Are thermal constants constant? A test using two species of ladybird.

There is a controversy about whether the thermal constants, lower developmental threshold, rate of development and corresponding degree days required for development, change when a species is reared under different developmental conditions. We present a more precise way of measuring these constants using the linear relationship between the rate of development and temperature. First we use the equation proposed by Ikemoto and Takai (2000) to determine the linear phase of development and then a generalised linear model having a different variance at low and high temperatures, specific for each condition, to estimate the parameters of the linear relationship. Using this method, we show that providing the difference in food quality is sufficiently great, an aphidophagous ladybird develops significantly faster and starts developing at a significantly lower temperature on a good than on a poor quality diet. Adaptive significance of the thermal constants not remaining constant is discussed in terms of a trade-off between growth and rate of development, when temperature and food quality varies.

The effect of relatedness on the response of Adalia bipunctata L. to oviposition deterring cues.

Larvae of aphidophagous ladybirds leave a cue in their tracks that deters oviposition. The influence of relatedness on this behaviour is for the first time explored in this paper. Two-spot ladybird females (Adalia bipunctata L.) under different conditions (young and naive, young and experienced, and old and naive) were exposed either to (i) clean filter paper, (ii) filter paper contaminated with unrelated larval tracks or (iii) filter paper contaminated with related larval tracks from their offspring. Oviposition time was recorded during nine hours. Oviposition was inhibited by larval tracks, and was more so by related than unrelated ones. Experienced females showed the same behaviour but with lower intensity. With old females, response was not significant with any type of larval tracks. This is the first report of the effect of relatedness on a ladybird's response to larval tracks.

Wheat antixenosis, antibiosis, and tolerance to infestation by Delphacodes kuscheli (Hemiptera: Delphacidae), a vector of "Mal de Río Cuarto" in Argentina.

"Mal de Rio Cuarto", is the most important virus disease of corn, Zea mays L., in Argentina. It is caused by the Mal de Rio Cuarto virus (family Reoviridae, genus Fijivirus. MRCV), which is a persistent virus transmitted by Delphacodes kuscheli (Fennah 1955) (Hemiptera: Delphacidae). Because corn is not a natural host of D. kuscheli, it has little protection from this pest. In contrast, wheat, Triticum aestivum L., is one of the main hosts of this vector and a reservoir of MRCV. The aim of this work was to identify genes involved in antixenosis, antibiosis, and tolerance of infestation by D. kuscheli in wheat, which might be used to reduce the population level of this vector on corn. A set of recombinant dihaploid (RDH) lines for chromosome 6A derived from the F1 cross between 'Chinese Spring' (CS) X 'Chinese Spring (Synthetic 6A)' (S6A) substitution line, was used for mapping. The S6A parental line is resistant to the MRCV vector. Antixenosis, antibiosis, and tolerance were evaluated using conventional tests in controlled environmental conditions. Most of the RDH and S6A showed higher levels of antixenosis against D. kuscheli than the parental line CS. The RDH lines showed highly significant antibiosis in terms of the duration of first, third, and fifth nymphal instars, developmental time (days), survival and fecundity. There were highly significant differences in the tolerance to D. kuscheli based on the chlorophyll content of the first and second leaves, foliar area, and aboveground fresh and dry weights. The duration of the fifth nymphal instar and the developmental period were significantly associated with Xgwm1017 marker loci, located at 48 cM on 6AL. Another quantitative trait locus accounting for the variation in chlorophyll content of the first leaf was associated with the interval between loci Xgwm459 and Xgwm334a, located in the telomeric region of the 6AS chromosome arm. The alleles with positive effects came from S6A. Antibiotic resistance of RDH could be useful for controlling the population increase of the MRCV vector on wheat, because prolonging the duration of development increases the period between two subsequent generations, so reducing the abundance of infective populations colonizing corn.

Sipha maydis: distribution and host range of a new aphid pest of winter cereals in Argentina.

