Induction of winged sexuparae in root-inhabiting colonies of the aphid Pemphigus betae

Abstract. Growth chamber experiments were conducted to determine what environmental factors induce the development of sexuparae in root-feeding colonies of the aphid Pemphigus betae Doane (Aphididae: Pemphiginae). Experimental colonies were derived from clones that had been reproducing for at least 15 months under constant laboratory conditions. At constant 20–23^oC, sexuparae were almost never produced regardless of the level of crowding or photoperiod. In contrast, every culture of several clones produced sexuparae when subjected to temperatures alternating between 20 and 15^oC during the light and dark phases respectively. This fluctuating temperature regime resulted in sexupara production regardless of photoperiod. All cultures also produced some sexuparae when cultures were kept in constant darkness at temperatures that were either alternated between 20 and 15^oC or kept constant at 15^oC. Constant 17^oC resulted in sexupara production in most cultures. At 17^oC, experimental manipulation of densities within cultures showed that crowding promotes sexupara production. Thus, within the range of conditions occurring at the time of sexupara induction in the field, photoperiod has no effect on sexupara production, temperature below a threshold between 17 and 20–23^oC triggers sexupara production, and crowding promotes sexupara production but only if temperature is low enough.     

Seasonal shifts in host usage in Uroleucon gravicorne (Homoptera: Aphididae) and implications for the evolution of host alternation in aphids

ABSTRACT.

• 1 Uroleucon gravicome (Patch) feeds on annual Erigeron species and on perennial Solidago species.
• 2 To test whether Solidago and Erigeron are analogous to winter and summer hosts of typical host-alternating aphids, reproductive performance, host preferences, and host associations were measured for E.strigosus, S.juncea and S.nemoralis during spring, summer and autumn.
• 3 Caged individuals can reproduce on both genera throughout the season, though colonies are rare on Solidago during summer.
• 4 Developmental rate, adult weight, and fecundity decline between May and August on all hosts; however, the drop is least on E.strigosus, intermediate on S.nemoralis, and greatest on S.juncea.
• 5 Throughout the season, reproductive performance is at least as great on Erigeron as on Solidago, the difference being least in spring.
• 6 Soluble nitrogen content shows the same seasonal trends as reproductive performance, declining in all hosts, but declining least in E.strigosus.
• 7 Preference tests and field records show that U.gravicorne moves from Solidago to Erigeron when the first alatae mature in late spring, feeds on Erigeron during the summer, and, in late summer, returns to Solidago where overwintering eggs are laid.
• 8 Solidago is not nutritionally superior during spring or autumn; its use during those seasons may result from the need for good oviposition sites.

     

Phenotypic plasticity in host choice behavior in black bean aphid, Aphis fabae (Homoptera: Aphididae)

The study of phenotypic plasticity in host choice behavior is crucial to predict evolutionary patterns of insect–plant interactions. The presence of sufficient variation in plasticity may facilitate host race formation and sympatric speciation. In this study, 13 Aphis fabae Scopoli genotypes reared both on broad bean and nasturtium exhibited statistically significant genotypic variability in host selection behavior. Some genotypes displayed increase in preference and acceptance in a novel host plant through generations. There are also strong conditioning effects of nasturtium as nasturtium reared genotypes are more willing to choose nasturtium over broad bean while broad bean reared genotypes do not show differences in choosing between the two host plants. There are also positive relationships between fitness and host choice behavior particularly for nasturtium. Results of the study supported the hypothesis that phenotypic plasticity in host choice behavior may be one of the major determinants of the evolutionary trajectory of a parasitic species, such as aphids.     

