Immune defence,a possible nonvisual selective factor behind the industrial melanism of moths (Lepidoptera)

It is generally believed that industrial melanism in Lepidoptera is mainly caused by differential predation by birds. In polluted areas, melanic individuals are favoured by natural selection because they are better camouflaged than pale moths on lichen‐free and sooty tree trunks. In this article, we show that, in the black arches moth (Lymantria monacha), melanic morphs have a stronger encapsulation response than pale morphs against nylon monofilament implants. This indicates that the melanic and pale morphs differ in the strength of their immune defence. The same chemical precursors and their end product, melanin pigment, are involved in the encapsulation response and in the external coloration. Thus, it seems that there may be two possible, not mutually exclusive, explanations for the frequency changes observed in the industrial melanism of moths. The dominant gene causes an increase in the amount of melanin pigment and its precursors. This increase causes two changes: an intensified immune defence as a form of improved encapsulation ability of foreign objects, and the well‐known protective dark coloration (a case of relational pleiotropy). It seems possible that industrial melanism is a by‐product of selection on the strength of immunity. In the field, these pleiotropic aspects are exceedingly difficult to distinguish from each other, and the factors may even be compensatory. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 831–838.     

The understanding of industrial melanism in the peppered moth (Biston betularia) (Lepidoptera: Geometridae)

Melanic polymorphism in B. betularia has been extensively studied. Correlations between high melanic frequency and high levels of air pollution have been demonstrated. Kettlewell and others have shown that differential bird predation has an important effect on the maintenance of the polymorphism, and coefficients of visual selection have been obtained on the assumption that the moth habitually rests on tree trunks. Computer models based on these selective coefficients show that they are not sufficient accurately to explain observed melanic frequencies. Other non-visual selective factors and weak frequency-dependent selection have been invoked to improve fits. Analysis of the resting positions of moths recorded in the wild demonstrates that B. betularia does not usually rest in exposed positions on tree trunks, but rather rests on the underside of branches, on trunks in shaded positions just below major branch joints or on foliate twigs. The results of a pilot selection experiment, while agreeing qualitatively with Kettlewell's results, suggest that fitness estimates that assume trunk-resting are quantitively incorrect. The error is greatest for melanic moths in rural areas. It is suggested that visual selective coefficients based on a true assessment of the resting behaviour of the moths may considerably improve the fit between computer predictions and observed phenotype frequency distributions.     

Background selection by the peppered moth (Biston betularia Linn.): individual differences

The hypothesis that dimorphically coloured, cryptic moths select appropriate rest sites by comparing their body scales to substrate reflectance was tested using typical and melanic morphs of the peppered moth, Biston betularia (L.). Experiments designed to block the individual's inspection of its inherited colour phenotype do not support Kettlewell's contrast/conflict (self-inspection) hypothesis. Instead, tracking of marked moths over successive days revealed individual differences in rest-site selection which were not related to treatments, experience (imprinting), nor closely to a moth's inherited colour pattern. Differences between family broods indicate that some genetic bias in background selection exists. The production of artificially selected lines with consistent but opposing preferences will allow us to investigate the co-evolution of pattern and behaviour.     

Behavioural studies on the peppered moth Biston betularia and a discussion of the role of pollution and lichens in industrial melanism

We examined the resting behaviour during pairing and egg-laying of bred female Biston betularia (L.) after they were set free at wooded sites in Somerset and Cardiff. Moths were not released immediately on emergence but were held for three nights to overcome an initial period of flight activity. Females (N= 257) were placed on trunks, and on large and small branches of trees, especially oaks, over periods of several weeks in 1984, 1985 and 1986. Releases were not made within 50 m of any known survivor. One hundred and thirty-one pairings were observed. We also examined oviposition behaviour in a cage experiment using different types of branches. Our observations support Mikkola's earlier conclusion from cage experiments with male moths that the species rests predominantly on branches and shows an appropriately specialized resting attitude, demonstrated here in a series of photographs. Many moths will rest underneath, or on the side of, narrow branches in the canopy. Once released females had settled and paired they only moved quite short distances, but positioning and egg-laying were profoundly influenced by the presence of foliose lichens. Females of all phenotypes show a strong preference to oviposit beneath a thallus of foliose lichens. Their tendency to rest against or close to the lichen during the day could markedly influence their crypsis. In the absence of foliose lichens, eggs are laid in cracks in the bark. The survivorship of released females both in different pairing combinations and during their whole reproductive life-span is analysed. These preliminary data suggest that bird predation can give rise to differential mortality of different pairing combinations and of the phenotypes. This technique provides a powerful method of measuring the relative crypsis and differential mortality of phenotypes with living females in natural resting sites. The significance of our observations on resting behaviour is discussed with regard to the role of air pollution and epiphytes in industrial melanism. Particular attention is given to the causal mechanisms underlying recent declines in the frequency of carbonaria melanics in parts of Britain. One important factor is the new growth made by trees since the marked reduction in particulate air pollution in the 1960s. It is emphasized that we need to know much more about the interactions between pollution, epiphytes and resting backgrounds, especially in the canopy, before we can be confident of our understanding of the evolution of industrial melanism.     

