Genetic differentiation as a result of adaptation to the phenologies of individual host trees in the galling aphid Kaltenbachiella japonica

Abstract.

• 1 The adaptation of the non-migratory galling aphid Kaltenbachiella japonica (Matsumura) to the budburst phenologies of individual host trees (Ulmus davidiana var. japonica Nakai) was investigated. There was a large variation in budburst time between individual host trees. We tested a hypothesis that aphid populations on respective host trees are genetically differentiated through adaptation to the budburst phenologies of host trees.
• 2 There was significant correlation between the budburst time of eight host trees in two successive years.
• 3 Tree-associated populations differed significantly in egg-hatching time, and the mean hatching time was significantly correlated with the mean budburst time of respective host trees.
• 4 Mating was made between male and female sexuales of the same clone (i.e. self-mating) to obtain selfed eggs. Hatching patterns of selfed eggs showed that there was a substantial amount of genetic variance in hatching time between clones (galls). Nested ANOVA demonstrated that the between-tree component contributed more to the phenotypic variance than the within-tree or within-gall components.
• 5 Reciprocal crosses between males and females of different clones furthermore demonstrated significant differences in hatching time between cross combinations, but no significant differences between the two reciprocals within combinations. This result suggests that there are no maternal effects for the timing of egg hatch and that the differences between selfed lines are attributable to genetic variance.
• 6 The fine-scale adaptation hypothesis was supported by egg-hatching experiments, which further suggest that budburst phenology varying between individual trees is acting as intense selective pressure on the egg-hatching time of associated insects.

     

Synchronization of larval emergence in winter moth (Operophtera brumata L.) and budburst in pedunculate oak (Quercus robur L.) under simulated climate change

1. The hypothesis that a 3 °C elevation in temperature and doubled CO₂ concentration would have no effect on the synchronization of winter moth egg hatch with budburst in oak was tested by comparing the separate and interactive effects of ambient and elevated (+ 3 °C) temperature and ambient and elevated (doubled to 340 p.p.m.) CO₂ in eight experimental Solardomes. In addition, an outdoor control was compared with the ambient temperature/CO₂ treatment combination.
2. Elevated temperature accelerated darkening (preceding egg hatch by about 5–10 days) and hatching of eggs developing off the trees; elevated CO₂ had no effect. The same effects were observed in eggs developing on the trees.
3. Within treatments, date of egg hatch was the same on trees with early or late budburst.
4. Egg darkening and budburst were closely synchronized at both ambient and elevated temperatures.
5. Both eggs and trees required fewer cumulative heat units (day degrees > 4 °C), for hatching and budburst, respectively, at ambient than elevated temperatures. The requirements in the outdoor control treatment were similar to those in the ambient Solardome treatment.
6. Egg hatch between 10 and 25 °C, on a temperature gradient in the laboratory, required a constant number of heat units; fewer were required below 10 °C.
7. Elevated temperatures, in the Solardomes and the field, delayed adult emergence from the pupae.
8. The results suggest that a general increase in temperature with climatic change would not affect the closeness of the synchronization between egg hatch of winter moth and budburst of oak.     

Are polyphagous geometrid moths with flightless females adapted to budburst phenology of local host species?

The majority of studies demonstrating local adaptation of insect herbivores involve sessile species, particularly those with a parthenogentic phase to their life history or endophagous "parasites" of plants. Current arguments suggest the strength of selection determines whether local adaptation can or cannot take place. Therefore local adaptation should not be limited to species with such traits. We studied the ability of three polyphagous geometrid moths with flightless adult females ( Erannis defoliaria , Operophtera brumata and O. fagata ) to synchronise their egg hatching with the budburst of a local host species in north east Scotland. A strong selection for hatching time is expected among generalist moths given the large variation in budburst phenology and an inability to hatch in synchrony with budburst decreases moth fitness substantially. In two successive seasons, we trapped emerging females from patches of five host species and recorded the temperature sum needed for 50% egg hatch of each brood laid by the trapped females. The hatching times of broods were compared against the average budburst time of the maternal host species in the study area. In addition, the trapping dates of each female were recorded. Only O. brumata showed synchrony with egg hatch and budburst which suggests local phenological adaptation to different host species . This could be maintained by selection and partial reproductive isolation between populations dwelling on different host species. No phenological adaptation was found in the other common geometrids of the study area.     

