Wingless Morph Research Articles

WING POLYMORPHISM OF RHIPIDOTHRIPS BRUNNEUS WILLIAMS, 1913 (THYSANOPTERA: AEOLOTHRIPIDAE

One of the small genera of the family Aeolothripidae (Thysanoptera: Terebrantia) Rhipidothrips Uzel includes 6 species. Up to now, 4 species of this genus have been recorded in Iran. The wingless morph of R. brunneus Williams is described for the first time based on specimens collected from pasture plants in Javanrud County, Kermanshah Province (western Iran). A key to Iranian species of Rhipidothrips is also provided.

Cell death and wing reduction during the metamorphosis of sex-specific flightless morphs in winter geometrid moths.

Winter geometrid moths show striking sexual dimorphism by having female-specific flightless morphs. The evolutionary grades of wing reduction in winter geometrid moths vary and range from having short wings, vestigial wings, to being wingless. Although the ontogenetic processes underlying the development of the wingless or short-wing morphs in Lepidoptera has been well studied, the mechanisms underlying the development of vestigial wing morphs in winter geometrid moths during metamorphosis are poorly understood. In the winter geometrid moth Sebastosema bubonaria Warren, 1896, the males have functional wings, but the females have vestigial wings. Here, we studied the ontogenetic processes affecting wing reduction in the winter geometrid moth S. bubonaria using light microscopy and transmission electron microscopy, and compared the ontogenetic process of wing reduction in this species with that in another species of the wingless-female winter moth that we investigated previously. Our results showed that, in the vestigial-wing morphs, the loss of pupal wing epithelium was terminated in the middle of the wing degeneration process, whereas in the wingless morph, the pupal wing epithelium disappeared almost completely and the final appearance of the wings differed slightly among flightless morphs. We propose that the extent and location of cell death in the pupal wing play an important role in the various patterns of reduced wings that are observed in flightless moths.

Salivary carbonic anhydrase II in winged aphid morph facilitates plant infection by viruses

Aphids are the most common insect vector transmitting hundreds of plant viruses. Aphid wing dimorphism (winged vs. wingless) not only showcases the phenotypic plasticity but also impacts virus transmission; however, the superiority of winged aphids in virus transmission over the wingless morph is not well understood. Here, we show that plant viruses were efficiently transmitted and highly infectious when associated with the winged morph of Myzus persicae and that a salivary protein contributed to this difference. The carbonic anhydrase II (CA-II) gene was identified by RNA-seq of salivary glands to have higher expression in the winged morph. Aphids secreted CA-II into the apoplastic region of plant cells, leading to elevated accumulation of H+. Apoplastic acidification further increased the activities of polygalacturonases, the cell wall homogalacturonan (HG)-modifying enzymes, promoting degradation of demethylesterified HGs. In response to apoplastic acidification, plants accelerated vesicle trafficking to enhance pectin transport and strengthen the cell wall, which also facilitated virus translocation from the endomembrane system to the apoplast. Secretion of a higher quantity of salivary CA-II by winged aphids promoted intercellular vesicle transport in the plant. The higher vesicle trafficking induced by winged aphids enhanced dispersal of virus particles from infected cells to neighboring cells, thus resulting in higher virus infection in plants relative to the wingless morph. These findings imply that the difference in the expression of salivary CA-II between winged and wingless morphs is correlated with the vector role of aphids during the posttransmission infection process, which influences the outcome of plant endurance of virus infection.

Can Mixed Intercropping Protect Cereals from Aphid-Borne Viruses? An Experimental Approach.