甘肃酒泉苹果蠹蛾的发生规律

苹果蠹蛾Cydia pomonella(L.)是我国重要的果树害虫和检疫对象。本文通过田间调查和性诱芯诱捕器诱捕的方法,研究了2008年苹果蠹蛾在甘肃酒泉的年发生规律和空间分布特性。结果表明,苹果蠹蛾在我国甘肃酒泉1年发生2代;越冬代和第1代成虫发生高峰期分别为5月上旬至中旬和7月上旬至中旬,成虫在树冠的中层、向阳的方向(东部和南部)、果实和叶背面产卵较多;幼虫发生的两次高峰分别发生在6月17日和7月26日,刚孵化幼虫很快蛀果隐蔽为害,给防治带来很大困难;幼虫化蛹前,主要从树的上部向下爬行,并常以老熟幼虫潜入树干或分枝的树皮下结茧化蛹或越冬,次年4月中下旬开始羽化。     

Shell colour polymorphism in a mangrove snail Littorina sp. (Prosobranchia: Littorinidae)

Shell colour polymorphism was examined in populations of a mangrove snail Littorina sp. in Queensland, Australia. Three morphs were recognized, yellow, red and brown, and morph frequencies varied both between widely spaced populations and between islands less than 1 km apart. Morph frequencies also varied with time of year. There was a relationship between shell colour and position on the tree, with yellow snails more often occurring amongst the foliage and brown snails more often on trunks and branches. In some populations yellow snails appeared to survive better than the other morphs, while in other populations there was no difference. The evidence for the maintenance of the polymorphism by natural selection is discussed.     

Seasonal polyphenism and leaf mimicry in the comma butterfly

The comma butterfly, Polygonia c-album, exhibits seasonal polyphenism with a darkish winter morph and a lighter summer one. Phylogenetic analysis suggests that the winter morph represents the ancestral condition. We suggest two hypotheses for the evolution of the summer morph and the maintenance of seasonal polyphenism in the comma: (1) that the summer morph is better protected against predation on summer roost sites, whereas the winter morph is better protected on hibernation sites, and (2) that the summer morph is energetically less expensive and results from deallocation of resources from soma (e.g. dark wing pigmentation) to reproduction. We tested the antipredation hypothesis in experiments using great tits, Parus major, as predators on winter and summer morph commas presented simultaneously on tree trunks or on nettles. However, this hypothesis was not supported as the winter morph was better protected than the summer morph on both backgrounds. Predation when both morphs were present was lower on nettles, and summer morphs placed in exposed positions on tree trunks outdoors disappeared sooner than winter morphs placed on the same background. In addition, in a final experiment, 18 summer morphs released in their natural habitat in the evening exclusively chose leaves for roost sites, whereas 12 of 19 winter morphs chose a tree trunk, branch or twig. We conclude that evolution of the summer morph is consistent with the life history hypothesis and that its choice of summer roost sites is associated with a low predation pressure.     

