The colour/pattern polymorphism of <Emphasis Type="Italic">Philaenus spumarius</Emphasis> (Hemiptera: Cercopidae) in the north-western Black Sea region of Turkey

The heritable elytral colour/pattern polymorphism of Philaenus spumarius in north-western Black Sea region of Turkey was investigated. A total of 2,530 adult spittlebugs, 1,020 females and 1,510 males were collected and analysed in terms of the 11 most common colour forms. Nine different phenotypes, expressed by six different alleles, were determined: populi (POP), typicus (TYP), trilineatus (TRI), marginellus (MAR), flavicollis (FLA), leucocephalus (LCE), gibbus (GIB), albomaculatus (ALB), and leucophtalmus (LOP). The common phenotypes lateralis (LAT) and quadrimaculatus (QUA) of previous studies were not encountered at all. The predominant phenotype was TYP (63%) in females and males followed by POP (31%). TRI (4%) and melanics (5%) were at close proportions. However, only one site had nine phenotypes altogether. The majority of the sites had only the POP and TYP morphs. The melanic forms were found only in females.     

Ventral pigmentation variation in the meadow spittlebug Philaenus spumarius (L.) (Hemiptera,Cicadomorpha: Cercopidae)

The meadow spittlebug Philaenus spumarius (L.) (Hemiptera, Cicadomorpha: Cercopidae) exhibits heritable colour/pattern variation on the dorsal and ventral sides of the adults. Eleven principal dorsal morphs are grouped as melanics and non-melanics according to the pigmentation of the dorsal surface. An investigation of ventral pigmentation variation in Philaenus spumarius was carried out with laboratory breeding stock obtained from Llysdinam, Wales. Analyses of the 1552 individuals, 753 females and 799 males, indicated that the pigmentation on the ventral surface of the adult individuals which vary significantly is associated with dorsal morph. Ventral parts of the dorsal melanics are usually darker than in non-melanics in both females (Kruskal-Wallis one-way ANOVA: H = 372.70, d.f. = 7, P < 0.001) and males (H = 407.36, d.f. = 7, P < 0.001). The combined “C” group morphs (flavicollis + gibbus + leuco- cephalus) are always darker than all other morphs in both sexes.     

Limited polymorphism in two spittlebugs,Philaenus spumarius (Linnaeus) and P. signatus Melichar (Hemiptera: Cercopidae), in island populations from Western Turkey

Two colour polymorphic spittlebug species, Philaenus spumarius and Philaenus signatus were recorded for the first time in Gökçeada, Turkey. Three non-melanic – POP (populi), TYP (typicus), and VIT (vittata) – and three melanic MAR – (marginellus), FLA (flavicollis), and LCE (leucocephalus) – colour/pattern phenotypes of P. signatus were found. Philaenus spumarius was represented only by POP and TYP, which are expressed by a single pigmentation allele. The other six alleles were absent on this island, although they are found in most populations throughout the world. Stochastic events such as random genetic drift may be the reason for shaping the polymorphism of these two sympatric species in Gökçeada. Thus, the absence of pigmentation alleles possibly indicates the founder effect.     

Genetic Basis of Heterogeneity for Morphological and Behavioral Characters in a Population of Archips podanaScop. (Lepidoptera: Tortricidae)

Inheritance of morphological characters related to male and female genitalia was examined in Archips podanaScop. The position and number of aedeagus prongs had a monogenic diallelic mode of inheritance. The gene determining this character is autosomal. Homozygous and heterozygous males have respectively one and two prongs. In contrast to homozygous for P ¹(prong A) and heterozygous (prong A+B) males, males homozygous for P ²(prong B) were characterized by much more active courtship. The gene correlates with a set of characters controlling individual ecology, exploring behavior of females, and feeding on different cultures. Morphological adult type 0 is an extremely weak form of phenotype B. Exploring activity of adults having phenotype B is the highest at the ratio of cis-11-tetradecenol to the other components of the female sex pheromone that leads to a somewhat lower activity of males of other phenotypes. In females, no regularities in inheritance of phenotypic characters were revealed. A hypothesis on a mechanism for maintenance of population stability for the given phenotypes is advanced and a possible mechanism for microevolution of a polymorphic species is proposed.     