Simple SummaryAphids are vectors of plant viruses and can cause important yield losses in numerous crops. Rhopalosiphum padi is the main vector of Barley yellow dwarf virus PAV (BYDV-PAV). The yellow dwarf disease (YDD), present in all cereal growing regions in the world, caused by BYDV-PAV, can induce up to 80% of yield losses in barley. In a context of global warming, reduction of pesticide uses, and development of organic farming, new control methods for YDD incidence are needed. In this study, the association of a winter barley with clover was tested for its impact(s) on the R. padi/BYDV-PAV pathosystem. The effect of clover was different according to the morph of the aphid introduced in a multi-plant (i.e., arena) experimental design in laboratory. A spatial effect on aphid distribution (observed for alate founder morph) and a reduction of the size of aphid populations (observed for wingless founder morph) are described. However, the presence of clover did not modify, under our experimental conditions, the efficiency of BYDV-PAV infections and within arena spread of YDD. Thus, clover used by cereal growers for other services (weeds reduction, nitrogen supply, and soil cover), would participate to lower the risks associated to the presence of BYDV vectors through a bottom-up regulation of aphid populations within the field. Barley/clover intercropping should be considered as a promising method for a future management strategy against vectors of yellow dwarf disease, especially in an insecticide-free agriculture.Intercropping, i.e., association of two or more species, is promising to reduce insect populations in fields. The cereal aphid Rhopalosiphum padi, a vector of the Barley yellow dwarf virus PAV (BYDV-PAV), represents a major threat for cereal grain production. In this study, we tested the potential of winter barley intercropped with clover to reduce the size of R. padi populations and to lower the BYDV-PAV incidence in fields. We used arenas (i.e., sets of 36 barley plants) intercropped with or without clover plants (at different sown densities). In each arena, a single viruliferous founder, R. padi, (with an alate or a wingless morph) was deposited to introduce aphids and viruses in the experiment. Thirteen days later, the number of aphids in the arena, the percentage of plants hosting aphids and the infection rates were monitored. Data produced through this experimental design showed that clover alters the distribution of the aphid progeny (lower aphid spread) produced by an alate founder morph. Moreover, clover reduces the size of aphid populations produced by a wingless founder morph. However, despite the effects of clover on biological parameters of R. padi, the presence of clover in barley arena did not modify BYDV infections, suggesting complex mechanisms between partners of the BYDV pathosystem for plant-to-plant virus spread.

No Evidence of Bacterial Symbionts Influencing Host Specificity in Aphis gossypii Glover (Hemiptera: Aphididae).

Simple SummaryThe cotton-melon aphid, Aphis gossypii Glover, is a polyphagous insect pest with many host-specialized biotypes. The mechanism of host specificity remains unknown in this aphid. In this study, we investigated whether bacterial symbionts control the host specificity of A. gossypii, as reported in some aphids. The two typical host-specialized biotypes used in this study produced significantly fewer nymphs on non-native hosts than on a native host, indicating a high host specificity of the two biotypes. We found that the winged morph of both biotypes had a significantly lower host specificity than its corresponding wingless morph. Bacterial analysis indicated that the composition of the bacterial symbionts was not different between the two biotypes, but within each biotype, the Buchnera abundance in the winged morph was only about 10% of that in the wingless morph. We suspected that a low Buchnera abundance was associated with a low host specificity. We compared the reproduction of A. gossypii with different Buchnera abundances, and did not find that a low Buchnera abundance resulted in a low host specificity. We then concluded that the host specificity of A. gossypii is not controlled by specific bacterial symbionts or Buchnera abundance.The cotton-melon aphid, Aphis gossypii Glover, is a polyphagous insect pest with many host-specialized biotypes, such as the Cucurbitaceae- and Malvaceae-specialized (CU and MA) biotypes. Bacterial symbionts were reported to determine the host range in some aphids. Whether this is the case in A. gossypii remains unknown. Here, we tested the host specificity of the CU and MA biotypes, compared the host specificity between the wingless and winged morph within the same biotype, and analyzed the composition of the bacterial symbionts. The reproduction of the CU and MA biotypes reduced by 66.67% and 82.79%, respectively, on non-native hosts, compared with on native hosts. The composition of bacterial symbionts was not significantly different between the CU and MA biotypes, with a Buchnera abundance >95% in both biotypes. Meanwhile, the winged morph produced significantly more nymphs than the wingless morph on non-native hosts, and the Buchnera abundance in the winged morph was only about 10% of that in the wingless morph. There seemed to be a relationship between the Buchnera abundance and host specificity. We regulated the Buchnera abundance by temperature and antibiotics, but did not find that a low Buchnera abundance resulted in the high reproduction on non-native hosts. We conclude that the host specificity of A. gossypii is not controlled by specific bacterial symbionts or by Buchnera abundance.

