Polymorphic fundatrices in thimbleberry aphid — ecology and maintenance

The thimbleberry aphid,Masonaphis maxima (Mason) lives on patches of plants that support 3,4 or 5 generations depending on site and weather. The life cycle requires sexual females and males to produce overwintering eggs. The eggs hatch in the spring to produce the first ’fundatrix’ generation; subsequent generations are produced parthenogenically. Males and other morphs are produced by wingless virginoparae, but sexual females are produced by ’gynoparae’, a winged morph that is specialized to produce only sexual females. The fundatrices have no indication of the number of generations that the plants will support in the current year. There are two fundatrix types that coexist in different ratios depending on the number of generations supported by the patch the previous year. One type produces sexual females in generations 3 and 5, and males in generations 4 and 5; the other type produces sexual females in generations 4 and 5, and males in generations 3, 4 and 5. The dimorphism adapts the aphid to its heterogeneous and somewhat unpredictable environment. The role of sex in the maintenance of the dimorphism is discussed. This is the first report of fundatrix polymorphism and consequent differential sex expression in aphids.     

Life-cycle and morph production in the Arctic aphid Acyrthosiphon brevicorne

 The life-cycle of the sub-Arctic, Dryas-feeding aphid, Acyrthosiphon brevicorne, is described and shown to be intermediate between that of the high-Arctic aphid (Acyrthosiphon svalbardicum) and typical holocyclic, monoecious, temperate species. The fundatrix of A. brevicorne can give rise directly to both sexual morphs, a necessary adaptation to the short Arctic summer. Production of males is predetermined; some males are produced by all-viviparous mothers, but at lower frequencies in the birth sequences of the fundatrix than in subsequent generations. By contrast, the production of oviparae is controlled, at least in part, by photoperiod. Received: 19 April 1994/Accepted: 4 August 1995     

越冬代梨木虱、梨二叉蚜产卵选择对子代种间竞争的影响

梨树萌芽期,越冬代梨木虱Cacopsylla chinensis和梨二叉蚜Schizaphis piricola子代共用同一食物资源,为了解其母代如何避免子代发生种间竞争,通过研究其产卵特点表明,梨二叉蚜和梨木虱在梨树枝条上的产卵位点不同。梨木虱主要在花芽枝上进行产卵,偏好于花芽芽鳞和枝条基部刻痕处产卵,以枝条基部产卵量最多,其次为顶花芽,产卵量随芽位次序依次下降,梨木虱还可在叶芽部位产卵,但数量较少;梨二叉蚜主要于叶芽枝上进行产卵,偏好在叶芽枝的第2、3芽位产卵,顶芽和枝条基部无产卵,未见梨二叉蚜在花芽部位产卵;在花芽枝上,梨木虱卵的空间生态位大于梨二叉蚜、叶芽枝上却相反,两者在梨树枝条上的产卵空间生态位呈分离状态;梨木虱和梨二叉蚜孵化的若虫数量及空间分布特点与卵一致;对梨木虱和梨二叉蚜产卵位点分析表明,其产卵位点的物理性状明显不同。本研究明确了梨木虱和梨二叉蚜母代通过不同产卵位点的选择,降低了子代种间的同位竞争。     

Wing and flight muscle polymorphism in a lygaeid bug, Horvathiolus gibbicollis: determinants and life history consequences

ABSTRACT.