Genetic control of colour /pattern polymorphism in British populations of the spittlebug Philaenus sputnarius (L.) (Homoptera: Aphrophoridae)

The results of breeding experiments designed to determine the genetic control of the colour/pattern polymorphism in Philaenus spumarius in certain British populations are described and contrasted to previously published results for material taken from populations in southern Finland. The 11 principal phenotypes are controlled by seven alleles at a single autosomal locus, with complex patterns of dominance and co-dominance. In Finland, and most of the rest of the species' extensive range, the eight melanic morphs are wholly or substantially limited to the females, in which they are dominant to the typical form. Previously, this sex limitation has been explained in terms of reduced penetrance of melanic alleles in males and/or dominance reversal between the sexes, such that the eight melanic phenotypes (controlled by five alleles which are adjacent in the dominance heirarchy) are dominant to typical in the females but recessive in the males. Published data on morph frequencies in S Finland and on breeding experiments using material collected from these populations are re-examined and it is shown that neither theory accounts adequately for all the evidence. By contrast, in most British populations there is greatly increased penetrance in males of the melanic morphs which are strictly female-limited elsewhere. Breeding experiments, using P. spumarius taken from selected British populations, provide clear evidence that melanics are dominant to typical in both sexes. This difference in genetic control is entirely consistent with the balance of morph frequencies between the sexes in the populations from which the breeding material for the two separate studies was taken. Experimental material for this study was taken from populations in the Cynon Valley in south Wales, which have the highest melanic frequencies in the species' range and show particularly clear expression of melanic phenotypes in males. Both of these features are strongly correlated with severe aerial pollution from a local smokeless fuel factory. Selection for melanism in these populations is clearly strong, which probably accounts for the high level of penetrance and the dominance of melanics to typical in both sexes. Although further experiments are needed, it is suggested that regulatory loci, controlling the penetrance and expression of melanic alleles in males, could account for the observed differences in genetic control between populations in Britain and Finland.     

Genetic diversity of Philaenus spumarius and P. tesselatus (Hemiptera,Aphrophoridae): implications for evolution and taxonomy

The meadow spittlebugs Philaenus spumarius and P. tesselatus are closely related taxa with uncertain taxonomic position in the light of previous morphological, ecological, cytological and molecular research. Despite morphological homogeneity of P. spumarius (with the exception of high colour and pattern polymorphism) across its wide Holarctic range, it is possible that additional taxa (species or subspecies) exist. Philaenus spumarius is a potentially important pest in parts of its range where it was introduced. We used DNA markers to describe the genetic diversity of P. spumarius and P. tesselatus and to verify the taxonomic status of P. tesselatus and remote populations of the former species. The mitochondrial (cytochrome B) data showed that there are two main groups encompassing a northeastern (Asia and north‐central Europe) and a southwestern (Mediterranean area and western Europe, including North American specimens) clade. According to the elongation factor‐1α gene, there are three main clades: northeastern (Eurasiatic clade, E1), southeastern (east Mediterranean – Caucasus clade, E2) and southwestern (Iberian clade, E3). These two or three mitochondrial and nuclear clades could be considered as separate taxonomic units. On the other hand, all studied individuals of both species possessed the same internal transcribed spacer 2 haplotype. American specimens most probably originated from some western European populations. All studied specimens of P. tesselatus belong to the southwestern clade and western Mediterranean cluster. Therefore, together with cytological data, its species status is doubtful. To definitively solve the taxonomic status of P. tesselatus and populations of P. spumarius, further research using more samples and more genetic markers are needed.     

Influence of diet and photoperiod on development and reproduction of European populations of Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae)

The current study examines the effect of photoperiod (16:08 or 12:12 h L:D) and diet (eggs of Ephestia kuehniella Zeller (Lepidoptera: Phycitidae) or the pea aphid Acyrthosiphon pisum (Harris) (Hemiptera: Aphididae)) on the development and reproduction of the multicoloured Asian lady beetle Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae). A long-term laboratory population of H. axyridis (since 1998) and a melanic and non-melanic population originating from field collected individuals of H. axyridis in Belgium were used in this study. Long day conditions (16 h photoperiod) shortened development of the field populations with 2–3 days when compared with short day conditions (12 h photoperiod). Oviposition in the field populations was delayed by 1–3 months when reared at a 12 h photoperiod. Dissections indicated that the females were in reproductive diapause. As compared with live pea aphids, a diet consisting of E. kuehniella eggs yielded heavier adult body weights (up to 12%) and increased the number of egg laying days (by 45–169%) for both field populations at a 16 h photoperiod and lengthened adult life span (by 45–92%) under both light regimens. The morph types differed in their response to the foods offered in terms of developmental rate, pre-oviposition period and number of oviposition days. The laboratory and field strains responded differentially to regimens of food and photoperiod. The study indicated a greater nutritional plasticity of the non-melanic morphs which may offer them a competitive advantage that may in part explain the predominance of non-melanic morphs in newly colonized areas.     