Comparative transcriptional analysis provides insights of possible molecular mechanisms of wing polyphenism induced by postnatal crowding in Aphis gossypii

BackgroundAphis gossypii is a worldwide sap-sucking pest with a variety of hosts and a vector of more than 50 plant viruses. The strategy of wing polyphenism, mostly resulting from population density increasing, contributes to the evolutionary success of this pest. However, the related molecular basis remains unclear. Here, we identified the effects of postnatal crowding on wing morph determination in cotton aphid, and examined the transcriptomic differences between wingless and wing morphs.ResultsEffect of postnatal crowding on wing determination in A. gossypii was evaluated firstly. Under the density of 5 nymphs·cm− 2, no wing aphids appeared. Proportion of wing morphs rised with the increase of density in a certain extent, and peaked to 56.1% at the density of 20 nymphs·cm− 2, and reduced afterwards. Then, transcriptomes of wingless and wing morphs were assembled and annotated separately to identify potentially exclusively or differentially expressed transcripts between these two morphs, in which 3 126 and 3 392 unigenes annotated in Nr (Non-redundant protein sequence) database were found in wingless or wing morphs exclusively. Moreover, 3 187 up- and 1 880 down-regulated genes were identified in wing versus wingless aphid. Pathways analysis suggested the involvement of differentially expressed genes in multiple cellular signaling pathways involved in wing morphs determination, including lipid catabolic and metabolism, insulin, ecdysone and juvenile hormone biosynthesis. The expression levels of related genes were validated by the reverse transcription quantitative real time polymerase chain reaction (RT-qPCR) soon afterwards.ConclusionsThe present study identified the effects of postnatal crowding on wing morphs induction and demonstrated that the critical population density for wing morphs formation in A. gossypii was 20 nymphs·cm− 2. Comparative transcriptome analysis provides transcripts potentially expressed exclusively in wingless or wing morph, respectively. Differentially expressed genes between wingless and wing morphs were identified and several signaling pathways potentially involved in cotton aphid wing differentiation were obtained.

Morph-specific differences in life history traits between the winged and wingless morphs of the aphid, <i>Sitobion avenae</i> (Fabricius) (Hemiptera: Aphididae)

Life history traits were evaluated in the wing polyphenic aphid, Sitobion avenae (Fabricius), by rearing the winged and wingless morphs under the laboratory conditions. Winged morph with large thoraces exhibited a significantly greater morphological investment in flight apparatus than wingless morph with small thoraces. Compared to the winged morph, the wingless morph produced significantlymore nymphs and exhibited significantly faster nymph development rates. In addition, the age at which reproduction first occurred for the winged morph was significantly delayed, and higher mortality was recorded.The results suggest that the fitness differences associated with wingsmay be related to nymph development, adult fecundity, and mortality. Based on these results, the trends and exceptions of life history traits for the wing polyphenic insects are discussed.

Achene dimorphism and protracted release: a trait syndrome allowing continuous reshaping of the seed-dispersal kernel in the Mediterranean species Pallenis spinosa

ABSTRACTBackground: Pallenis spinosa (Asteraceae) produces both winged and wingless achenes. Both achene morphs are non-dormant and show a similar embryo size, rendering dispersal ability as their only apparent functional difference.Aims: We studied morph-specific release and spatial dispersal patterns to ascertain whether the common view of seed dimorphism as a mixed strategy, that is functionally fully differentiated morphs, is appropriate for this system.Methods: For three years, at the onset of achene release, in early autumn, we placed achene traps at different distances from source plants, censusing achene arrival at 3–4 day intervals. We constructed morph-specific dispersal kernels and related release intensity to prevailing meteorological conditions in census intervals. Selected kernel models were used to describe dispersal effects of observed changes in the proportion of winged achenes (pw) in successive released fractions.Results: Achene release extended up to early-mid winter, peaking in rainy, windy intervals. Throughout the season, pw decreased progressively. Unexpectedly, the wingless morph produced the longest dispersal tails and it only showed ability for fat-tailed dispersal. Consequently, maximum dispersal distances steadily increased throughout the season.Conclusions: Achene dimorphism in P. spinosa appears to allow a within-season continuous reshaping of the seed-dispersal kernel instead of representing a mixed strategy.