Natural selection and the distribution of shell colour morphs in three species of Littoraria (Gastropoda: Littorinidae) in Moreton Bay, Queensland

The distribution of yellow, brown and red morphs of sympatric species of Littoraria were recorded on mangrove trees of the genus Avicennia within Moreton Bay, Queensland. The roles of background mimicry (leaf vs. bark, dark vs. light), niche selection and thermal tolerance (sunny vs. shaded positions and height above ground) were examined. The yellow advantage found previously in the area was tested. Total yellow morph frequency adjusted to a reduction in leaf background on pruned trees. Morph frequencies in Littoraria species reflect differences in habitat use. L. filosa (high yellow frequency) was more frequently found on leaves at the highest tree levels, while L. luteola (high brown frequency) was more frequently found on branches at lower levels. It is therefore argued that morphs mimic background elements. Previously reported niche selection by yellow and brown morphs of leaf and bark backgrounds is shown to be a result of the distribution of L. luteola on branches and L. filosa on leaves. At warmer times of the year, yellow L. filosa were more common in sunny positions; this is thought to be a result of thermal tolerance. There appears to be some advantage to particular morphs on particular tree types, but this relationship needs to be examined further. Mangrove-dwelling Littoraria are a promising model to investigate molluscan polymorphism. In the past, erroneous identification of sympatric species may have influenced the accuracy of reported patterns. We used allozyme electrophoretic markers as a precise identification technique. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 75 , 219–232.     

Movement of plum curculio adults toward host trees and traps: flight versus walking

Adult plum curculios, Conotrachelus nenuphar (Herbst), were dislodged from branches of apple trees by tapping on 14 days between blossom petal fall and 3 weeks thereafter. They fell onto a small square of framed white cloth that was quickly but carefully moved to a position beneath or adjacent to the canopy of another apple tree.There, adults were observed for 50-min periods from 0800–1950 h for propensity to fly from or walk off the cloth or move to hide beneath foliage on the cloth. At each position, the mean time from observation initiation until flight, walking off or hiding did not differ significantly among these behaviors (averaging 21–30 min), but significantly more flew (31–40%) than crawled off (16–19%) or hid (18–20%). Substantial walking off but no flights occurred at air temperatures below 20 °C, whereas at temperatures of 20 °C or greater, propensity for flight was usually significantly greater than that for walking off.Flights were predominantly toward the tree canopy or inter-tree space, with only a small proportion (17% or less) toward the tree trunk or (in a parallel study) toward an unbaited black pyramid trap (mimicking a tree trunk). Walking was overwhelmingly toward the tree trunk or a black pyramid trap adjacent to the trunk. Our observations lead us to conclude that unbaited black pyramid traps next to trunks of host trees (the most attractive position) can be effective at air temperatures below 20 °C by attracting plum curculio adults that would have entered host trees by walking toward and up tree trunks.Such traps appear to be less effective at 20 °C or greater because under such conditions, adults tend to enter host trees by flight into the canopy.     

An experimental investigation of the behaviour and mortality of artificial and natural morphs of Cepaea nemoralis (L.)

Two population cage experiments which examined the mortality and behaviour of artificial and natural morphs of the landsnail C. nemoralis ( L .) are described. In the first experiment artificial 'morphs' were manufactured from one genetically homogeneous founder group of snails by painting the shells black or white. Differences in behaviour and mortality between these two 'morphs' were observed during the course of the experiment. Daylight and weather conditions were important in determining the activity patterns of the two types of snail.
In the second experiment naturally occurring brown and yellow colour morphs were used. These exhibited very similar behaviour patterns to those of their artificial 'mimics' and it is concluded that phenotype must play an important role in determining the behavioural responses of the snails to their environment. Thermal relationships are suggested as causal factors for the differences between the experimental morphs.     

Evolutionary dynamics of declining melanism in the peppered moth in the Netherlands

Populations of Biston betularia in the region of The Netherlands around Leiden and Rotterdam were resampled. A comparison of three sets of data for 1969-1973, 1988 and 1999 enabled a further examination of declines in melanism. Unlike parallel changes for the black carbonaria form of this species in urban regions of Britain, those inThe Netherlands involve substantial changes in frequencies of at least two of the intermediate insularia morphs as well as an increase in the non-melanic typica morph. The darkest of the three insularia morphs has shown a transitory pulse of increased frequency in The Netherlands. The dynamics are discussed in relation to the history of air pollution and to straightforward predictions about selection.     