Opposing survivorship and fecundity effects of host phenology on the gall-forming aphid Hormaphis hamamelidis

Abstract  1. This study examined the synchronisation and roles of host and insect phenology for fundatrices of the gall-forming aphid Hormaphis hamamelidis , which forms galls on the leaves of the understorey tree Hamamelis virginiana .
2. Host and insect phenologies varied widely among years and individual host trees.
3. Most winter eggs (≈ 85%) hatched before budbreak and galls could be initiated but in 2 years there was no inherent risk for earlier hatching fundatrices in terms of greater mortality. Egg hatch was not synchronised tightly with host trees but instead seemed to occur well in advance of budbreak as a conservative, bet-hedging strategy to ensure the opportunity to form galls.
4. Fundatrices that formed galls later had higher fecundity, contrary to typical patterns for insects that feed on the first flush of leaves in temperate forests, where later herbivores have lower fecundity. Also, unlike many other systems, trees that had later budbreak had greater densities of galls.     

Influence of intra-tree variation in time of budburst of white spruce on herbivory and the behaviour and survivorship of Zeiraphera canadensis

Abstract.

• 1 Field studies were carried out to determine the effects of intra-tree variation in the time of budburst of white spruce, Picea glauca Moench (Voss.), on the behaviour and survivorship of, and herbivory by, the spruce bud moth, Zeiraphera canadensis Mutt. & Free.
• 2 There was significant variation in the time of budburst among whorls, shoots and buds. Budburst was acropetal, with buds in the interior of the lower crown bursting first and terminal buds on terminal shoots in the upper crown bursting last.
• 3 Bud moths laid the greatest proportion of their eggs in the middle of the crown and egg hatch was usually best synchronized to budburst in this region. Many eggs hatched before terminal buds on terminal shoots in the upper crown had burst and thus intra-tree variation in budburst decreased the probability that first-instar larvae would colonize the most important plant parts for growth.
• 4 However, many later instars dispersed upwards and outwards in the crown and colonized the late bursting buds in the upper crown. Such dispersal reduces the effectiveness of intra-plant variation in budburst to reduce herbivory and permits Z.canadensis to eat young nutritious buds for a longer period of time.
• 5 Intra-tree variance in the date of budburst was greater than that between trees but there were no consistent differences between the intra-tree variance of trees in half-sib families with high or low susceptibility to Z.canadensis. Differences between trees in herbivory, bud moth density and survivorship were not related to the amount of intra-tree variation in budburst.

     

Influence of inter-tree variation in time of budburst of white spruce on herbivory and the behaviour and survivorship of Zeiraphera canadensis

Abstract.

• 1 Field studies were carried out during 4 years to determine the effects of inter-tree variability in the time of budburst of white spruce, Picea glauca Moench (Voss.), on the distribution, behaviour and survivorship of, and herbivory by, the spruce bud moth, Zeiraphera canadensis Mutt. & Free.
• 2 Differences between trees in herbivory, bud moth density and survivorship were inversely related to the mean date of budburst. The most and least susceptible half-sib‘families’evaluated were the first and last to flush, respectively. Differences in herbivory for trees in two families with similar phenologies were partially due to oviposition preference.
• 3 Oviposition and larval survivorship were highest on trees in the most susceptible families. Although egg hatch was more closely synchronized with budburst in warm than in cool springs, between tree differences in the time of budburst were consistent over years.

     

Local adaptation and host race formation of a gall-forming aphid in relation to environmental heterogeneity

Summary The process of host race formation in the aphid Tetraneura yezoensis is examined in relation to its population structure. T. yezoensis induces pouch galls on new leaves of Ulmus davidiana and U. laciniata. Its populations on the two host species are often sympatric. Fundatrices found on one elm species, when reciprocally transplanted to the other, suffered greatly reduced average fitness. This shows that aphid populations associated with the two elm species are genetically differentiated in physiological traits. Individual trees of each elm species showed large differences in susceptibility to gall formation and in bud burst time, and such between-tree variations were consistent over years. Overwintered eggs taken in early spring from four trees (two from each species) were incubated under the same temperature conditions. The average hatching time differed significantly even between populations from conspecific trees, and the sequence of egg hatching paralleled that of the leafing of those four trees. This between-tree difference in hatching time was consistent over years and was found to be genetic, showing that gene flow between aphid populations on separate trees is often restricted. The heterogeneity in host traits may have promoted the evolution of philopatry in this aphid. Of the fundatrices that hatched on a tree of one elm species, a few precent were preadapted to gall formation on the other elm species. This suggests that the formation of a new host race proceeds parapatrically under disruptive selection and at a low level of gene flow. Evidence was actually obtained that a small fraction of Tetraneura alates are passively transported and land on non-host plants.     