Sensilla Morphology and Complex Expression Pattern of Odorant Binding Proteins in the Vetch Aphid Megoura viciae (Hemiptera: Aphididae).

Chemoreception in insects is mediated by several components interacting at different levels and including odorant-binding proteins (OBPs). Although recent studies demonstrate that the function of OBPs cannot be restricted to an exclusively olfactory role, and that OBPs have been found also in organs generally not related to chemoreception, their feature of binding molecules remains undisputed. Studying the vetch aphid Megoura viciae (Buckton), we used a transcriptomic approach to identify ten OBPs in the antennae and we examined the ultrastructural morphology of sensilla and their distribution on the antennae, legs, mouthparts and cauda of wingless and winged adults by scanning electron microscopy (SEM). Three types of sensilla, trichoid, coeloconic and placoid, differently localized and distributed on antennae, mouthparts, legs and cauda, were described. The expression analysis of the ten OBPs was performed by RT-qPCR in the antennae and other body parts of the wingless adults and at different developmental stages and morphs. Five of the ten OBPs (MvicOBP1, MvicOBP3, MvicOBP6, MvicOBP7, and MvicOBP8), whose antibodies were already available, were selected for experiments of whole-mount immunolocalization on antennae, mouthparts, cornicles and cauda of adult aphids. Most of the ten OBPs were more expressed in antennae than in other body parts; MvicOBP1, MvicOBP3, MvicOBP6, MvicOBP7 were also immunolocalized in the sensilla on the antennae, suggesting a possible involvement of these proteins in chemoreception. MvicOBP6, MvicOBP7, MvicOBP8, MvicOBP9 were highly expressed in the heads and three of them (MvicOBP6, MvicOBP7, MvicOBP8) were immunolocalized in the sensilla on the mouthparts, supporting the hypothesis that also mouthparts may be involved in chemoreception. MvicOBP2, MvicOBP3, MvicOBP5, MvicOBP8 were highly expressed in the cornicles-cauda and two of them (MvicOBP3, MvicOBP8) were immunolocalized in cornicles and in cauda, suggesting a possible new function not related to chemoreception. Moreover, the response of M. viciae to different components of the alarm pheromone was assessed by behavioral assays on wingless adult morph; (-)-α-pinene and (+)-limonene were found to be the components mainly eliciting an alarm response. Taken together, our results represent a road map for subsequent in-depth analyses of the OBPs involved in several physiological functions in M. viciae, including chemoreception.

Morph-specific differences in biochemical composition related to flight capability in the wing-polyphenic Sitobion avenae

Flight performance at various times after emergence in the alate morph and age-dependent changes in biochemical composition of winged and wingless morphs were evaluated in the wing-polyphenic aphid Sitobion avenae (Fabricius) (Hemiptera: Aphididae). Alates exhibited the highest flight activity at 18–36 h after adult emergence. Throughout the nymphal and adult development, the whole-body content of total lipid was significantly higher in the winged vs. wingless morph, whereas the content of water, soluble sugar, glycogen, phospholipid, and soluble protein showed significantly higher levels in the wingless vs. winged morph. There were no significant differences in the content of triglyceride and free fatty acid during nymphal and adult stages in both morphs. However, triglyceride content was significantly higher in the winged vs. wingless morph during adulthood. Differences in biochemical composition between morphs indicate that there is an energetic cost of flight capability. Our results from S. avenae adults showed that total lipid and triglyceride for the winged morph accumulated significantly to a maximum, and water content decreased significantly to a minimum, on days 1 and 2 after the final molt, exactly when the highest flight activity was reached. This study suggests that flight activity is associated with triglyceride and water content.