Selective bird predation on the peppered moth: the last experiment of Michael Majerus

Colour variation in the peppered moth Biston betularia was long accepted to be under strong natural selection. Melanics were believed to be fitter than pale morphs because of lower predation at daytime resting sites on dark, sooty bark. Melanics became common during the industrial revolution, but since 1970 there has been a rapid reversal, assumed to have been caused by predators selecting against melanics resting on today's less sooty bark. Recently, these classical explanations of melanism were attacked, and there has been general scepticism about birds as selective agents. Experiments and observations were accordingly carried out by Michael Majerus to address perceived weaknesses of earlier work. Unfortunately, he did not live to publish the results, which are analysed and presented here by the authors. Majerus released 4864 moths in his six-year experiment, the largest ever attempted for any similar study. There was strong differential bird predation against melanic peppered moths. Daily selection against melanics (s ≈ 0.1) was sufficient in magnitude and direction to explain the recent rapid decline of melanism in post-industrial Britain. These data provide the most direct evidence yet to implicate camouflage and bird predation as the overriding explanation for the rise and fall of melanism in moths.     

Differences in the iliolumbar ligament and the transverse process of the L5 vertebra in young white and black people

The anatomy of the iliolumbar ligament (ILL) and the spatial orientation of the transverse process of the L5 vertebra were studied in 62 young black (n = 29) and white (n = 33) men and women during routine autopsy. The aim of the study was to determine possible racial differences in the structure and attachments of the iliolumbar ligament. The present study also investigated the spatial orientation of the transverse process of the L5 vertebra since the ILL has been reported to attach to the transverse process. The measurements of the iliolumbar ligament were carried out with a digital vernier caliper while the transverse process angles were measured with an adjustable protractor. The ligament in black people was made up of a single, markedly longer band compared to white people, where the ligament was made up of two shorter bands. The ILL measured 61.8 +/- 1.3 mm in black and 33.2 +/- 1.5 mm white men, and in black women 61.3 +/- 0.9 mm versus 32.2 +/- 1.2 mm in white women (p <0.01). Further, the ILL was markedly wider in black than white subjects (p <0.01). The horizontal and vertical angle also varied greatly between black and white subjects (p <0.01). These are previously unrecognized observations. Albeit unsubstantiated, these findings may have implications for understanding the etiology of various low back stress problems.     

Web-building and predatory behaviour of Spartaeus spinimanus and Spartaeus thailandicus, primitive jumping spiders (Araneae, Salticidae) from South-east Asia

Behavioural data are provided for the first time for two species of Spartaeus, S. spinimanus from Singapore and Thailand and S. thailandicus from Thailand. These jumping spiders (Araneae, Salticidae), from the primitive subfamily Spartaeinae, build large sheet webs on tree trunks and they capture prey—especially moths—on or near the web by walking to and lunging at them. When walking, Spartaeus waves its palps and legs in an unusual way. In formal tests, Spartaeus captured moths more frequently than flies and captured both moths and flies more readily when with their webs than without their webs.     

Experiments on resting site selection by nocturnal moths

When different species of moths are presented with a choice between black and white resting backgrounds, there is a strong correlation between the colour selected and the reflectance of the forewings. Under more natural conditions, light-coloured moths usually rest on fresh vegetation whilst dark-winged species select tree bark or rest upon the ground, and different defensive strategies appear to have been adopted by species in these two latter situations. Studies on the mechanism of background selection, and on background selection in polymorphic species, are reviewed.     

Nest position,breeding success and diet of the Black-crowned Night Heron,Nycticorax nycticorax in the Anzali Wetland,Northern Iran (Aves: Ardeidae)

Breeding ecology of the Black-crowned Night Heron (Nycticorax nycticorax) was studied in a mono-specific colony in the Anzali wetland, Northern Iran during the breeding season of 2016. The breeding period lasted from mid-May to late July. The average clutch size was 3.1±0.6 eggs and the breeding success 77.6%. No significant differences were found between nests built on trunks and those built on branches of trees. The clutch size and breeding success appeared to be independent of the structural variables of the nesting site (diameter of trees, height from the ground, height of nests from the canopy, nest number per tree, location of nests on trunks and branches). No significant difference was found between the timing of the start of incubation and the height of nests above the ground. The average vertical and horizontal distances between nests was one metre. Fish, particularly Carassius gibelio, dominated the diet of the nestlings.     