Effects of elevated temperature on multi-species interactions: the case of Pedunculate Oak, Winter Moth and Tits

1. The effects of temperature on the Oak–Winter Moth–Tit food chain were studied at Wytham Wood, Oxford, and experimentally in the controlled environment solardomes at the Institute of Terrestrial Ecology, Bangor.
2. Tree cores from Wytham indicated that mature Oaks grew best at high temperatures and rainfall, but with low caterpillar populations. Young trees grew less well at elevated temperature, probably because they lost more water than they gained. Elevated temperatures advanced budburst, reduced foliar nitrogen and increased leaf toughness.
3. Moth eggs laid later or maintained at cooler temperatures than average required fewer heat units to hatch. Caterpillars took up to 50 days to complete growth at field temperatures but did so in only 20 days at a constant 15 °C.
4. The mass of Tit chicks at day 15 (day 1 = egg hatch) was positively correlated with temperature and negatively correlated with rainfall during the growing period.
5. At elevated temperature, budburst and moth egg hatch were synchronized, but earlier. Late feeding larvae and larvae fed on leaves from trees grown at elevated temperature produced smaller pupae. Pupal mass was unaffected when caterpillars and trees were maintained together under the same conditions.
6. Delaying egg hatch in Tits, to simulate conditions at elevated spring temperatures, resulted in reduced chick mass, body size and fledging success. This occurred because the chicks were fed later and prey quality was poorer, because the peak of caterpillar biomass was missed.
7. We predict that moth reproductive output will be retained at elevated temperatures because both leaves and caterpillars develop faster. Brood size in birds may be reduced because they cannot lay early enough to coincide with the narrower peak of food abundance.     

Adaptation of a generalist moth, Operophtera brumata, to variable budburst phenology of host plants

The adaptation of three allopatric populations of a generalist moth, Operophtera brumata (L.), on two tree species, Prunus padus (L.) and Quercus robur (L.) which represent the extremes of the timing of budburst in spring, was studied in Finland and Sweden. The synchrony of the hatching and budbursting was monitored, and its importance to dispersal and growth of larvae was assessed by rearing cohorts of larvae, whose hatching dates were manipulated, on both hosts. In addition, the realised heritability of the hatching time was estimated.Experimental populations hatched in approximate synchrony with the budburst of their original host species. As a result of the manipulation of the hatching dates of larvae, the growth rates of larvae decreased and the dispersal rates increased on both hosts in relation to the ageing of foliage. The realised heritability of hatching times was rather high (0.63). There were fewer differences in the host use efficiency and behaviour of the experimental populations than in the hatching phenology. The synchrony of hatching with the budburst of the local dominant host plant is probably a result of stabilising selection.     

Enemy-free space and the feeding niche ofan aphid

1. The nymphs of the aphid Monaphis antennata occupy an unusual feeding niche, being restricted to the upper surfaces of leaves and petioles. The possibility that this is a predator-avoidance strategy was investigated.
2. Nymphs could be induced to feed on the under surfaces of petioles and the mortality of these nymphs was then compared with the mortality of nymphs feeding from the upper surfaces when exposed to coccinellid predators.
3. Coccinellid larvae spent less time foraging on the upper surfaces of leaves and petioles than on the under surfaces of leaves and petioles.
4. The nymphs occupying the upper surfaces of petioles, the normal feeding position, were encountered by the coccinellids later and less often, and fewer were attacked compared with those occupying the under surfaces.
5. It is concluded that the feeding niche of the nymphs of this aphid is adaptive in that it reduces exposure to foraging predators and is the result of selection to occupy enemy-free space.     

Predicting adaptation of phenology in response to climate change, an insect herbivore example

Climate change has led to an advance in phenology in many species. Synchrony in phenology between different species within a food chain may be disrupted if an increase in temperature affects the phenology of the different species differently, as is the case in the winter moth egg hatch–oak bud burst system. Operophtera brumata (winter moth) egg hatch date has advanced more than Quercus robur (pedunculate oak) bud burst date over the past two decades. Disrupted synchrony will lead to selection, and a response in phenology to this selection may lead to species genetically adapting to their changing environment. However, a prerequisite for such genetic change is that there is sufficient genetic variation and severe enough fitness consequences. So far, examples of observed genetic change have been few. Using a half-sib design, we demonstrate here that O. brumata egg-hatching reaction norm is heritable, and that genetic variation exists. Fitness consequences of even a few days difference between egg hatch and tree bud opening are severe, as we experimentally determined. Estimates of genetic variation and of fitness were then combined with a climate scenario to predict the rate and the amount of change in the eggs' response to temperature. We predict a rapid response to selection, leading to a restoration of synchrony of egg hatch with Q. robur bud opening. This study shows that in this case there is a clear potential to adapt – rapidly – to environmental change. The current observed asynchrony is therefore not due to a lack of genetic variation and at present it is unclear what is constraining O. brumata to adapt. This kind of model may be particularly useful in gaining insight in the predicted amount and rate of change due to environmental changes, given a certain genetic variation and selection pressure.     