• 1 Horvathiolus gibbicollis (Costa), a ground-living seed-feeding bug of the mediterranean region, has two wing morphs. In macrop-terous bugs both pairs of wings are fully developed. In brachypterous ones forewings are reduced to about two-thirds and hindwings to less than a third of their length in macropters.
• 2 Each morph bred true with regard to wing length when reared under variable density, food and temperature conditions for several generations.
• 3 All F₁ offspring between crosses of the two morphs were brachypterous. In F₂ approximately 25% were macropters and 75% brachypters implying monogenic control of wingform.
• 4 Flight muscles in macropters vary from fully developed to totally reduced. This variation is determined by environmental conditions during adult life. Most young adult bugs have flight muscles, and totally starved or unmated bugs retain their flight muscles. Fed and mated females histolyse flight muscles as they start laying eggs, while most males of the same group retain their muscles.
• 5 Brachypterous bugs have a smaller thorax and larger abdomen than macropterous ones.
• 6 Brachypterous bugs reach adulthood slightly before macropterous ones, and they have a distinctly shorter adult preoviposition period.
• 7 Lifetime egg production does not differ significantly between the two wing morphs. However, the temporal pattern of egg laying is different in the two morphs. The m_x-curve of macropters starts later, then attains a higher peak and finally decreases faster than that of brachypters.
• 8 Initially, macropters lay smaller eggs than brachypters, but egg volume increases with age in macropters and eventually approaches that of brachypters.
• 9 The initial increase in reproductive effort (egg volumexegg number) of macropters is concomitant with wing muscle histolysis and the mobilization of thorax space for reproduction.
• 10 Adult survival rate does not differ between the morphs.

     

Autumnal emergence of Anagrus wasps,egg parasitoids of Empoasca vitis,from grapevine leaves and their migration towards brambles

• 1 The Anagrus‘atomus’ parasitoid group (Hymenoptera: Mymaridae), associated with Empoasca vitis (Göthe) (Homoptera: Cicadellidae), overwinters on vegetation surrounding vineyards. The emergence of parasitoid adults from grapevine leaves in autumn was studied in north‐eastern Italy, both in relation to the E. vitis egg‐laying period and to the presence of leafhoppers overwintering as eggs on Rubus bushes.
• 2 Autumnal peaks of Anagrus captured using yellow sticky traps were observed first on grapevines and then on brambles. Parasitoid captures in vineyards were observed for more than 1 month after the last first‐instar nymphs of the grape leafhoppers were noticed. Two species belonging to the A. ‘atomus’ group, Anagrus atomus and Anagrus ustulatus, were captured both on grapevines and brambles.
• 3 Parasitoids of the A.‘atomus’ group can emerge from third‐generation grape leafhopper eggs in accordance with two different development time patterns (i.e. normal or delayed). Individuals with delayed emergence required up to 2.2‐fold more time to develop from an egg to adult than individuals with normal emergence. This meant that some parasitoid adults emerged in autumn from eggs of grape leafhopper laid in August.
• 4 A delayed emergence as a result of a slower development ensures that the A.‘atomus’ parasitoid group is synchronized with the egg‐laying of leafhoppers that overwinter as eggs on Rubus spp.
• 5 Consequently, leafhoppers overwintering as eggs on brambles play a key role in the ecology of the relationship between grape leafhoppers and the A.‘atomus’ parasitoid group.

     

Spatial and temporal patterns of predation by ants on eggs of Cactoblastis cactorum

ABSTRACT.

• 1 Predation of Cactoblastis cactorum (Berg) (Lepidoptera: Pyralidae) eggs by ants was studied for a summer and a winter generation on prickly pear Opuntia ficus-indica (L.) Miller plants in the field.
• 2 Egg Predation between plants was spatially density-dependent in the winter but not in the summer generation.
• 3 To test for the presence of delayed density-dependent mortality within generations, percentage egg Predation during 5-day periods was plotted against the number of eggs available in each period. Data points joined in a time sequence moved in an anticlockwise direction in the summer generation, suggesting a delayed density-dependent response by the ants, whereas in the winter generation the points moved in a clockwise direction.
• 4 The degree of coincidence of the C. cactorum eggs with the peak larval production periods of the ants is suggested as the main reason for the temporal changes in egg Predation.

     

Overwintering of reproductively mature females of a pro‐ovigenic parasitic wasp,Leptopilina heterotoma (Hymenoptera: Figitidae)

Leptopilina heterotoma is a Drosophila parasitoid mainly occurring in temperate regions, and females of this species are pro‐ovigenic (i.e. all or nearly all of their lifetime eggs are already mature at emergence). Here I investigated how L. heterotoma overwinters in Sapporo, Japan, a cool temperate region, by outdoor and laboratory experiments. Females of this species had mature eggs at emergence as expected, and they did not resorb eggs even if they were exposed to cold or starvation. Nevertheless, females were able to overwinter, and post‐overwintering females retained parasitization capacity. In this species, thus, adult overwintering is not associated with reproductive diapause. However, females grown in late autumn or at low temperatures (e.g. 5°C) had a fewer number of mature eggs and higher winter survival. At low temperatures, female larvae could save energy by reducing egg production and allocate the saved energy to expenditure for overwintering. In contrast to females, males and pre‐adult individuals were not able to overwinter.     

