The evolution of Müllerian mimicry

It is now 130 years since Fritz Müller proposed an evolutionary explanation for the close similarity of co-existing unpalatable prey species, a phenomenon now known as Müllerian mimicry. Müller's hypothesis was that unpalatable species evolve a similar appearance to reduce the mortality involved in training predators to avoid them, and he backed up his arguments with a mathematical model in which predators attack a fixed number (n) of each distinct unpalatable type in a given season before avoiding them. Here, I review what has since been discovered about Müllerian mimicry and consider in particular its relationship to other forms of mimicry. Müller's specific model of associative learning involving a "fixed n" in a given season has not been supported, and several experiments now suggest that two distinct unpalatable prey types may be just as easy to learn to avoid as one. Nevertheless, Müller's general insight that novel unpalatable forms have higher mortality than common unpalatable forms as a result of predation has been well supported by field experiments. From its inception, there has been a heated debate over the nature of the relationship between Müllerian co-mimics that differ in their level of defence. There is now a growing awareness that this relationship can be mediated by many factors, including synergistic effects between co-mimics that differ in their mode of defence, rates of generalisation among warning signals and concomitant changes in prey density as mimicry evolves. I highlight areas for future enquiry, including the possibility of Müllerian mimicry systems based on profitability rather than unprofitability and the co-evolution of defence.     

Stenusine,an antimicrobial agent in the rove beetle genus <Emphasis Type="Italic">Stenus</Emphasis> (Coleoptera,Staphylinidae)

Stenusine is well known as the alkaloid, discharged by the rove beetle, genus Stenus Latreille (Coleoptera, Staphylinidae). The Stenus beetles employ the alkaloid as an escape mechanism when on water surfaces. In the case of danger, they lower their abdomen and emit stenusine from their pygidial glands. Stenusine shows a low surface tension and therefore a high spreading pressure; these properties propel the beetle quickly over the water. Many Steninae do not live in habitats with open waters, but in detritus, leaf litter, mosses, etc. This raises the possibility that stenusine might also have another function, e.g., as antibiotic or fungicide. Stenus beetles show an intense grooming behaviour. With gas chromatography-mass spectrometry analyses we could prove that they cover themselves with their secretion. To tests its antimicrobial properties we conducted agar diffusion tests with stenusine and norstenusine, another substance that is abundant in most Stenus species. Both compounds have an antimicrobial effect on entomopathogenic bacteria and fungi. Stenusine not only allows for an extraordinary method of locomotion on water surfaces, it also protects the Steninae from being infested with microorganisms.     

Costs of female odour in males of the parasitic wasp <Emphasis Type="Italic">Lariophagus distinguendus</Emphasis> (Hymenoptera: Pteromalidae)

The display of female traits by males is widespread in the animal kingdom. In several species, this phenomenon has been shown to function adaptively as a male mating strategy to deceive sexual rivals (female mimicry). Freshly emerged males of the parasitic wasp Lariophagus distinguendus (Hymenoptera: Pteromalidae) are perceived by other males as if they were females because of a very similar composition of cuticular hydrocarbons which function as a sex pheromone in this species inducing courtship behaviour in males. Within 32 h, however, males deactivate the pheromone and are no longer courted by other males. In this paper, behavioural experiments were performed to test hypotheses on potential costs and benefits associated with the female odour in young males. We did not find any benefits, but demonstrated that young males were significantly more often outrivaled in male-male contests when competing with two older males for a female. Also, young males were significantly more often mounted in homosexual courtship events during these contests. Thus, display of female traits by males is not necessarily beneficial, and in fact, can be disadvantageous. We suggest that these costs have favoured the evolution of the pheromone deactivation mechanism in L. distinguendus males. The function of cuticular hydrocarbons as a female courtship pheromone in L. distinguendus might have evolved secondarily from a primary function relevant for both genders, and the deactivation of the signal in males might have caused a shift of specificity of the chemical signal from the species level to the sex level.     

