte, and lose their eosinophily. The filaments are beneath the follicular cells; according to preliminary TEM observations, they derive from the zona radiata. When the oocyte is mature, the theca externa disintegrates, freeing the filaments. These rather conspicuous and complex structures are markedly different from the adhesive egg fibrils observed in other teleosts.     

Clutch predation in great tinamous Tinamus major and implications for the evolution of egg color

Egg camouflage has been found to reduce predation in several ground‐nesting species. Therefore, the evolution of eggs that lack camouflage in ground nesting birds is puzzling. Even though clutch predation in the tropics is high, tinamous are the only tropical ground‐nesting birds that do not build a nest and do not lay cryptic eggs. I studied predation of great tinamou clutches in a lowland tropical forest and found that risk of predation was higher during incubation when the eggs are covered by the parent, than during laying when they are exposed, suggesting that predators primarily use cues from the incubating males to locate the clutch and not cues from the eggs. Clutch size had no effect on predation rate, even though larger clutches are more conspicuous to a human observer. Predation by visual cues is likely reduced during incubation by the camouflaged plumage and high nest attendance of males. If most predators use cues from the incubating male and not the eggs to locate clutches, then conspicuous egg color may have evolved in great tinamous as an intra‐specific signal. I evaluate hypotheses that may explain the maintenance of conspicuous egg color in tinamous.     

A simulation of evasive mimicry in the wild

The hypothesis of evasive mimicry proposes that: (a) the conspicuous coloration of some animals has evolved to warn predators that they are good at avoiding capture; and (b) this conspicuous coloration is mimicked by other, less evasive species which thus gain protection from predation. We have tested these ideas with wild birds as the predators and red and yellow pastry 'baits' as the prey. We used a specially constructed bird table to make one of the colours good at 'escaping': when a bird was about to eat a bait the observer pulled a cord which activated a mechanism that made the baits of one colour (but not the other) drop out of the bird's reach. We first measured the overall selection by the naive birds by presenting, for one day, a 'control' of equal numbers of non-evasive red and non-evasive yellow baits. The birds were then trained for 20 days on equal numbers of evasive yellow and non-evasive red baits, and were then again given a second control (in which the yellows were now non-evasive 'mimics'). The training was repeated for 18 days but with red as the evasive colour, and a third control was then presented (with reds now as the mimics). The proportions attacked daily in the two training sessions suggested that the birds were learning to ignore yellows in the first and reds in the second. In comparison with the first control, fewer yellows were eaten after they had been trained to ignore escaping yellows, and more were eaten after they had been trained on escaping reds. We conclude that the results are further evidence for the hypothesis of evasive mimicry.     

Selection for social signalling drives the evolution of chameleon colour change

Rapid colour change is a remarkable natural phenomenon that has evolved in several vertebrate and invertebrate lineages. The two principal explanations for the evolution of this adaptive strategy are (1) natural selection for crypsis (camouflage) against a range of different backgrounds and (2) selection for conspicuous social signals that maximise detectability to conspecifics, yet minimise exposure to predators because they are only briefly displayed. Here we show that evolutionary shifts in capacity for colour change in southern African dwarf chameleons (Bradypodion spp.) are associated with increasingly conspicuous signals used in male contests and courtship. To the chameleon visual system, species showing the most dramatic colour change display social signals that contrast most against the environmental background and amongst adjacent body regions. We found no evidence for the crypsis hypothesis, a finding reinforced by visual models of how both chameleons and their avian predators perceive chameleon colour variation. Instead, our results suggest that selection for conspicuous social signals drives the evolution of colour change in this system, supporting the view that transitory display traits should be under strong selection for signal detectability.     

Predator‐Associated Vocalizations in North American Red Squirrels (Tamiasciurus hudsonicus): To Whom are Alarm Calls Addressed and How do They Function?