The hydraulic architecture of balsam fir (Abies balsamea)

Leaf-specific conductivities (LSCs – hydraulic conductivity per dry weight of supplied leaves). Huber values (transverse sapwood area per dry weight of supplied leaves), specific conductivity (hydraulic conductivity per transverse sapwood area) and tracheid diameters were measured throughout the trunk and crown of 20-year-old trees of Abies balsamca (L.) Mill. Measured specific conductivity was proportional to the radius to the fourth power of tracheids. LSCs, which indicate the relative water availability to different plant parts, are much higher in the trunk than in first order branches, and lowest in second order branches. The structural basis for this "hydraulic hierarchy" lies both in Huber values and in tracheid diameters. For similar diameter stem segments, there was no statistically significant difference for trunks versus branches in specific conductivity. However, in old parts of the tree, trunks are wider than supported branches and producer wider tracheids resulting in greater specific conductivities than in branches. In vigorous trees with strong apical control, Huber values were 12.0 times greater in the trunk than in similar diameter branch segments. In slow-growing trees with weak apical control, Huber values were 2.2 times greater in the trunk versus similar branch segments.     

Hanging in the treetops: an in situ experiment in ancient hemlock assessing outbreak defoliator performance among crown levels

1. Past studies have shown that both egg lay and larval feeding of a generalist defoliator, pale‐winged gray (Iridopsis ephyraria Walker) (Lepidoptera: Geometridae), are concentrated in the mid‐lower crown of eastern hemlock [Tsuga canadensis (L.) Carr.]. 2. Using tree‐climbing techniques, we carried out in situ bioassays in large hemlock trees (～25 m) to determine whether the observed foraging preferences are adaptive and how they reflect associated intra‐tree variations in microhabitat quality associated with temperature and foliage nutritional quality. 3. In 1 of 2 years, larval survival was significantly higher in the shaded lower versus sunlit upper crown; however, in both years, groups that fed in the sunlit upper crown branches had larger male and female moths and more female‐biased sex ratios. 4. Differences in pale‐winged gray performance among crown levels were somewhat supported by trends in foliage nutritional chemistry but not well‐correlated with variation in temperature. 5. The present study is one of only a few to carry out bioassays on large mature trees and results reinforce the idea that foraging patterns of herbivores are likely to reflect trade‐offs among several factors that vary between sunlit and shaded branches within forest canopies.     

Tests of hypotheses on predation as a factor maintaining polymorphic melanism in coastalplain fox squirrels (Sciurus niger L.)

Fox squirrels ( Sciurus niger ) in the south-eastern U.S. coastal plain differ from those in the rest of the species' range by having black heads with white nose and ears. Postcranially on the dorsum, they also show interindividual colour variation, ranging from all-light agouti to all-black non-agouti. I present results of experiments undertaken to test whether the evolution of these pigmentary features can be attributed to interactions with predators. Comparative static crypsis (when the squirrels are still) was tested by determining how well specimens of the squirrel morphs matched their backgrounds in terms of intensity (brightness) distributions and patch size. Comparative dynamic effects (when the squirrels are moving) were tested using captive red-tailed hawks ( Buteo jamaicensis ) and models of the fox squirrel morphs. Results indicated that morphs of coastal-plain fox squirrels with all-light backs are better static matches to unburned backgrounds than are the darker morphs with which they coexist. Against fire-blackened backgrounds, fox squirrel morphs with intermediate and all-black backs are better intensity matches than those with all-light backs, but the dark morphs are not better patch-length matches than light morphs. The superiority of dark morphs in intensity matching for a short time after a fire does not seem sufficient to account for their long-term maintenance. Hawks responded more slowly to moving fox squirrel models with intermediate amounts of black on the back than to all-light or all-dark morphs; this result suggests a possible factor that would favour retention of genes for dorsal blackness in the coastal-plain population. Patchy black and white heads appear to promote static crypsis of south-eastern fox squirrels, and hawks reacted more slowly to moving squirrel models with such colouration than to those with plain heads.     

Effect of papaya trunk angle on infestation by white peach scale,Pseudaulacaspis pentagona (Hemiptera: Diaspididae)

Two papaya (Carica papaya L.) seedlings growing in one planting hole often results in angular or non-vertical growth of the trees. Data on trunk angularity, or leaning, (deviation from the vertical line of reference) and white peach scale, Pseudaulacaspis pentagona Targioni-Tozzetti (Hemiptera: Diaspididae), densities on paired papaya trees were collected approximately 1 year after infestation of a papaya field was discovered. Paired trees showed a significantly higher degree of leaning than single trees. White peach scale densities were significantly higher on tree trunks with a greater departure from vertical in paired comparisons. Therefore, paired tree planting practices may facilitate the development of economic infestations of white peach scale populations in papaya orchards.