Sipha maydis (Passerini) is a new aphid pest of cereals and cultivated and wild grasses in Argentina. This species was recently introduced into America, and nothing is known of its distribution or host range in South America. A better understanding of its biology is likely to facilitate control. This article records 1) the distribution and 2) the host range of S. maydis in Argentina. Over the period 2004-2006 samples were collected from 32 populations at several localities in Argentina. The number of S. maydis, accompanying aphid species, and the host from which they were collected were recorded. The distribution of S. maydis ranged from 32 degrees 52' to 42 degrees 03' S, and from 57 degrees 41' to 71 degrees 24' W, bounded by isothermals 18 and 10 degrees C and isohyets 200-400 and <1,200 mm. No S. maydis were found in the subtropical region, even in winter. In the field, the different populations showed very different host preferences. S. maydis was found mainly on cultivated barley and wheat and on wild Bromus spp. and Sorghum halepense (L.) Pers. No aphids were found on maize, Zea mays L. Most of the damage to winter cereal crops occurred at the seedling stage in early autumn and of adult plants when infestations occurred in late spring. In the 4 yr after the first record of S. maydis in Argentina, it colonized a huge area similar to that colonized by Diuraphis noxia (Mordvilko) in 10 yr. The wide range of regions, hosts and climatic conditions this species is adapted to is likely to make the control of this pest very difficult.

Comparing growth patterns among field populations of cereal aphids reveals factors limiting their maximum abundance.

Cereal stands in central Europe are commonly infested with three species of aphids that may become serious pests. With increasing abundance, the proportion of a particular species in the total aphid population may remain constant, suggesting a density-independent exponential growth, or the proportion can change, suggesting density-dependent constraints on growth. The constraints that affect particular species, and thus their relative abundance, were studied. The proportionality between maximum abundances of the cereal aphids was studied using a 10-year census of the numbers of aphids infesting 268 winter wheat plots. For two species their abundance on leaves and ears was compared. With increasing aphid density the maximum abundance of Rhopalosiphum padi (Linnaeus) remained proportional, but not that of Sitobion avenae (Fabricius), which was constrained by the smaller surface area of ears compared to leaves. There was no evidence of inter-specific competition. Maximum abundance of R. padi and Metopolophium dirhodum (Walker) on leaves did not change proportionally as the proportion of M. dirhodum decreased with increasing overall aphid density. This decrease was probably caused by the restricted distribution of M. dirhodum, which is confined to leaves, where space is limiting. No change in proportion between populations was detected when the average densities were below 0.54 aphids per leaf or ear. Non-proportional relationships between aphid populations appeared to be due to spatial constraints, acting upon the more abundant population. Detecting the limitation of population growth can help with the assessment of when density-independent exponential growth is limited by density-dependent factors. This information may help in the development of models of cereal aphid population dynamics.

Resource tracking in aphids: programmed reproductive strategies anticipate seasonal trends in habitat quality.

Most aphids show cyclical parthenogenesis, have short generation times and complete several generations each season. Two hypotheses have been proposed to account for the adaptive seasonal trends observed in the reproductive strategies of aphids. Firstly, individuals of each generation modify their reproductive strategy in direct response to the conditions they experience during their development. Secondly, the reproductive strategies of the different generations are to a large extent programmed and anticipate seasonal trends in habitat quality. These hypotheses were tested by rearing individuals of three generations of the host-alternating willow-carrot aphid, Cavariella aegopodii, on both willow and carrot. This revealed that the way this aphid allocates resources to gonads and lipid reserves is independent of an aphid's weight and the host plant on which it is reared. In addition each generation shows a specific relationship between offspring size and adult size, which tends to keep the absolute investment in individual offspring relatively constant from generation to generation, inspite of the big differences in adult size between generations. That is, through programmed allometric engineering aphids anticipate the predictable seasonal trends in habitat quality and so more closely track their resources, investing relatively more in gonads when food quality is high and relatively more in lipoidal reserves when food quality is poor.

Assessment of patch quality by ladybirds: role of larval tracks.