桃蚜不同蚜型DNA多态性的RAPD研究

采用RAPD方法,对全周期桃蚜的有翅产雌性母蚜、无翅性母蚜、雄蚜、卵、干母、干雌、有翅迁移蚜等蚜型的DNA遗传多态性进行了分析。结果表明：卵的DNA多态性最大,性蚜次之,孤雌生殖蚜最小;卵与其它蚜型之间在遗传上具最大差异,其中与孤雌生蚜的差异大于与性蚜的;干母、干雌和迁移蚜之间的遗传关系最近。     

Inducing the sexual forms and hatching the eggs of pea aphids

In temperate climates, pea aphids (Acyrthosiphon pisum) produce a single sexual generation each year in response to declining photoperiod and temperature. Mating occurs in the fall and the eggs have an obligatory winter diapause. Genetic recombination during the sexual phase is thought to be an important source of genetic variability within cyclically parthenogenetic aphid populations. Methods for reliably producing sexual forms and hatching the eggs of aphids are therefore central not only to the study of evolutionary change in aphid populations, but also for a general understanding of the origin of agriculturally important variation in destructiveness within pest species.Here, sexual forms of six pea aphid clones were induced in the laboratory and eggs were successfully hatched by creating conditions that closely mimicked those found in field situations. A declining photoperiod was produced by controlling artificial lighting using a timer with variable cycle length. Using these conditions, sexual forms were successfully produced for all six clones tested, which were then mated in all combinations. Eggs were exposed to a daily cycle of freezing and thawing in an incubator under a short-day photoperiod. Egg hatch averaged 60%, but was as high as 89% for some crosses. These methods will permit testing of evolutionary hypotheses and execution of detailed genetic studies of sources of variability within pea aphid populations. They are thus important tools for both evolutionary and agricultural studies.     

Temporal changes in host adaptation in the pea aphid, Acyrthosiphon pisum

Abstract.

• 1 Temporal changes in host adaptation were followed in a local population of the pea aphid, Acyrthosiphon pisum. Aphid clones were collected in one alfalfa and one clover field at three different times. In the spring, first-generation females were collected. Later, in the autumn, females belonging to the last parthenogenetic generation were collected. Lastly, sexual females were collected after mating in autumn and allowed to produce eggs which were hatched. The performance was evaluated on alfalfa and clover. The spring-collected individuals were also assessed on peas.
• 2 On the overwintering hosts clover and alfalfa, the clones performed best on the plant of origin, i.e. negative correlations in performance. Correlations between performance on the temporary summer host, pea, and that on clover/alfalfa were weak or nonsignificant.
• 3 Significant variation in host performance was found within both host fields at spring, which is a prerequisite for changes in clone composition due to selection/migration.
• 4 The clones from alfalfa showed an increase in mean performance on alfalfa between spring and autumn, whereas no changes among the clones from the clover field were observed. This difference in seasonal response between the two fields could have been the result of larger variation in performance among the alfalfa clones and/or a differential tendency to migrate among clones in both fields.
• 5 After sexual recombination in the autumn, mean performance in the alfalfa field returned to the spring level, probably as a result of emergence of new genetic combinations. In the clover field, mean performance did not change significantly over time.