Morph‐specific differences in metabolism related to flight in the wing‐dimorphicAphis gossypii

Abstract The cotton aphid,Aphis gossypiiGlover, is a wing‐dimorphic species, which causes globally important agricultural losses. In this present study, we compared the biochemical basis of wing polymorphism inA. gossypiiwith respect to trade‐off of energy resources, including glycogen, trehalose, lipids (total lipid, triglyceride and phospholipid), free fatty acids, and soluble protein between dispersal and reproduction morphs during the wing‐bud nymph and adulthood. Total lipid, triglyceride and free fatty acids were significantly higher in winged versus wingless morphs at 12 h of adulthood, the period during which alates are able to fly. By contrast, the wingless morph contained more glycogen than the winged morph from the 4th nymphal stage to adulthood. Trehalose content in the wingless morph was also higher than that in the winged morph during the 3rd and 4th nymphal stages, butvice versaat 12 h of adulthood. Finally, soluble protein content increased from nymphs to adults and was higher during adulthood in aptera versus alate. Whole‐body water content in 12‐h adults was significantly higher in apterae than that in alatae. These results indicate significant physiological differences between morphs related to specialization for flight.

Morph-Dependent Expression and Subcellular Localization of Host Serine Carboxypeptidase in Bacteriocytes of the Pea Aphid Associated with Degradation of the Endosymbiotic Bacterium Buchnera

Pea aphids form a mutualistic association with the endosymbiotic bacterium Buchnera, which is harbored in specialized host cells called bacteriocytes. The adult aphids display dimorphism In which there are winged and wingless morphs. We previously reported that the Buchnera density in bacteriocytes of the winged morph (alate) decreases around final ecdysis, whereas that in the wingless morph (aptera) does not decrease; the decrease in density in alatae is accompanied by activation of the host lysosomal system and by Buchnera degradation. In the present study, we performed a proteomic analysis to clarify the molecular mechanisms underlying the decrease in Buchnera density. By comparing the protein expression profiles of bacteriocytes in alatae and apterae Just after final ecdysis, we identified three and one protein spots that were preferentially expressed in alatae and apterae, respectively. Among the three alate-preferential spots, two were an identical aphid protein, carboxypeptidase vitellogenic-like (CPVL), whereas the other was a mixture of four proteins: gamma-glutamyl hydrolase, acyl-CoA dehydrogenase, aphid short chain acyl-CoA dehydrogenase, and Buchnera S-adenosylmethionine synthetase. The aptera-preferential spot was Buchnera outer membrane protein A. Immunoblot and immunohistochemical analyses using aphid bacteriocytes Just after final ecdysis revealed that expression of aphid CPVL was preferentially upregulated in alatae and was localized around Buchnera cells in the bacterlocytes, suggesting the involvement of CPVL in Buchnera degradation in alatae.

Development of a wingless morph in the ladybird beetle, Adalia bipunctata

Many taxa of winged insects have independently lost the ability to fly and often possess reduced wings. Species exhibiting natural variation in wing morphology provide opportunities to investigate the genetics and developmental processes underlying the evolution of alternative wing morphs. Although many wing dimorphic species of beetles are known, the underlying mechanisms of variation are not well understood in this insect order. Here, we examine wing development of wild type and natural wingless morphs of the two-spot ladybird beetle, Adalia bipunctata. We show that both pairs of wings are distally truncated in the wingless adults. A laboratory population of the wingless morph displays heritable variation in the degree of wing truncation, reflecting reduced growth of the larval wing discs. The coexistence of variable wingless morphs supports the idea that typical monomorphic wingless insects may be the result of a gradual evolution of wing loss. Gene expression patterns in wing discs suggest that the conserved gene network controlling wing development in wild-type Adalia is disrupted in the dorsoventral patterning pathway in the wingless morphs. Previous research on several species of ant has revealed that the anteroposterior wing patterning pathway is disrupted in wingless workers. Future investigations should confirm whether interruptions in both taxa are limited to the patterning pathways found thus far, or whether there are also shared interruption points. Nevertheless, our results highlight that diverse mechanisms of development are likely to underlie the evolution of wingless insects.