Alarm vocalizations produced by prey species encountering predators can serve a variety of functions. North American red squirrels are a small-bodied mammal popularly known for producing loud, conspicuous alarm calls, but functional accounts of calling in this species are few and contradictory. We conducted research over a 3-yr period on a sample of 47 marked red squirrels in the foothills of the Canadian Rockies. We recorded the production of alarm calls during encounters with natural predators and in a series of simulated predator experiments. We tested for variation in call production patterns consistent with three traditional hypotheses concerning the conspecific warning functions of alarm calling: namely that they serve as warnings to kin, to potential mates, or to territorial neighbors with which callers have an established relationship. Patterns of calling did not provide clear support for any of these hypothesized functions. We consider several possible qualifications to our results. We also consider the possibility that conspicuous calls given by red squirrels during encounters with predators are directed at the predators themselves and function to announce their detection and possibly deter them. This possibility is consistent with additional life-history features of red squirrels including that they are a relatively solitary and territorial, food-hoarding species that produces the same conspicuous vocalizations in response to other squirrels intruding on their territory to steal cones. An important corollary of this account is that red squirrel alarm calls probably do not entail referentially specific messages about different types of predator, as proposed previously.     

Immunohistochemical studies on the spinal dorsal horn of the turtle Chrysemys d'orbigny.

Immunohistochemical methods were used to characterize some of the systems of nerve fibers occurring in the spinal dorsal horns of the turtle Chrysemys d'orbigny. Substance P (SP), calcitonin gene-related peptide (CGRP) and leuenkephalin (Enk) immunoreactive fibers were found concentrated in the superficial horn region, termed here synaptic field Ia. From this zone the immunoreactive fibers project to deeper dorsal horn regions. Comparison with histological images obtained after HRP labeling of dorsal root axons indicates that SP-, CGRP- and Enk-immunoreactive fibers are small-diameter primary sensory fibers entering the cord via synaptic field Ia. It is posulated here that these three substances may coexist in the same fibers. Enk-positive fibers also occur randomly scattered in the lateral funiculi, showing a conspicuous increase in density at the perimedullary plexus level. Tyrosine hydroxylase (TH) immunoreactive fibers were found in the more compact dorsal horn neuropil (synaptic field II) and also forming bilateral conspicuous bundles in the lateral funiculi. TH-immunoreactive cell bodies were found in the epithelium lining the central canal. Taking into account data derived from Golgi impregnated material it is proposed that they represent epithelial cells undergoing neural differentiation.     

Predator experience on cryptic prey affects the survival of conspicuous aposematic prey

Initially, aposematism, which is an unprofitable trait, e.g. noxiousness conspicuously advertised to predators, appears to be a paradox since conspicuousness should increase predation by naive predators. However, reluctance of predators for eating novel prey (e.g. neophobia) might balance the initial predation caused by inexperienced predators. We tested the novelty effects on initial predation and avoidance learning in two separate conspicuousness levels of aposematic prey by using a 'novel world' method. Half of the wild great tits (Parus major) were trained to eat cryptic prey prior to the introduction of an aposematic prey, which potentially creates a bias against the aposematic morph. Both prey types were equally novel for control birds and they should not have shown any biased reluctance for eating an aposematic prey. Knowledge of cryptic prey reduced the expected initial mortality of the conspicuous morph to a random level whereas control birds initially ate the conspicuous morph according to the visibility risk. Birds learned to avoid conspicuous prey in both treatments but knowledge of cryptic prey did not increase the rate of avoidance learning. Predators' knowledge of cryptic prey did not reduce the predation of the less conspicuous aposematic prey and additionally predators did not learn to avoid the less conspicuous prey. These results indicate that predator psychology, which was shown as reluctance for attacking novel conspicuous prey, might have been important in the evolution of aposematism.     

A preference for a sexual signal keeps females safe

Predation is generally thought to constrain sexual selection by female choice and limit the evolution of conspicuous sexual signals. Under high predation risk, females usually become less choosy, because they reduce their exposure to their predators by reducing the extent of their mate searching. However, predation need not weaken sexual selection if, under high predation risk, females exhibit stronger preferences for males that use conspicuous signals that help females avoid their predators. We tested this prediction in the fiddler crab Uca terpsichores by increasing females' perceived predation risk from crab-eating birds and measuring the attractiveness of a courtship signal that females use to find mates. The sexual signal is an arching mound of sand that males build at the openings of their burrows to which they attract females for mating. We found that the greater the risk, the more attractive were males with those structures. The benefits of mate preferences for sexual signals are usually thought to be linked to males' reproductive contributions to females or their young. Our study provides the first evidence that a female preference for a sexual signal can yield direct survival benefits by keeping females safe as they search for mates.     