PHENOTYPE FIXATION AND GENOTYPIC DIVERSITY IN THE COMPLEX LIFE CYCLE OF THE APHID PEMPHIGUS BETAE

The aphid Pemphigus betae typically shows a complex life cycle, with annual alternation between cottonwood trees, where it forms leaf galls, and herbaceous plants, where it lives on roots. Distinct phenotypes are associated with each phase. In a population in Utah, aphid clones vary in their tendencies to undergo the cottonwood phase of the life cycle, with certain clones rarely producing the winged migrants that initiate the cottonwood phase. To examine the extent of genotypic variability in life cycle traits and to determine the consequences of phase deletion for fitness in the remaining phase of the life cycle, I compared performances of sets of clones that differed in their tendencies to delete the cottonwood phase. Performances of 35 aphid clones were compared on root hosts in the laboratory, by quantifying size (a correlate of fecundity), developmental time, and mortality for clonal sublines. Clones that tend to delete the cottonwood phase and remain on roots developed faster, showed lower mortality, and were larger as compared to clones that tend to undergo an annual migration to cottonwood. These laboratory comparisons of individual clones agree with earlier findings from field experiments. Performance also varied significantly among clones within sets, with the set of clones not previously subjected to selection on root hosts showing higher variance than clones isolated following selection in the root-inhabiting phase. These data suggest that selection in the cottonwood phase opposes selection in the root phase. Thus, even in cases of polyphenism, in which divergent developmental pathways confer some independence of phenotypic expression in alternative phases, antagonistic pleiotropy limits adaptation within a phase. These findings on the correlates of intrapopulational variation in frequency of phase expression provide some of the best evidence for character release following reduction of a complex life cycle, and they support the hypothesis that the advantages of polyphenism may be limited by negative genetic correlations among fitness traits of alternative phenotypes.     

Molecular data support a rapid radiation of aphids in the Cretaceous and multiple origins of host alternation

Many aphids display a remarkably complex life cycle of host alternation, in which cyclical parthenogenesis is combined with the obligate use of two unrelated host plants. We used mitochondrial ribosomal DNA (partial 12S and 16S) sequences to reconstruct the phylogeny of aphids, to determine how many origins of host alternation and correlated major host-plant shifts have occurred. Our results agreed with previous morphological studies in that species clustered with good support at the level of tribes. There was little well-supported phylogenetic structure at levels deeper than tribes, however, except for the monophyly of two subfamilies, Aphidinae and Lachninae. We argue that aphids experienced a rapid radiation at the tribal level, after host shifting from gymnosperms to angiosperms. A rapid radiation is consistent with aphid fossils, which record the presence of few subfamilies in the late Cretaceous, but most extant tribes by the early Tertiary. Plant fossils also record host plants of aphid tribes diversifying during this time. A hypothesized mechanism by which host alternation has evolved (fundatrix specialization), coupled with the rapid radiation, implies that this life cycle may have originated as often as in the ancestor of each tribe that displays it. We also consider, however, an alternative hypothesis of fewer origins. The basal radiation of Aphididae was dated from molecular sequences to have occurred at approximately 80–150 Mya.     

The distribution of aphids in response to variation among individual host plants: Uroleucon rudbeckiae (Homoptera: Aphididae) andRudbeckia laciniata (Asteraceae)

Abstract. 1. An investigation was conducted to determine the effects of variation among individual plants of Rudbeckia laciniata L. on the distribution and abundance of the aphid Uroleucon rudbeckiae (Fitch).
2. Wild plants were examined repeatedly for up to 2¹/₂ years to obtain information about plant growth and flowering, and about aphid densities on individual plants.
3. Aphid densities on individual plants were positively associated with several measures of plant vigour, including height, growth since previous year, and occurrence of flowers.
4. The colonization of plants by alate aphids was more highly correlated with plant height than with plant exposure. Aphid densities on individual plants were more strongly associated with plant height than with number of colonists.
5. In all years there were many hosts which were never colonized by aphids. Some of these plants produced shoots but did not flower. Experimental infestation of these plants showed that they were unsuitable for aphid growth and reproduction. Some plants which did flower and which escaped aphid infestation were nevertheless suitable hosts. Previously unsuitable host individuals were able to support aphids when grown under glass-house conditions.     

Geographic variation in Pemphigus populicaulis (Insecta: Aphididae) in Eastern North America*,†