Intraspecific variation in host plant quality: the aphid Cinara cupressi on the Mexican cypress, Cupressus lusitanica

Abstract.

• 1 In Kenya, the aphid Cinara cupressi (Buckton) has an aggregated distribution across individuals of its host plant, the Mexican cypress, Cupressus lusitanica (Mill).
• 2 There are significant differences in host plant quality: C.cupressi nymphs transplanted onto cypress trees with low aphid infestations are significantly less likely to establish than aphids transplanted onto trees with high aphid infestations.
• 3 Given that some transplanted nymphs establish, the previous level of infestation does not influence aphid survival.
• 4 Evidence for local adaptation to individual plants was not found: nymphs transplanted to the trees from which their parents were collected neither established more readily nor showed lower mortality than those transplanted to other trees with similar levels of infestation. Therefore the variation in host plant quality has not led to the development of local adaptation to particular host plants.

     

The role of cereal growth stages and crowding in the induction of alatae in Sitobion avenae and its consequences for population growth

Abstract. 1. Alatiform nymphs of Sitobion avenae were first recorded on wheat in 1977 and 1978 when there were 0.6 aphids per tiller and reached a maximum proportion of 0.9 of the population despite a seventeen-fold difference in aphid density between years.
2. At the same aphid density per tiller there was a higher proportion of alatiform nymphs on the tillers at an advanced than at an earlier stage of plant growth.
3. Both pre- and post-natal crowding, and late stages of plant growth, were important in inducing the development of alatae in the laboratory.
4. Changes in the quality of the host plant through its previously reported effect on reproductive and survival rates, and its effect on alate induction and emigration was important in shaping the population dynamics of S.avenae on cereals.     

Preference and performance of the hyperparasitoid Syrphophagus aphidivorus (Hymenoptera: Encyrtidae): fitness consequences of selecting hosts in live aphids or aphid mummies

Abstract.  1. Theoretical models predict that ovipositional decisions of parasitoid females should lead to the selection of the most profitable host for parasitoid development. Most parasitoid species have evolved specific adaptations to exploit a single host stage. However, females of the aphid hyperparasitoid Syrphophagous aphidivorus (Mayr) (Hymenoptera: Encyrtidae) display a unique and atypical oviposition behaviour by attacking either primary parasitoid larvae in live aphids, or parasitoid pupae in dead, mummified aphids.
2. In the laboratory, the correlation between host suitability and host preference of S. aphidivorus on the host Aphidius nigripes Ashmead parasitising the aphid Macrosiphum euphorbiae (Thomas) was investigated.
3. The relative suitability of the two host stages was determined by measuring hyperparasitoid fitness parameters (survival, development time, fecundity, sex ratio, and adult size of progeny), and calculating the intrinsic rate of population increase ( r _m). Host preference by S. aphidivorus females and the influence of aphid defence behaviour on host selection was also examined.
4. Hyperparasitoid offspring performance was highest when developing from hosts in aphid mummies and females consistently preferred this host to hosts in parasitised aphids. Although aphid defensive behaviour may influence host selection, it was not a determining factor. Ecological and evolutionary processes that might have led to dual oviposition behaviour in S. aphidivorus are discussed.     

Intraspecific variability in herbivore performance and host quality: a field study of Uroleucon caligatum (Homoptera: Aphididae) and its Solidago hosts (Asteraceae)

Abstract. 1. Performances of two clones of Uroleucon caligatum on eleven clones of Solidago were measured by caging aphids on plants in the field.
2. Several measures were obtained, including developmental time from birth to adulthood, size of first instar nymphs, adult weight, and total colony weight.
3. All measures of performance were strongly affected by plant clone.
4. Effect of aphid clone-plant clone interaction was significant only for first instar size.
5. In a subsequent screenhouse experiment, plant clones were subjected to uniform conditions and still exhibited large differences in host quality.     