     

Embryonic stage of obligatory diapause and effects of abiotic conditions on egg hatching in the balsam twig aphid,Mindarus abietinus

Diapause‐mediated dormancy in overwintering insect eggs has rarely been studied with regard to the ecological factors controlling postdiapause development. In insects of temperate latitudes, water availability at the end of winter, in interaction with temperature, could control the resumption of development for insect stages in postdiapause quiescence. The balsam twig aphid, Mindarus abietinus Koch (Hemiptera: Aphididae), overwinters as eggs in southern Québec, Canada, on balsam fir, Abies balsamea (L.) Miller (Pinaceae), in Christmas tree plantations, where it is known as a pest. Previous work has shown that eggs of this aphid maintain low water content during winter, presumably to survive sub‐zero temperatures. Conversely, in late winter and early spring, they passively or actively absorb surrounding moisture, which is accompanied by notable changes in size, shape, and fresh mass. The primary objective here was to determine the embryonic stage at which winter diapause starts and is maintained in M. abietinus, a relatively primitive aphid. Secondly, we tested the hypothesis that free water availability to postdiapause eggs, in combination with temperatures above developmental threshold, is essential for embryonic development and hatching, by experimentally soaking field‐collected eggs in water at controlled frequencies. We observed that embryogenesis starts at the time of egg laying and stops after a few days, before the anatrepsis stage of blastokinesis is complete, when the germ band has not yet entirely immersed itself into the yolk. We also found that water surrounding overwintered eggs on fir shoots, in interaction with temperature regime, significantly increases M. abietinus egg hatching rates. Potential impacts of environmental factors such as precipitation are discussed in relation to M. abietinus egg hatching rates and potential for population growth in spring.     

A transgenerational interval timer inhibits unseasonal sexual morph production in damson-hop aphid, Phorodon humuli

Abstract.  The induction of sexual and parthenogenetic morphs of the damson-hop aphid, Phorodon humuli , on hops is controlled by daylength. The ability of P. humuli , to produce winged pre-sexual females (gynoparae) in the short-day conditions of spring is inhibited by an interval timer present in generations immediately after hatching of the overwintering egg. The inhibition expires after three generations when nymphs are born and reared in short days (LD 12 : 12 h), irrespective of whether their parents are reared in short or long days (LD 18 : 6 h). No gynoparae are produced by aphids maintained for 13 generations in long days. Two wingless aphids from 35 survive transfer from Prunus spinosa to hops. No winged females are produced during nine generations among their progeny maintained in long days on hops, but gynoparae, followed by males, are produced one generation after these aphids are transferred to short days.     

Relating rainfall and vegetation greenness to the biology of spur‐throated and Australian plague locusts

• 1 Rainfall and vegetation greenness are widely claimed to influence fat content and egg development in locusts. Body size, abdominal fat and egg length of spur‐throated Austracris guttulosa and Australian plague Chortoicetes terminifera locusts were related to the normalized difference vegetation index (NDVI; a greenness indicator) to test these assumptions and quantify biological responses.
• 2 Cumulative rainfall was highly correlated with NDVI values of an area (25 × 25 km) surrounding the trap to which locusts were attracted. Catches of locusts were greater in hotter periods preceding rain, associated with an increase in NDVI indicative of brown (0.298) versus green (0.465) vegetation.
• 3 Pre‐overwintering descendent A. guttulosa were larger and fattier than post‐overwintering antecedents. Egg maturation of antecedents occurred coincident with increasing NDVI. Higher abdominal fat in pre‐overwintering locusts was positively associated with higher NDVI.
• 4 Male C. terminifera of two descendent generations were larger than their antecedents but only pre‐overwintering locusts were significantly fattier than their predecessors. Only pre‐overwintering females were significantly larger than their predecessors and no generation was significantly fattier than any other. Fat content was negatively correlated with egg maturation and differed significantly with NDVI, although strong and consistent relationships were not obtained.
• 5 The findings suggest that locust–environment interactions are species‐ and habitat‐specific. If A. guttulosa hoppers develop during periods when regional NDVI exceeds 0.4, fatter adults will arise. The elucidation of relationships between greenness, fat and egg development in C. terminifera is unlikely using NDVI values from areas comprising mixtures of monocotyledonous and dicotyledonous vegetation.

     

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.     