Middle-Eocene artiodactyls from Shanghuang (Jiangsu Province,Coastal China) and the diversity of basal dichobunoids in Asia

A new assemblage of basal dichobunoid artiodactyls from the middle-Eocene Shanghuang fissure fillings includes the diacodexeid Jiangsudon shanghuangensis gen. and sp. nov., a new species of the lantianine dichobunoid Elaschitotherium, Elaschitotherium crepaturus sp. nov., and an indeterminate suoid which is presently the earliest record of this clade. Diacodexeids are also represented by two forms provisionally referred to cf. Diacodexis sp. and to an indeterminate Diacodexeidae, respectively. The occurrence of diacodexeids in Shanghuang contrasts with the early and earliest middle-Eocene chronological range of the family in Europe and North America and suggests that the stratigraphic range of the family in Asia extends up to the middle Eocene. This may reflect particular habitats in coastal China that may have been relatively stable during the early and middle Eocene, thus preserving forest-dwelling artiodactyls that became extinct in the other Holarctic regions. Compared to other supposedly coeval North American, European, and Asian faunas, the Shanghuang mammalian assemblage is most similar to early Uintan faunas of North America but is also remarkable in recording forms close to taxa that are characteristic of the Wasatchian and Bridgerian North American Land Mammal Ages. The Irdinmanhan age of the Shanghuang fauna is supported by the mammalian assemblage recovered from the fissure D, but an Arshantan age cannot be completely ruled out at this point. Although the Shanghuang assemblage is biased towards preservation of small components of the mammalian fauna, the Shanghuang fauna provide an important and unique window into the Eocene diversity and early evolution of cetartiodactyls in eastern Asia.     

Interspecific mating of the introduced bumblebee <Emphasis Type="Italic">Bombus terrestris</Emphasis> and the native Japanese bumblebee <Emphasis Type="Italic">Bombus hypocrita sapporoensis</Emphasis> results in inviable hybrids

The bumblebee Bombus terrestris is not only an effective pollinator, but also a potential invasive alien species outside its native range. Recently, nearly 30% of queens of the Japanese native species Bombus hypocrita sapporoensis and B. hypocrita hypocrita were estimated to copulate with B. terrestris males in the field, suggesting that indigenous bumblebees could be genetically deteriorated through hybrid production with the introduced species. In this study, we evaluated hybrid production between the introduced B. terrestris and the indigenous B. hypocrita sapporoensis under laboratory conditions. The hatching rate of eggs derived from interspecific matings was 0% and 8.6% depending on the direction of the cross, which was significantly lower than that from intraspecific matings of B. terrestris (76.9%) and B. hypocrita sapporoensis (78.9%). Genetic studies using microsatellite markers revealed that both haploid and diploid individuals were present in the egg stage, whereas all hatched larvae were haploid. In addition, histological studies revealed that eggs derived from interspecific matings terminated development 2 days after oviposition. These results strongly suggested that eggs derived from interspecific matings are inviable due to post-mating isolation mechanisms. Mass release of exotic pollinators could cause serious population declines of native bumblebee species.     

Mantophasmatodea now in the Jurassic

The Mantophasmatodea is the most recently discovered insect order. The fossil records of all other ‘polyneopteran’ orders extend far in the past, but the current absence of pre-Cenozoic fossils of the Mantophasmatodea contradicts a long evolutionary history, which has to be assumed from the morphological distinctness of the group. In this paper, we report the first Mesozoic evidence of a mantophasmatodean from the Middle Jurassic of Daohugou, Inner Mongolia, China. Furthermore, the new fossil shares apomorphic characters with Cenozoic and recent Mantophasmatodea, suggesting a longer evolutionary history of this order.     