Gravid females of the two-spot ladybird, Adalia bipunctata (L.), were deterred from ovipositing when kept in petri dishes that had previously contained conspecific larvae but not conspecific adults, or the larvae of another two species of ladybird, Adalia decempunctata (L.) and Coccinella septempunctata L. The deterrent effect was density dependent and mediated via a chloroform-soluble contact pheromone present in the larval tracks. Similarly, gravid females of C. septempunctata were deterred from ovipositing by conspecific larval tracks and chloroform extracts of these tracks, but not by the tracks or extracts of tracks of A. bipunctata larvae. That is, in ladybirds the larvae produce a species-specific oviposition-deterring pheromone. In the field, the incidence of egg cannibalism in ladybirds increases very rapidly with the density of conspecific eggs or larvae per unit area. Thus, in responding to the species specific oviposition deterring pheromone female ladybirds reduce the risk of their eggs being eaten and spread their offspring more equally between patches.

Mate recognition in the two-spot ladybird beetle, Adalia bipunctata: role of chemical and behavioural cues.

On encountering a mature female, a male of the two-spot ladybird, Adalia bipunctata (L.), first palpated her elytra with his maxillary palps, then mounted her, extruded his penis and mated. Copulation never occurred between active males but males copulated with dummies bearing male elytra as frequently as with dummies with female elytra of their own species. Similarly, males attempted mating with immobilised conspecifics of both sexes. However elytra washed in chloroform failed to stimulate mating. Analysis of the chloroform extracts of the elytra revealed that male and female ladybirds are coated by the same blend of hydrocarbons among which 9- and 7-methyl tricosane are dominant. Our results are consistent with a role of these cuticulars hydrocarbons in species recognition and show that behaviour, in particular movement, is necessary for discrimination between males and females.

The role of food quality and competition in shaping the seasonal cycle in the reproductive activity of the sycamore aphid.

The hypothesis that seasonal changes in sycamore aphid,Drepanosiphum platanoidis (Schr.), recruitment are determined by changes in food quality and aphid population density was tested. There was no clear association between the reproductive activity of the sycamore aphid and the seasonal changes in specific amino acids or groups of amino acids in extracts of sycamore,Acer pseudoplatanus L., leaves. Seasonal changes in reproductive activity tracked the changes in total amino acids of the leaf tissue of the host, but with a short time delay. High numbers of adult aphids appeared to depress reproductive activity. A regression analysis of the results revealed that total amino acids the previous week and current numbers of adult aphids significantly affected sycamore aphid reproductive activity. The results of this analysis support the above hypothesis, that the marked seasonal changes in the total quantity of amino acids in sycamore leaves and intraspecific competition for this resource, through its effect on adult weight, shape the seasonal cycle in the reproductive activity of the sycamore aphid.

Attack strategy of ladybird beetles (Coccinellidae): factors shaping their numerical response.

Temporal changes in aphid abundance pose a considerable challenge to ovipositing aphidophagous ladybirds, as in order to maximize their fitness they need to synchronize their reproduction with the early development of aphid populations. Field census data and laboratory experiments were used to determine how ovipositing females of the two-spot ladybird, Adalia bipunctata (L.), assess whether an aphid population is suitable for exploitation. In the field, two-spot ladybirds usually laid eggs well before aphid populations peaked in abundance. In the laboratory they showed a marked reduction in their reproductive numerical response in the presence of larvae of their own species but not of other aphidophagous ladybirds. At the highest aphid density this was not a consequence of competition for food between larvae and ovipositing females. In the presence of conspecific larvae gravid females were very active and as a consequence more likely to leave an area, and when confined with other conspecific females or larvae laid fewer eggs and later than females kept on their own. The extent of the inhibition of egg laying is negatively correlated with the rate of encounter with larvae. Thus it is proposed that gravid females appear mainly to use the presence of conspecific larvae to assess the potential of an aphid colony for supporting the development of their offspring.

Temperature prediction and the timing of sex in aphids.

The aphid life cycle contains a series of parthenogenetic, viviparous generations, followed usually by a sexual generation that produces resistant overwintering eggs. Since the decision to produce sexually-reproducing offspring ends the period of rapid growth of the clone, it should be postponed as late as is compatible with successful oviposition. The time of leaf fall determines the latest possible time of oviposition, and is itself determined mainly by daylength. The time required for the development of the final generations of aphid depends on temperature. The decision to end the sequence of parthenogenetic generations should thus depend on temperature and photoperiod. This paper calculates the optimal combination of daylength and temperature for the cueing of this decision in Rhopalosiphum padi. It is shown that the experimental data of Dixon and Glen (1971) are in agreement with the predictions, and that, in the field, the time of production of male R. padi does vary adaptively with July temperature.