The geographic variation of 33 morphological characters of the gall-forming aphid Pemphigus populicaulis is studied for 118 localities east of 100^oW longitude. Variation can be partitioned into within-gall, among-gall and among-locality components. Among localities variation ranges from 26 to 54%, being significant for all characters. Variation among galls within localities ranges from 24 to. 56%, that within-galls from 8 to 4796. The design of the study permits computation of character correlation matrices within and among localities. Gall size is correlated with tnorphometric characters only on an interlocality but not on an intralocality basis. Interlocalily correlations are a function of intralocality correlations, confirming earlier predictions. There is little correlation between characters of stem mother and alate morphotypes within localities, whereas among localities such correlation is appreciable. This phenomenon may be caused by aspects ol the environment that vary among localities but remain reasonably constant through the earlv life cycle of the aphid. When subjected to factor analysis both correlation matrices yield four factors. Multiple discriminant analysis of the data set results in five interpretable significant axes. Maps are furnished for characters representing the independent dimensions of variation and for discriminant function scores. The patterns of variation can be shown to be significantly nonrandom by Mantel's test and by spatial autocorrelation analysis. All variables are significantly positively autocorrelated at 200 km, many at 400 km and a few at 600 km; few general statements can be made about significant autocorrelations at higher distances. The positive autocorrelation at relatively short distances may be related to the pool of clones from which the genotypes of any one locality sample are taken. There are three correlogram patterns that can be associated with four clusters of variation patterns of characters. The separate patterns presumably cannot be explained by a single microevolutionary process.     

Defensive Behavior in Primary- and Secondary-Host Generations of the Soldier-Producing Aphid, Pemphigus bursarius (Hemiptera: Aphididae)

The genus Pemphigus comprises several species that produce soldiers (defensive morphs) in galls on the primary host. At the moment, it is unclear if host-alternating species also produce defenders on their secondary host. We therefore examined how P. bursarius morphs of the secondary host generations respond to predators to test whether they show defensive behavior. We further examined how this response compares with the antipredator behavior of soldiers in P. bursarius and P. spyrothecae occurring on the primary host. We performed two manipulative experiments using two predatory species to quantify the behavior of the different morphs in response to predators. In both experiments, secondary host morphs of P. bursarius showed no attacking behavior and antipredator behavior in these morphs was limited to escaping natural enemies by walking away. In contrast, the first instars of the primary host generations in both Pemphigus species showed attacking behavior and were capable of killing the predators.     

不同寄主植物上瓜蚜体内主要构成成分的变化

为揭示寄主植物与瓜蚜Aphis gossypii相互作用的机理, 采用微量凯氏定氮法、索氏提取法、蒽酮比色法、盐酸水解法和气相色谱法分别测定了黄瓜、南瓜、搅瓜、瓢葫芦和哈密瓜等5种寄主植物对瓜蚜体内粗蛋白、脂肪、糖、氨基酸和脂肪酸等主要构成成分的影响。结果表明, 不同寄主植物上瓜蚜体内上述主要构成成分均存在显著差异; 瓜蚜体内主要构成成分以粗蛋白含量最高, 占虫体干重的35.40%~45.25%, 其次为脂肪（24.01%~30.33%）, 最低为总糖（4.09%~7.91%）. 瓜蚜体内含有18种氨基酸, 其中谷氨酸含量最高（5.15~6.97 mg/100 mg）, 其次为天门冬氨酸（3.75~5.33 mg/100 mg）, 再次为亮氨酸（2.59~3.60 mg/100 mg）, 最少为半胱氨酸（0.44~0.49 mg/mg）。瓜蚜体内含有8种脂肪酸。 除瓢葫芦外, 其他4种供试寄主植物上瓜蚜体内棕榈酸含量最高（10.53%~15.55%）, 其次为肉豆蔻酸（3.32%~9.93%）, 亚麻酸含量最低（0.04%~0.17%）。瓢葫芦上瓜蚜体内肉豆蔻酸含量最高, 为16.20%, 其次为棕榈酸（7.15%）。瓜蚜体内饱和脂肪酸含量占虫体干重的20.08%~27.72%, 而不饱和脂肪酸只占2.58%~3.84%。结果提示寄主植物对瓜蚜体内主要构成成分均有显著的影响。     

Temperature and host plant effects on development and fecundity of Brevicoryne brassicae (L.) (Homoptera: Aphididae)

The development, survival and reproduction of the cabbage aphid, Brevicoryne brassicae (L.) were evaluated at three constant temperatures (20, 25 and 30°C) on cabbage, cauliflower, red cabbage, turnip and radish. The development periods of immature stages ranged from 10.7 d at 20°C to 7.60 d at 30°C for red cabbage. Total percentages of survivorship of immature stages varied from 39.40 and 82.50 within the temperature range of 25–30°C on radish. The average progeny per female was 31.15, 28.95 and 23.77 at 20, 25 and 30°C on cabbage.     