Within-population variation in ejaculate characteristics in a prolonged breeder,Peron’s tree frog, <Emphasis Type="Italic">Litoria peronii</Emphasis>

Sperm number is often a good predictor of success in sperm competition; however, it has become increasingly clear that, for some species, variation in probability of paternity cannot be explained by sperm number alone. Intraspecific variation in ejaculate characteristics, such as the number of viable sperm and sperm longevity, may play an equally important role in determining fertilization success. Here, we assess variation among ejaculates in three factors that may contribute to fertilization success (number of sperm per ejaculate, viability, and longevity), in a population of Peron’s tree frog (Litoria peronii). We detected large variation among males in the number of sperm per ejaculate and the proportion of viable sperm within ejaculates, which could not be explained by variation in either male size or body condition. However, the proportion of viable sperm released by males increased over the season. Finally, we assessed sperm longevity (proportion viable sperm determined using a dual-fluorochrome vital dye) at two different temperatures. At 23°C, on average, 75% of sperm remained viable after 2 h, but there were significant differences amongst males with the percentage of viable sperm ranging from 43% to 95%. For sperm incubated at 4°C, ejaculates varied fivefold in sperm longevity with some males having 50% viable sperm after 5 days. Our data suggest that ejaculate characteristics (sperm number, viability, and longevity) vary widely in Peron’s tree frog and may therefore play an important role in determining siring success both in the presence and absence of sperm competition. We discuss the results in relation to selection on ejaculate traits via natural and sexual selection in this and other amphibians.     

Why do <Emphasis Type="Italic">Manduca sexta</Emphasis> feed from white flowers? Innate and learnt colour preferences in a hawkmoth

Flower colour is an important signal used by flowering plants to attract pollinators. Many anthophilous insects have an innate colour preference that is displayed during their first foraging bouts and which could help them locate their first nectar reward. Nevertheless, learning capabilities allow insects to switch their colour preferences with experience and thus, to track variation in floral nectar availability. Manduca sexta, a crepuscular hawkmoth widely studied as a model system for sensory physiology and behaviour, visits mostly white, night-blooming flowers lacking UV reflectance throughout its range in the Americas. Nevertheless, the spectral sensitivity of the feeding behaviour of naïve moths shows a narrow peak around 450 nm wavelengths, suggesting an innate preference for the colour blue. Under more natural conditions (i.e. broader wavelength reflectance) than in previous studies, we used dual choice experiments with blue- and white-coloured feeders to investigate the innate preference of naïve moths and trained different groups to each colour to evaluate their learning capabilities. We confirmed the innate preference of M. sexta for blue and found that these moths were able to switch colour preferences after training experience. These results unequivocally demonstrate that M. sexta moths innately prefer blue when presented against white flower models and offer novel experimental evidence supporting the hypothesis that learning capabilities could be involved in their foraging preferences, including their widely observed attraction to white flowers in nature.     

Ventral polarization vision in tabanids: horseflies and deerflies (Diptera: Tabanidae) are attracted to horizontally polarized light

Adult tabanid flies (horseflies and deerflies) are terrestrial and lay their eggs onto marsh plants near bodies of fresh water because the larvae develop in water or mud. To know how tabanids locate their host animals, terrestrial rendezvous sites and egg-laying places would be very useful for control measures against them, because the hematophagous females are primary/secondary vectors of some severe animal/human diseases/parasites. Thus, in choice experiments performed in the field we studied the behavior of tabanids governed by linearly polarized light. We present here evidence for positive polarotaxis, i.e., attraction to horizontally polarized light stimulating the ventral eye region, in both males and females of 27 tabanid species. The novelty of our findings is that positive polarotaxis has been described earlier only in connection with the water detection of some aquatic insects ovipositing directly into water. A further particularity of our discovery is that in the order Diptera and among blood-sucking insects the studied tabanids are the first known species possessing ventral polarization vision and definite polarization-sensitive behavior with known functions. The polarotaxis in tabanid flies makes it possible to develop new optically luring traps being more efficient than the existing ones based on the attraction of tabanids by the intensity and/or color of reflected light.