Age- and morph-dependent activation of the lysosomal system and Buchnera degradation in aphid endosymbiosis

Endosymbiosis in aphids is maintained through a mutualistic association between the host and a symbiotic bacterium, Buchnera, which is harbored in specialized host cells called bacteriocytes. Here, we examined the changes in the Buchnera density in bacteriocytes in relation to the development and polyphenism of the host aphid. Buchnera density in the winged morph aphids, alatae, decreased drastically around the final ecdysis, whereas in the wingless morph aphids, apterae, Buchnera density decreased after the final ecdysis. Thereafter, in both apterae and alatae, Buchnera density was maintained at a constant level until 10 days and then again decreased gradually until 18 days after the final ecdysis. Cytochemical analysis with LysoTracker reagent and quantitative RT-PCR analysis revealed that the number of lysosome-like acidic organelles and the amount of lysosome-related gene ( lysozyme and cathepsin L) transcripts increased drastically in the bacteriocytes of alatae around the final ecdysis. Electron microscopy of alatae bacteriocytes around the final ecdysis revealed many Buchnera with irregular electron-dense areas in their cytoplasm that were enclosed by a distended symbiosome membrane. These findings indicated that age- and morph-dependent decreases in Buchnera density coincided with activation of the host lysosomal system and the increased degradation of Buchnera.

The cabbage aphid: a walking mustard oil bomb

The cabbage aphid, Brevicoryne brassicae, has developed a chemical defence system that exploits and mimics that of its host plants, involving sequestration of the major plant secondary metabolites (glucosinolates). Like its host plants, the aphid produces a myrosinase (beta-thioglucoside glucohydrolase) to catalyse the hydrolysis of glucosinolates, yielding biologically active products. Here, we demonstrate that aphid myrosinase expression in head/thoracic muscle starts during embryonic development and protein levels continue to accumulate after the nymphs are born. However, aphids are entirely dependent on the host plant for the glucosinolate substrate, which they store in the haemolymph. Uptake of a glucosinolate (sinigrin) was investigated when aphids fed on plants or an in vitro system and followed a different developmental pattern in winged and wingless aphid morphs. In nymphs of the wingless aphid morph, glucosinolate level continued to increase throughout the development to the adult stage, but the quantity in nymphs of the winged form peaked before eclosion (at day 7) and subsequently declined. Winged aphids excreted significantly higher amounts of glucosinolate in the honeydew when compared with wingless aphids, suggesting regulated transport across the gut. The higher level of sinigrin in wingless aphids had a significant negative impact on survival of a ladybird predator. Larvae of Adalia bipunctata were unable to survive when fed adult wingless aphids from a 1% sinigrin diet, but survived successfully when fed aphids from a glucosinolate-free diet (wingless or winged), or winged aphids from 1% sinigrin. The apparent lack of an effective chemical defence system in adult winged aphids possibly reflects their energetic investment in flight as an alternative predator avoidance mechanism.

The extraordinary mating system of Zeus bugs (Heteroptera:Veliidae:Phoreticovelia sp.)

Wingless female Zeus bugs (genus: Phoreticovelia) produce a secretion from dorsal glands that males feed upon when riding on females. This unique form of sex-role-reversed nuptial feeding may have set the stage for an unusual mating system. Here, we provide natural history details of the mating behaviour for two Zeus bug species. While these species have many mating behaviours in common, the wing morphs within species exhibit entirely different mating strategies. Adult wingless females are ridden permanently by adult wingless males. In the wild, adult sex-ratios among the wingless morph are male-biased; few unmounted adult females exist and many males instead ride immature females who also produce glandular secretions. In contrast, sex-ratios among the winged morph is not male-biased, sexual size dimorphism is less pronounced, females have no dorsal glands and are, consequently, not ridden by males. Field and laboratory observations show that mating is strongly assortative by wing morph. This assortment may allow evolutionary divergence between the two morphs. We discuss the implications of this mating system and suggest that it adds to those studies showing that sexually antagonistic coevolution can be a driver of mating system evolution.