Genotypic and phenotypic variation in transmission traits of a complex life cycle parasite

Characterizing genetic variation in parasite transmission traits and its contribution to parasite vigor is essential for understanding the evolution of parasite life‐history traits. We measured genetic variation in output, activity, survival, and infection success of clonal transmission stages (cercaria larvae) of a complex life cycle parasite (Diplostomum pseudospathaceum). We further tested if variation in host nutritional stage had an effect on these traits by keeping hosts on limited or ad libitum diet. The traits we measured were highly variable among parasite genotypes indicating significant genetic variation in these life‐history traits. Traits were also phenotypically variable, for example, there was significant variation in the measured traits over time within each genotype. However, host nutritional stage had no effect on the parasite traits suggesting that a short‐term reduction in host resources was not limiting the cercarial output or performance. Overall, these results suggest significant interclonal and phenotypic variation in parasite transmission traits that are not affected by host nutritional status.     

Diurnal Patterns in Host Finding by Potato Aphids, Macrosiphum euphorbiae (Homoptera: Aphididae)

Potato aphid, Macrosiphum euphorbiae (Homoptera: Aphididae), is an abundant potato pest and vector of potato leaf-roll virus and potato virus Y in Maine and other potato growing areas. We investigated the circadian rhythmicity of its movement towards host plant odor. Effects of daily cycle (day or night) and illumination (light or dark) on the proportion of aphids colonizing potato leaflets were determined in a Petri plate arena and in a Y-tube olfactometer. In Petri dishes, both daily cycle and light had a highly significant effect on plant colonization. Increasing temperature reduced aphid colonization of the leaflets. In the olfactometer, light had a significant effect on the proportion of aphids walking towards the host plant. Interaction between time and light was also statistically significant, with the effect of illumination being smaller during the day than during the night. Our results suggest that circadian rhythm in host-finding behavior of the potato aphid is regulated by both exogenous and endogenous mechanisms.     

温度对三叶草彩斑蚜种群参数的影响

为了探讨温度对三叶草彩斑蚜种群增长的影响,在实验室研究了15～35℃范围内共9个温度下三叶草彩斑蚜的发育、繁殖和生命表.结果表明:三叶草彩斑蚜种群在35℃下不能存活.若虫的发育历期随着温度的上升而显著缩短,在15 ～ 32℃内为18.33～4.02 d,存活率在40.0％～83.6％,25℃下若虫的存活率最高、32℃下存活率最低.成蚜的平均寿命在10.64 ～ 20.87 d,23℃下寿命最长、32℃下最短.产蚜高峰期随温度的上升而提前,除15℃为15 d以外,其余温度下均在3～6d.平均繁殖力和单头最高繁殖力以25℃下最高,分别为82.0和149.0头.平均世代周期随着温度的上升从15℃的31.17 d逐渐缩短到32℃的10.17 d.净增殖率在25℃下最大(68.62),32℃下最小(13.96).内禀增长率在0.10 ～0.30 d-1,其中28 ℃下最高、15℃下最低.若虫的发育起点温度和有效积温分别为9.35℃和97.83 d·℃.繁殖力、净增殖率和内禀增长率与温度的关系均可用一元二次方程描述.     

An interval timer controls the production of sexual morphs in Myzus persicae (Homoptera: Aphididae)

Abstract The interval timer mechanism was examined in two clones of Myzus persicae (Sulzer) originating from peach in Naousa, North Greece. Overwintering eggs were collected in April 1997 and after egg hatch the morphs of progeny were recorded at LD 10 : 14 h and 17 °C until the appearance of sexuals. In one clone, observations were continued until the 21st generation after egg hatch. In addition, the production of sexual morphs was examined in two other clones originating from Lehonia, Central Greece, when apterae (wingless females) were transferred from long to short days and 17 °C in the second and eighth generation of parthenogenetic rearing after their collection from peach trees. In clones from Naousa, the first gynoparae appear and comprise 10% of alate (winged) females in the second half of the third generation after egg hatch. The percentage of gynoparae increased in each consecutive generation, reaching 100% in fifth and seventh generations in clones Nb and Na, respectively. First males appeared later than gynoparae, in fifth and sixth generation in clones Nb and Na, respectively. In clone Nb, the number of males, after their appearance in fifth generation, was relatively constant from the sixth to the ninth generation (10.8–12.3). Then a significant increase was observed in the 10th generation, thereafter remaining relatively constant until the 21st generation (18.1–19.8). A similar trend in the number of males produced was observed in clones deriving from Lehonia.     