Effect of oestrogenic, and androgenic and gestagenic hormones on the gametogenesis (oogenesis and spermatogenesis) in the snail Helix pomatia

In the ovotestis of Helix pomatia both oogenesis and spermatogenesis were influenced by treatments with steroid hormones produced in the gonads of higher vertebrates. Testosterone influenced gametogenesis to a small degree. Progesterone and oestrone-acetate at first stimulated ovogenesis, but they also acted on spermatogenesis. All three hormones examined influenced oogenesis in a conspicuous and significant way, while their effect on spermatogenesis was indistinct.     

Sympathetic innervation of the carotid bifurcation in the rabbit and cat: Blood vessels,carotid body and carotid sinus

Summary Two postganglionic branches of the superior cervical ganglion enter the area of the carotid bifurcation in the rabbit and the cat. The common and external carotid arteries receive a rich adrenergic nerve supply, which can be demonstrated by fluorophores of biogenic amines appearing after formaldehyde treatment. The internal carotid artery is only sparsely innervated; however, it shows a dense sympathetic supply at the site of pressor receptors. Following removal of the superior cervical ganglion, a total loss of fluorescent adrenergic nerves occurs and degeneration of nerve endings possessing dense core vesicles is conspicuous. These nerve terminals are situated mainly subendothelially in the carotid body sinusoids; they only rarely terminate on type I cells.     

Perspective: the evolution of warning coloration is not paradoxical

Animals that are brightly colored have intrigued scientists since the time of Darwin, because it seems surprising that prey should have evolved to be clearly visible to predators. Often this self-advertisement is explained by the prey being unprofitable in some way, with the conspicuous warning coloration helping to protect the prey because it signals to potential predators that the prey is unprofitable. However, such signals only work in this way once predators have learned to associate the conspicuous color with the unprofitability of the prey. The evolution of warning coloration is still widely considered to be a paradox, because it has traditionally been assumed that the very first brightly colored individuals would be at an immediate selective disadvantage because of their greater conspicuousness to predators that are naive to the meaning of the signal. As a result, it has been difficult to understand how a novel conspicuous color morph could ever avoid extinction for long enough for predators to become educated about the signal. Thus, the traditional view that the evolution of warning coloration is difficult to explain rests entirely on assumptions about the foraging behavior of predators. However, we review recent evidence from a range of studies of predator foraging decisions, which refute these established assumptions. These studies show that: (1) Many predators are so conservative in their food preferences that even very conspicuous novel prey morphs are not necessarily at a selective disadvantage. (2) The survival and spread of novel color morphs can be simulated in field and aviary experiments using real predators (birds) foraging on successive generations of artificial prey populations. This work demonstrates that the foraging preferences of predators can regularly (though not always) result in the increase to fixation of a novel morph appearing in a population of familiar-colored prey. Such fixation events occur even if both novel and familiar prey are fully palatable and despite the novel food being much more conspicuous than the familiar prey. These studies therefore provide strong empirical evidence that conspicuous coloration can evolve readily, and repeatedly, as a result of the conservative foraging decisions of predators.     

Subterranean “oasis” in the Brazilian semiarid region: neglected sources of biodiversity

Biodiversity and Conservation - Semiarid regions experience conspicuous seasonal variations, especially related to precipitation. Caves in these areas can be exceptions since they are less affected...     

Losing the trait without losing the signal: Evolutionary shifts in communicative colour signalling

Colour signalling traits are often lost over evolutionary time, perhaps because they increase vulnerability to visual predators or lose relevance in terms of sexual selection. Here, we used spectrometric and phylogenetic comparative analyses to ask whether four independent losses of a sexually selected blue patch are spectrally similar, and whether these losses equate to a decrease in conspicuousness or to loss of a signal. We found that patches were lost in two distinct ways: either increasing reflectance primarily at very long or at very short wavelengths, and that species with additional colour elements (UV, green and pink) may be evolutionary intermediates. In addition, we found that patch spectral profiles of all species were closely aligned with visual receptors in the receiver's retina. We found that loss of the blue patch makes males less conspicuous in terms of chromatic conspicuousness, but more conspicuous in terms of achromatic contrast, and that sexual dimorphism often persists regardless of patch loss. Dorsal surfaces were considerably more cryptic than were ventral surfaces, and species in which male bellies were the most similar in conspicuousness to their dorsal surfaces were also the most sexually dimorphic. These results emphasize the consistent importance of sexual selection and its flexible impact on different signal components through evolutionary time.