A transitional stage between the ergatoid and winged male morph in the ant Cardiocondyla obscurior

Males of the ant Cardiocondyla show a dispersal dimorphism of a winged and wingless morph. The loss of flight has lead to morphological reductions in the wingless (ergatoid) males and also affected body size, eye size and pigmentation. As ergatoid males mate exclusively inside the maternal nest, they underlie increased male-male competition and therefore have also evolved additional changes in behaviour and physiology: in contrast to winged males, ergatoid males are highly aggressive towards each other and their spermatogenesis is prolonged compared to all other hymenopteran males. In addition to these two male morphs, we found males with an intermediate appearance. These "intermorphic" males provide a transitional stage between normal males in most investigated morphological and physiological parameters. As they are produced extremely rarely and only in colonies that switch between pure ergatoid to mixed male production, we argue that they likely represent a developmental mistake. Parallels between the determination of male morphs and female castes (queen-worker dimorphism and worker polymorphism) might help to understand how the large potential of phenotypic plasticity in both sexes of social insects is realised during development.

Life history, ecology and communal gall occupation in the manzanita leaf-gall aphid, Tamalia coweni (cockerell) (Homoptera: Aphididae)

I describe life history details for a California population of Tamalia coweni on the host plants Arctostaphylos patula and Arctostaphylos viscida. In particular, studies on the sequence of gall initiation and the identity of gall-making morphs reveal that later generations of wingless females, as well as the stem mother, are capable of inducing galls. Winged females disperse to other sites suitable for gall induction and deposit wingless gall-forming females. The life cycles of T. coweni on the two species of Arctostaphylos studied differ considerably and affect the ecology and population biology of the aphid. I discuss the possible role of a newly observed wingless morph in the life history of T. coweni. I document communal gall occupation by multiple aphid foundresses and place it into a sociobiological context.

Photoperiodic Induction of Wing Morphs in the Waterstrider Limnoporus canaliculatus (Gerridae: Hemiptera)

The influence of photoperiod on wing morph induction was studied in the wing-dimorphic waterstrider Limnoporus canaliculatus Say. The winged morph was strongly induced under a short 12:12 (L:D) stationary photoperiod and to a greater degree by a single transfer from a long (16:8) to a short, stationary photoperiod (12:12) at hatch. Morph frequencies obtained in the transfer from a long to a short photoregime closely approximated those in a natural population during the fall, when the winged morph is almost exclusively produced. Long, stationary daylength or a single change from short to long daylength resulted in an equivalent, maximal production of the wingless morph in the laboratory. These results are consistent with the maximal production of the wingless morph during late June–July in the field when daylength is longest. Continuous change in daylength (incremental or decremental) elicited no important effect on morph induction beyond that elicited by a single change in daylength. Parental chilling, field generation of the parent (summer versus fall), or environmentally-induced wing phenotype of the parent did not affect morph induction in progeny. Developmental stages that were sensitive to photoperiod were exclusively or nearly exclusively restricted to the egg-second stadium. Several key aspects of photoperiodically mediated morph induction in L. canaliculatus are inconsistent with the general model of morph induction proposed for waterstriders.

DIFFERENCES IN SURVIVORSHIP, DEVELOPMENT RATE AND FERTILITY BETWEEN THE LONGWINGED AND WINGLESS MORPHS OF THE WATERSTRIDER, LIMNOPORUS CANALICULATUS.

Species of waterstriders (Gerridae: Hemiptera) exhibit a remarkable diversity in degree of winglessness (Vepsalainen, 1978; Calabrese, 1980; Zera, 1981). Some species consist exclusively of fully-winged individuals, while other species are composed almost exclusively of wingless individuals; many species exhibit the intermediate case of wing polymorphism and consist of various proportions of fully-winged, short-winged and/or wingless morphs. Wing-polymorphic species often exhibit dramatic spatial and/or temporal variation in morph ratios, both among populations of the same species and among species. Because of this diversity, waterstriders, especially wing polymorphic species, are ideal candidates for the study of the evolution of winglessness (or conversely, the evolution of dispersal). One of the major questions in the study of the evolutionary forces which influence wing polymorphism concerns fitness differences among the morphs. In many non-gerrids, dramatic differences between the fully-winged and the shortwinged or functionally equivalent (Anderson, 1973) wingless morph have been documented in a variety of fitness-associated traits (for reviews, see Harrison, 1980; Dingle, 1982). These include differences in such traits as survivorship under stress, duration of larval or nymphal development, age of first reproduction, and fecundity. Moreover, one of the most important results of these studies is the demonstration that differences in fitness traits are often consistently associated