Plant structure and the searching efficiency of coccinellid larvae.

| 1. To determine the effect of plant structure on the searching efficiency of Coccinella septempunctata L. larvae, their functional response on pea and bean plants was compared. 2. The attack coefficient a was lower on pea than on bean plants. 3. This was not due to a difference in the coincidence of prey distribution and predator searching effort, but was due to larvae falling off the smooth leaves of pea plants significantly more frequently than off bean plants. 4. It was concluded that plant structure is an important factor in determining the quality of a habitat for coccinellids.

The effect of nutrient stress on life history parameters of the black bean aphid, Aphis fabae scop.

Experiments were conducted to investigate the effects of brief spells of starvation on the longevity and age-specific reproduction of A. fabae. Increasing the duration of the starvation period reduced the aphids' ability to resettle when returned to a host, but no longer-term effects on mortality were found. Secondly, as predicted, starvation reduced life-time fecundity, but increased the reproductive rate immediately after nutrition improved.

Migratory urge and reproductive investment in aphids: Variation within clones.

Winged aphids with a small reproductive investment take-off more readily and at a steeper angle, delay wing muscle autolysis for longer, are more resistant to starvation and have relatively more olfactory sense organs than aphids with a large reproductive investment.It is proposed that by means of such programmed tactically diverse but labile flight and reproductive strategies aphid clones are better able to exploit resources that vary both spatially and temporally.

Metabolic acclimatization to seasonal changes in temperature in the sycamore aphid, Drepanosiphum platanoides (Schr.), and lime aphid, Eucallipterus tiliae L.

Both sycamore and lime aphids live on the leaves of their respective host trees. The sycamore aphid unlike the lime aphid has the ability to adapt metabolically to changes in temperature. The difference in ability of these two tree-dwelling aphids to adapt metabolically to changes in temperature can be related to their mode of life. The sycamore aphid lives on sycamore which is native to the mountainous areas of southern and central Europe where conditions are cool. Sycamore also has a longer growth period than lime as its buds burst earlier and it sheds its leaves later. Sycamore aphids are therefore naturally exposed to a wider range of temperatures than lime aphids.

The "interval timer", photoperiod and temperature in the seasonal development of parthenogenetic and sexual morphs in the lime aphid, Eucallipterus tiliae L.

The ability of the lime aphid to produce sexuals (males and females) as an alternative to parthenogenetic females is regulated by a timing mechanism ('interval timer') which restrains the appearance of these morphs early in the year. In this species there are two 'interval timers': one controls the production of sexual females, and the other controls the production of males. Both 'intervals timers' are sensitive to day-length and temperature but they respond in different ways.With the approach of autumn the waning effect of the 'interval timer' inhibiting female production combined with the short day-lengths results in an increasing proportion of the aphids developing into sexual females. The restraining effect of the 'interval timer' results in a gradual transition from parthenogenetic to gamic reproduction over a period of several generations and is still operational in the autumn. However, in this species even relatively long day conditions (17 h) can induce the development of oviparae. This low threshold of response to day-length combined with the short generation time results in the sexual morphs appearing very early in the year. This is of considerable adaptive significance in years when, as frequently happens, the aphids disappear locally before the onset of autumn.

The role of intra-specific mechanisms and predation in regulating the numbers of the lime aphid, Eucallipterus tiliae L.

Analysis of changes in numbers of the lime aphid, Eucallipterus tiliae L., reveals that there is an overcompensated density dependent factor acting within years, and an inverse density dependent factor acting between years. Because of its effect on the aphid's rate of development and reproduction the variation in temperature from year to year acts as a density disturbing factor.There is no evidence from laboratory experiments to suggest that the quality of its food influences the numbers of the lime aphid. However, qualitative changes in the aphid following period of aphid abundance are shown to be important in the overcompensated density dependent response. The more prevalent insect predators influence the regulating process as they eat most of the aphids that remain after an aphid population has declined from a very high level of abundance. Their action results in a more overcompensated response than would occur in their absence.