Overwintering of the sycamore aphid, Drepanosiphum platanoidis

Spatial and temporal aspects of overwintering in the holocyclic sycamore aphid Drepanosiphum platanoidis (Schrank) (Homoptera: Callaphididae) were investigated. Eggs of D. platanoidis were principally laid on trunks and branches of Acer pseudoplatanusL., at considerable distances from the buds. Trunks with rough bark, followed by branches with moderate roughness supported more overwintering eggs than the relatively exposed smooth surfaces of twigs and terminal buds. Eggs were aggregated on rough bark. Aspect had no effect on the number of eggs laid. Experimental manipulations varying ovipara densities showed intra-specific competition among oviparae for optimal oviposition sites. Overwintering egg mortality was a function of time, with the greatest mortality rate occurring in late winter. Egg mortality varied from 80.4% to 76.9% on trunks and 83.0% to 65.9% on branches in 1996 and 1997.     

Formation of long‐lasting galls by overwintered nymphs in the Japanese aphid Quadrartus yoshinomiyai (Hemiptera: Aphididae: Hormaphidinae)

This paper examines the life history of a generation of galls created by the aphid Quadrartus yoshinomiyai (Hormaphidinae: Nipponaphidini) on its primary host plant, Distylium racemosum. First‐instar fundatrix nymphs of Q. yoshinomiyai initiated galls on stems of developing shoots in early April and incipient enclosed galls were found from later the same month. The galls lasted for up to 14 months, during which they grew to maturity, opened in early or mid‐April of the following year and dried up by the end of June. First‐instar fundatrix nymphs were found on winter buds, indicating that they hatched from eggs in autumn and overwintered as nymphs. These results suggest that Q. yoshinomiyai has a three‐year life cycle.     

Symbionin,an aphid endosymbiont-specific protein—III: Symbionin present in the male,ovipara and fundatrix

Electron microscopic observations demonstrated that the male of the kondo aphid, A. kondoi harbors intracellular symbionts different in shape from those in the viviparous female. Two-dimensional gel electrophoresis indicated that the endosymbiont in the male is less active in synthesizing symbionin, an aphid endosymbiont-specific protein than that in the viviparous female. Symbionin was also found in the winter egg though it was much less in amount than proteins related to the yolk formation. In the fundatrix which hatches out of the fertilized winter egg, symbionin was the most abundant protein.     

用微卫星引物PCR分析棉蚜不同蚜型的DNA多态性

用微卫星引物PCR方法, 对棉蚜Aphis gossypii干母、干雌、孤雌蚜(有翅迁飞孤雌蚜和无翅孤雌蚜)、性母、性雌蚜和雄蚜的DNA多态性进行分析。 结果表明：① 棉蚜的有性蚜型和无性蚜型之间在遗传上有较大的差异; ② 性母与性雌蚜和雄蚜的遗传关系都十分接近,说明同一性母既产性雌蚜,也产雄蚜;③ 干母和干雌的遗传关系很近, 但孤雌蚜已与二者有分化。     

Monitoring the European pine sawfly with pheromone traps in maturing Scots pine stands

Abstract

• 1 During 1989–93, field studies were conducted in Finland to develop a method based on pheromone traps to monitor and forecast population levels of the European pine sawfly (Neodiprion sertifer Geoffr.) and tree defoliation.
• 2 Three traps per site were baited with 100 µg of the N. sertifer sex pheromone, the acetate ester of (2S,3S,7S)‐3,7‐dimethyl‐2‐pentadecanol (diprionol), in maturing pine stands in southern and central Finland. In addition, three different dosages (1, 10 and 100 µg) of the pheromone were tested in 1991–92.
• 3 The highest number of males was observed in traps baited with the highest dose. On average, there was a 10‐fold increase in trap catch between lure doses.
• 4 Density of overwintering eggs was used to evaluate the effectiveness of pheromone traps in predicting sawfly populations. The proportion of healthy overwintering eggs was determined each year. A model based on the number of current shoots on sample trees, diameter at breast height and tree height was formulated to estimate eggs per hectare.
• 5 Linear regression analysis produced high coefficients of determination between number of males in traps and density of total eggs in the subsequent generation, when populations were at peak densities. The relationships were not significant for low population densities. The results indicate a risk of moderate defoliation when the seasonal trap catch is 800–1000 males per trap or higher.