New data on the nature and origin of colour polymorphism in the spittlebug genus Philaenus (Hemiptera: Aphorophoridae)

In Mediterranean populations of Philaenus colour polymorphism is maintained mainly by selection taking place during the long aestivation period, when adults are forced to migrate from their oviposition and larval hosts to other plants. Melanic morphs are more frequent in the south and also non-melanic morphs of P. spumarius are more strongly pigmented in the Mediterranean than in more northern populations. Industrial melanism is not indicated. Colour polymorphism is correlated with the phylogenetic relationships of the species. The possible origin of polymorphism is discussed in comparison to the theory of Halkka & Halkka (1990). Suggestions are made about the geographic area where further studies on the polymorphism should be focused.     

Temperature and the colour polymorphism of Philaenus spumarius (Homoptera: Aphrophoridae)

ABSTRACT.

• 1 The spittle bug, Philaenus spumarius (L.), has eleven distinct elytral colour pattern morphs in Britain, of which three are designated non-melanic and eight melanic. Frequencies of different morphs vary between populations.
• 2 The role of thermal factors in the maintenance of the balanced polymorphism was investigated by measurements of dorsal reflectance and body temperature when artifically irradiated.
• 3 Non-melanic forms had higher reflectances and showed lower temperature excesses when irradiated. There was considerable variation in reflectance for the populi–typicus morph group, and trilineatus is therefore the only true‘non-melanic’morph.
• 4 For samples along altitudinal transects in three Scottish glens, trilineatus frequencies were found to be negatively and populi-typicus and melanic frequencies positively correlated with altitude.
• 5 Thermal considerations do therefore play some role in the broad and complex selection regime of P.spumarius.

     

Differential survival and reproduction in colour forms of Philaenus spumarius give new insights to the study of its balanced polymorphism

1. Colour polymorphisms are common across animals and are often the result of complex selection regimes. Philaenus spumarius (Linnaeus) (Hemiptera, Aphrophoridae) shows a widely studied dorsal colour polymorphism with several described phenotypes whose variation in their occurrence and frequency, as well as their maintenance across time, have been reported. Several selective influences have been suggested to play a role, but the mechanisms underlying the maintenance of this polymorphism are still poorly understood. 2. To explore the adaptive significance of the colour polymorphism of P. spumarius, an experiment was conducted in captivity under semi‐natural conditions to measure survival, reproductive success, and duration of egg maturation. 3. It was found that there was higher longevity, a higher number of oviposition events, and a higher number of eggs laid by trilineatus phenotype females than by typicus and marginellus, supporting previous reports of an increase in trilineatus frequency during the season. The duration of egg maturation did not differ among phenotypes. 4. The higher longevity and fertility of the trilineatus phenotype may compensate, for example, the higher rate of attack by parasitoids and/or higher solar radiation reflectance in this phenotype, which have already been reported in previous studies, constituting a possible mechanism for the maintenance of this polymorphism.     

Variation in the response of <Emphasis Type="Italic">Mimulus guttatus</Emphasis> (Scrophulariaceae) to herbivore and virus attack

Plants are often simultaneously attacked by insect herbivores and plant pathogens, yet relatively few studies have investigated the potential interactive effects of herbivores and pathogens on plant fitness. We studied the effects of simultaneous attack by meadow spittlebugs, Philaenus spumarius (Homoptera: Cercopidae), and a plant virus, cucumber mosaic virus (CMV), on the fitness of Mimulus guttatus (Scrophulariaceae). We wanted to determine: (1) if trade-offs in defense against meadow spittlebugs and CMV exist, (2) if meadow spittlebugs and CMV interact to affect plant fitness, and (3) if genetic variation is associated with these interactive effects. We found no evidence of trade-offs in defense against meadow spittlebugs and CMV in M. guttatus in a greenhouse experiment. Thus, the ability of M. guttatus to defend itself against one of these enemies is unlikely to preclude the evolution of adequate defenses against the second enemy. We did, however, find strong evidence that spittlebugs and CMV interacted to affect plant fitness and that genetic variation underlies the degree and direction of this interaction. This suggests that selection can act on the genetic variation underlying the interaction between the two enemies and that strong selection imposed by one will alter the response of M. guttatus populations to the second. To our knowledge this is the first study to demonstrate genetic variation associated with the non-additive effect of an herbivore and a pathogen on plant fitness. We suggest that future studies of the mechanisms underlying the defensive properties of plants need to consider variation associated with defense mechanisms and the potential effect of this variation on the response of plant populations to selection by multiple enemies.Co-ordinating editor: Steufer     