Choice of host plant as a factor in reproductive isolation of the aphid genus Cryptomyzus (Homoptera, Aphididae)

Abstract. 1. Host plant preference experiments were conducted with closely related taxa of the aphid genus Cryptomyzus. Males, and presexual morphs (sexuparae and gynoparae), were used to determine the impact of host plant choice on reproductive isolation. In the case of host-alternating species these morphs are migratory and so will select the host plant.
2. Host plant preference of two closely related taxa of C. alboapicalis (Theobald) was found to promote their reproductive isolation. The preference of sexuparae of these monoecious taxa was more pronounced than that of the males.
3. Host plant preference and subsequent production of oviparae showed that C. galeopsidis (Kaltenbach) consists of two host races restricted to Ribes rubrum L. and R. nigrum L., respectively. The existence of clones, intermediate in their preference and reproductive performance on these plants, suggests that hybridization occurs.     

Life cycles of three Stomaphis species (Homoptera: Aphididae) observed in Kyoto,Japan: possible host alternation of S. japonica

Aphids of the genus Stomaphis, which have a very long stylet, are parasitic on tree trunks and feed on phloem through thick xylem. Stomaphis japonica on Quercus acutissima, Q. serrata and Q. variabilis; S. yanonis on Celtis sinensis and Zelkova serrata; and S. malloti on Mallotus japonicus were surveyed every season from 1981 to 1998 at six sites situated in natural forests in Kyoto, Japan. From fragmentary observations, their life cycles were inferred and possible host alternation of S. japonica is discussed. Stomaphis yanonis and S. malloti are monoecious and holocyclic in Kyoto, as has also been observed near Tokyo and at Ise city, Mie Prefecture, respectively. Stomaphis japonica was found throughout the year on Q. serrata passing through a holocycle, but was found only from late May to mid‐November on Q. acutissima and Q. variabilis, where neither oviparae nor males occurred. Alatae flew to Q. acutissima and Q. variabilis in late May to mid‐June and settled there. However, all virginoparae emerged as alatae from October and left these trees in mid‐October to early November. These observations suggest that Stomaphis japonica in this area comprise two sympatric populations: a monoecious population living on Q. serrata all year round and a heteroecious population alternating between Q. serrata as a primary host and Q. acutissima and Q. variabilis as secondary hosts.     

Host-dependent association and its implications on the relative abundance of blue alfalfa aphid and pea aphid (Homoptera: Aphididae) on alfalfa in Oklahoma

Field populations of blue alfalfa aphid and pea aphid on alfalfa were sampled during 1985 and 1986 to determine the association of co-occurrence, interspecific interactions and comparative temporal variations in the spatial dispersion patterns of these species in Oklahoma. Relative abundance of these species is discussed in the light of above analyses.Cole' s coefficient revealed a high degree of association between these species in terms of their occurrence on the same alfalfa stems in the field. Regression analyses indicated that the species populations tended to increase in concert on the same stems without evidence of competitive displacement. Spatial dispersion patterns of both species were highly aggregated at low population densities early in the season. Over time, both species tended to disperse and became less aggregated as numbers increased. It was concluded that magnitude of interspecific interactions between the blue alfalfa aphid and the pea aphid were not of a nature that they could be termed as competing species. On the contrary, a concept of an “ecospecies” is proposed for practical applications such as sampling plans and economic threshold determinations.     