Vibrational playbacks and microscopy to study the signalling behaviour and female physiology of Philaenus spumarius

The meadow spittlebug Philaenus spumarius, vector of the bacterium Xylella fastidiosa, relies on vibrational communication to accomplish mating: the female calls to establish a duet with a male. A deeper knowledge of the species’ reproductive biology and behaviour would provide useful information for developing control techniques based on principles of ‘biotremology’, which studies the vibrational behaviour of animals. Playback tests were conducted on single females and male–female pairs of P. spumarius from June to October 2018, and the features of the recorded calling signals were analysed using a wavelet decomposition. Dissections were performed on females to evaluate the relationship between calling activity and ovarioles development. From August onwards, females started to emit calling signals and to develop ovarioles. Female calling activity, duration of their chirps and their responsiveness to mating increased as the season progressed, and they were correlated with ovarioles’ development and presence of mature eggs. Hence, the ovarian maturation represents a key factor in association with the development of the sexual behaviour of P. spumarius females. Conversely, males produced advertisement signals soon after adult eclosion in May, but these signals were not involved in the pair formation process. Mating was achieved only when males produced courtship signals in response to female calling signals and established with them vibrational duets starting from August. Here, we provide new information regarding the P. spumarius’ ethology and hypothesize that potential mating disruption techniques should consider the insect physiology and be applied when both sexes are responsive to mating signals.     

Intrapopulation gender variation in common ragweed (Asteraceae: Ambrosia artemisiifolia L.), a monoecious,annual herb

Summary We measured variation in gender among individuals within populations of ragweed (Ambrosia artemisiifolia) in an abandoned old-field and in the greenhouse. There was great variability in sex expression, from all-female to approximately 78% male. Plants differed significantly in gender in different locations within the field. Plants in an area abandoned from agriculture one year previously were more male than plants in a nearby area abandoned four years previously. In the greenhouse, soil moisture treatments and levels of attack by spittlebugs (Philaenus spumarius) did not affect gender. Height was positively correlated with relative maleness in both populations. Plants with greater shoot weight were relatively more male in the greenhouse, but not in the field. The gender variation we observed either has a genetic basis or is controlled by environmental variables other than those we investigated.     

Differences in Fecal Androgen Patterns of Breeding and Nonbreeding Kori Bustards (Ardeotis kori)

To better understand breeding conditions to promote reproduction in captive kori bustards, fundamental endocrine studies measuring fecal androgen metabolites in male and female kori bustards were conducted. Feces collected weekly from males and females were analyzed for testosterone using enzyme‐linked immunoassay. Results from adult males (n = 5), adult females (n = 10), immature males (n = 10), and immature females (n = 10) revealed seasonally elevated testosterone concentrations in fertile, but not nonfertile adult males and females (P > 0.05). Adult females that were not maintained in a breeding group, or that did not produce eggs, did not demonstrate increases in testosterone compared to egg laying counterparts. In males, but not females, seasonal testosterone increases were accompanied by weight gain. Peaks in male fecal androgen metabolites ranged from 10‐ to 22‐fold higher than nonbreeding season (181.5 ± 19.1 vs. 17.0 ± 0.94 ng/g; P < 0.05). Mean breeding season values for adult males were 83.6 ± 6.1 ng/g vs. nonbreeding season values of 12.3 ± 0.73 ng/g (P < 0.05). In females, average breeding season testosterone concentrations were approximately 4‐fold higher than nonbreeding season (55.9 ± 6.0 vs. 14.5 ± 1.8 ng/g), with peaks 10‐ to 30‐fold higher. Results show that noninvasive fecal androgen metabolite analysis can provide a means of predicting fertility potential of male and female kori bustards and might be utilized to assess effects of modifying captive environments to promote reproduction in this species. Zoo Biol. 32:54‐62, 2013. © 2012 Wiley Periodicals, Inc.     