寄主作物对B型烟粉虱生长发育和种群增殖的影响

在恒温28℃、湿度80％的条件下研究了5种寄主植物对B型烟粉虱(Bemisia tubaci)生长、发育和繁殖的影响。结果表明：取食棉花、烟草、花生、大豆和玉米的烟粉虱。其形态、发育历期、存活率、成虫繁殖力和寿命等生命参数均有明显差异。棉花、大豆和玉米不利于烟粉虱卵和若虫的生长发育。表现为虫体较小、卵期和生命周期延长、存活率降低。烟粉虱在棉花、大豆、花生、烟草和玉米上的生命周期分别为32．03d、32．11d、25．69d、24．43d和20．68d。其存活率分别为49．86％、54．41％、86．86％、69．93％和29．38％。与烟草和玉米相比。棉花、大豆和花生明显有利于烟粉虱的种群繁殖．在棉花、大豆、花生、烟草和玉米上生长发育的成虫寿命和单雌产卵量分别是27．8d和235．O粒、23．2d和191．1粒、22．Od和131．1粒、6．25d和28．O粒、2．42d和5．1粒。在花生、大豆、棉花、烟草和玉米上的内禀增长率(rm)分别为O．1590、O．1364、O．1236、O．0841和-O．0285．其种群趋势指数(I)分别为113．85、117．38、103．98、19．58和O．4274。     

Horn polyphenism and related head shape variation in a single-horned dung beetle: Onthophagus (Palaeonthophagus) fracticornis (Coleoptera: Scarabaeidae)

Horns of Onthophagus beetles are typical examples of phenotypically plastic traits: they are expressed as a function of environmental (nutritional) stimuli, and their reaction norm (i.e. the full set of horn lengths expressed as a response to different degrees of nutritional states) can be either linear or threshold-dependent. Horned males of Onthophagus ( Palaeonthophagus ) fracticornis (Preyssler, 1790) bear a single triangular cephalic protrusion of vertex carina, which has received phylogenetic support as the most primitive horn shape in the genus. Inter- and intra-sexual patterns of horn expression were studied in O. fracticornis by means of static allometries while associated variations in head shape were assessed using geometric morphometric techniques. The relation between log-transformed measurements of body size and vertex carina supported an isometric scaling in females. On the contrary, a sigmoidal model described better the horn length-body size allometry in males, with a switch point between alternative morphs at a pronotum width of 3.88 mm. Sigmoidal static allometries of horns in Onthophagus populations arise from a threshold-dependent developmental process of horn growth. This process underlies the expression of both plesiomorphic and apomorphic horn shapes in the genus. Given that the single-horn model has been identified as primitive, we propose that such a developmental process giving rise to it may be evolutionarily ancient as well. Horn expression was accompanied by a deformation of the head which makes minor and major morphs appear even more different. Therefore, in this species both horn and head shape expression contribute to male dimorphism.     

Presence of two host races of Aphis gossypii Glover (Hemiptera: Aphididae) collected in Turkey

The aphid, Aphis gossypii, is a primary pest of citrus, cotton, cucurbits and greenhouse‐grown vegetables in Turkey and throughout Europe. There is some previous empirical data suggesting that host‐adapted genotypes of this aphid exist which may in fact be host‐races. To determine if host races of A. gossypii are indeed present in the eastern Mediterranean region of Turkey, reciprocal host transfer experiments and life table analyses were performed with multiple asexual lineages (= clones) of the aphid collected from different hosts. The collection hosts included citrus, cucumber, eggplant, okra, sweet pepper and cotton. Aphid developmental times on the host from which the aphid was originally collected (= collection or natal host) were shorter (5.2–6.0 days) and had a higher intrinsic rate of population growth (r_m = 0.25–0.44) than the 6.6–7.3 days required when the aphid was reared on a non‐original collection host (= non‐collection host or non‐natal host) and had r_m = 0.03–0.30. Total immature mortality of the cotton clone, especially in the first nymphal stage, was high (51–100%) with low r_m (0–0.03) on cucumber, citrus and sweet pepper. Aphid populations transferred from citrus, eggplant and okra to cotton (r_m = 0.29–0.30) did not differ significantly in their performance from that of the cotton population on cotton (r_m = 0.34), whereas that from sweet pepper and cucumber populations (r_m = 0.22–0.24) were significantly lower. These data have allowed us to separate A. gossypii into two distinct biological groups: (a) a ‘generalist’ population obtained from cucumber, sweet pepper, citrus, eggplant and okra which exhibited statistically better development on cotton; versus (b) a population from cotton which, by comparison on reciprocal hosts, developed poorly on non‐natal hosts except on eggplant. Development of the cotton clone on cucumber and okra was not improved after four successive generations on the non‐natal host. The good development of A. gossypii from eggplant and cotton on these reciprocal hosts suggests that these particular clones were similar, if not identical, host races.