Plant genotype and induced responses affect resistance to herbivores on evening primrose (Oenothera biennis)

Abstract. 1. Although both genotype and induced responses affect a plant's resistance to herbivores, little is known about their relative and interactive effects. This study examined how plant genotype of a native plant (Oenothera biennis) and induced plant responses to herbivory affect resistance to, and interactions among, several herbivores. 2. In a field experiment, genetic and environmental variation among habitats led to variation in the amount of early season damage and plant quality. The pattern of variation in early season infestation by spittlebugs (Philaenus spumarius, a piercing–sucking herbivore) negatively correlated with oviposition preference by a later feeding specialist weevil (Tyloderma foveolatum, a leaf‐chewer). 3. To determine if plant genotype and induced responses to herbivory might be responsible for these field patterns, we performed no‐choice and choice bioassays using four genotypes of O. biennis that varied in resistance. Plants were induced by either spittlebugs or weevils and assays measured the responses of the same specialist weevil as well as a generalist caterpillar (Spodoptera exigua). 4. Resistance to adult weevils was largely unaffected by plant genotype, while they experienced induced resistance following damage by conspecific weevils in no‐choice assays. Caterpillars were more strongly affected by plant genotype than induced responses in both no‐choice and choice assays, but they also fed less and experienced higher mortality on plants previously damaged by weevils. In contrast to the pattern suggested by the field experiment, spittlebugs did consistently induce resistance against either weevils or caterpillars in the bioassay experiment. 5. These results support recent findings that show herbivore species can compete via induced plant responses. Additionally, a quantitative review of the literature demonstrates that plant genotype tends to be more important than interspecific competition among herbivores (plant‐mediated or otherwise) in affecting herbivore preference and performance.     

The effect of resistant solanaceous plants on the sex ratio of Heterodera rostochiensis and the use of the sex ratio to assess the frequency and genetic constitution of pathotypes

The sex ratios of two Heterodera rostochiensis populations were compared on fifteen solanaceous plants. Larvae did not mature in the roots of Atropa belladona, Datura stramonium or Nicotiana tabacum but adults were produced on the other plants, on some only males and on others, males and females. With two other populations, but not with a third, the percentage of larvae that became female on a potato hybrid with a dominant gene for resistance derived from Solanum tuberosum ssp. andigena increased during successive generations on the hybrid in accordance with the hypothesis that such larvae are double recessives (aa). Fewer larvae became adult in plants with two dominant genes for resistance than in plants with one, and such plants probably exerted selection of both males and females. A population from St Brelades (Jersey) produced fewer females and more males on the susceptible variety Arran Banner than did either the Thorney (Cambs.) or Aberystwyth populations. A much greater percentage of larvae from the St Brelades population became female on an ex andigena hybrid derived from CPC 1690 than on one derived from CPC 1685.     

Geographic variation in the colour/ pattern polymorphism of British Philaenus spumarius (L) (Homoptera: Aphrophoridae) populations *

Geographic variation in the frequency of colour/pattern morphs of Philaenus spumarius is described at 48 localities in England and Wales. The sites have been divided into four broad environmental categories, two urban and two rural. There is a distinction between the urban and rural groupings in the relative proportions of the colour morphs; in urban localities there is a higher combined frequency of the eight ‘melanic’ morphs. Differences also exist between locality groupings, and between the sexes, in the relative frequencies of the eight morphs within the ‘melanic’ phenotypic category. Morphs limited to the female sex in some other parts of the species range occur among males in British populations. It is suggested that the different dominance hierarchy between the sexes which exists in Fennoscandian populations may not be characteristic of British populations.     

Agonistic behavior correlated with hydrocarbon phenotypes in dampwood termites,Zootermopsis (Isoptera: Termopsidae)

The dampwood termite genus ZootermopsisEmerson contains three recognized species with four distinct and consistent hydrocarbon phenotypes. Agonistic behaviors among nonreproductive insects from colonies of the same and different hydrocarbon phenotypes were observed in the laboratory. Various combinations of soldier versus nymphs, pseudergate versus pseudergate, and soldier versus soldier encounters were used in experimental trials. Soldiers or pseudergates seldom attack individuals of the same hydrocarbon phenotype. Z. angusticollis(Hagen) (phenotype II) is typically aggressive toward phenotype III of Z. nevadensis(Hagen) but not always aggressive against phenotype I of Z. nevadensis.The variation in response is dependent on which castes are placed in the bioassay arena: soldier versus soldier bouts result in consistent aggression, while pseudergate versus pseudergate or soldier versus nymphs contacts do not. Both pseudergates and soldiers of Z. laticeps(Banks) (phenotype IV) respond agonistically toward the other three phenotypes: Z. angusticollis (II) and Z. nevadensis(I and III). Although hydrocarbon phenotypes I and III, both Z. nevadensis,are morphologically indistinguishable, agonistic behavioral responses between phenotype I and phenotype III are not equivalent to I versus I or III versus III behavioral responses. The I versus III engagements, regardless of the castes involved, display a greater proportion of avoidance and aggressive responses than I or III intraphenotype encounters. We interpret the lack of avoidance or aggressive behavior within each of the two phenotypes of Z. nevadensisand the significant avoidance and aggressive behavior between phenotypes as definite evidence of discrimination between disparate hydrocarbon phenotypes. These agonistic bioassays along with data on distinct hydrocarbon patterns and geographic distributions serve as the basis for creating two subspecies of Z. nevadensis: Z. n. nevadensis(Hagen) and Z. n. nuttingiHaverty and Thorne, ssp. nov.     

SEXUAL TETRAMORPHISM IN THYMELAEA HIRSUTA (THYMELAEACEAE): EVIDENCE OF THE PATHWAY FROM HETERODICHOGAMY TO DIOECY AT THE INFRASPECIFIC LEVEL

After more than 5 years of study on natural populations of Thymelaea hirsuta (L.) Endl. (Thymelaeaceae), four distinct sexual phenotypes are shown to occur: protogynous, protandrous, subandroecious (male), and subgynoecious (female) individuals. The four populations studied differ significantly in the relative abundance of sexual types. Plant size and sexual phenotype are independent in three populations. Each sexual form produces viable seeds. Growth and morphogenesis of the aerial vegetative and sexual parts are described and found not to be related to sexual phenotype. The system, called sexual tetramorphism, combines characteristics of subdioecy (subandroecious and subgynoecious individuals) and heterodichogamy (protogynous and protandrous individuals). High fruit production in females and protandrous plants vs. low fruit production in males and protogynous plants provides evidence of a strong tendency toward functional dimorphism. The probable specialization of protogynous types toward the male function and the obvious specialization of protandrous types toward the female function suggest an evolution from heterodichogamy to dioecy, sexual tetramorphism being the intermediate stage. Tetramorphism in T. hirsuta provides further support for the hypothesis of an evolutionary pathway from heterodichogamy to dioecy.     

Industrial melanism in the spittlebug Philaenus spumarius (L) (Homoptera: Aphrophoridae)*

The visible polymorphism of the spittlebug, Philaenus spumarius, has been investigated in the vicinity of a smokeless fuel factory in the Cynon Valley of south Wales. The factory is a significant source of local particulate air pollution. A striking relationship exists between the combined frequencies of the eight dark (melanic) morphs and proximity to the factory. Maximum melanic phenotype frequencies of over 95“, occur in both sexes immediately adjacent to it and decline to levels normal for south Wales 1.5–6 km away, depending on direction. This relationship is largely confined to sites within the valley; samples from adjacent localities outside it have melanic frequencies within normal limits for the south Wales area. Maximum melanic phenotype frequencies in the Cynon Valley are far higher than any known from elsewhere in the species range in Europe, Asia and North America. No consistent difference is apparent in total melanic frequency between males and females at any of the sites in this study. However, marked differences exist between the sexes in the relative contributions of the eight melanic phenotypes to the overall association with the factory. For females the industrial melanism is entirely attributable to the group flanicollis + gibbus + leucocephalus (mainly leucocephalus) whereas in males both this group and the group quadnmaculatus + albomaculatus + leucopthalmus contribute to the relationship. It is suggested that this relationship is due to the selective effects of the local air pollution from the factory. The exact nature of the selection involved is as yet uncertain; it would appear to be strong since the local adaptation involved has developed in a maximum of 40 generations since the factory was opened. Finally, comparison is made with two other insects, a ladybird and a moth, in which high frequencies of melanic forms are also associated with this pollution source.