Phenological variation as protection against defoliating insects: the case of<Emphasis Type="Italic"> Quercus robur</Emphasis> and<Emphasis Type="Italic"> Operophtera brumata</Emphasis>

Phenological synchrony between budburst and emergence of larvae is critical for the fitness of many spring-feeding insect herbivores. Therefore, large intraspecific variation in timing of budburst of the host may have a negative effect on the herbivore. We studied how asynchrony between emergence of larvae and budburst affects the fitness of Operophtera brumata (Lepidoptera: Geometridae), a major defoliator of Quercus robur, which can adapt to the phenology of a single tree. It is known that, in maturing leaves of Q. robur, accumulation of condensed tannins has a negative effect on growth of O. brumata. However, there is no information about the effect of hydrolysable tannins and other phenolics that are potential antifeedants. In this study, we also analysed changes in secondary chemistry of the foliage of Q. robur and how different compounds are correlated with growth and survival of O. brumata. The effect of asynchrony on O. brumata was studied in rearing experiments. The neonate larvae were incubated without food for different periods of time. The decline in nutritional quality of foliage was estimated by rearing cohorts of larvae with manipulated hatching times on the leaves of ten individual Q. robur trees. For the chemical analysis, the foliage of these trees was sampled at regular intervals. In the absence of foliage, mortality of neonate larvae started to increase exponentially soon after the larvae emerged. If the larvae missed budburst, the decline in nutritional quality of the foliage led to increased mortality and lower body mass (= fecundity). Hydrolysable tannins were not significantly correlated with performance of the larvae. Only condensed tannins were found to correlate negatively with the growth and survival of O. brumata. Certain individual trees were unsuitable hosts for O. brumata because the decline in quality of the foliage was very rapid. Based on regression equations for increasing rate of mortality and decreasing fecundity, we calculated that a relatively small mismatch of +/-30 degree days between budburst and hatching of larvae leads to a 50% decrease in the fitness of O. brumata. Thus, large phenological variation within a Q. robur stand can limit the colonisation of neighbouring trees by dispersing larvae. Furthermore, the hybridisation of moths adapted to phenologically different trees may lead to maladapted phenology of their offspring.     

Influence of balsam fir (Abies balsamea) budburst phenology on hemlock looper (Lambdina fiscellaria)

Many herbivorous insects emerge synchronously with budburst of their host plant, as the nutritional quality of foliage often decreases rapidly following budburst. We carried out manipulative field experiments to evaluate the influence of bud and shoot phenology on performance of the hemlock looper, Lambdina fiscellaria Guenée (Lepidoptera: Geometridae: Ourapterygini), on balsam fir, Abies balsamea (L.) Mill. (Pinaceae), in NF, Canada. Hemlock looper survival, pupal weight, and realized fecundity, which were then combined to estimate fitness, were all highest when newly emerged first instars were placed on foliage of current‐year shoots that had completed approximately 25–35% of their elongation, and lower when placed on younger or older foliage. Survival of a small portion of larvae placed on buds a week before budburst suggests that newly emerged first instars either entered unburst buds or survived for a week without food. In laboratory experiments, approximately half of larvae survived for 4 days without food or water at 10 °C and 65% r.h. The timing of egg hatch in the field appeared to be adaptive, but the short duration of egg hatch suggests that another factor in addition to host plant phenology exerts stabilizing selection pressure on the timing of egg hatch.     

Life table parameters affecting the population development of the woolly beech aphid, Phyllaphis fagi

A detailed study of the basic biological parameters affecting the population development of the woolly beech aphid, Phyllaphis fagi L. (Homoptera: Drepanosiphidae), was performed under field and laboratory conditions in order to obtain information about factors responsible for outbreak situations in forest nurseries. In the nurseries, the aphid eggs have found to hatch before budburst. The newly hatched nymphs were highly active in searching for feeding places, which resulted in higher nymph mortality in the first generation than in the following generations. Ten aphid generations were recorded in the nursery during one growing season. In the laboratory, development time, nymph mortality, reproduction, and adult lifetimes were assessed for P. fagi at four constant temperatures, i.e., 12, 15, 20, and 25 °C through four successive generations. Both temperature and aphid generation were found to affect life table parameters. The first generation, however, was less susceptible to changing temperatures than the following generations. Phyllaphis fagi reproductive effort was more dependent on temperature than on aphid generation. No significant difference was found in reproductive effort between apterous and alate females. The potential for outbreak situations therefore exists throughout the summer. The formation of winged morphs was not restricted to a few generations but rather continued for several months, with activity peaking in all years in mid-June. This had implications for migration to new seedlings in the nurseries. The highest r_m value was obtained at 20 °C. Nymphs born at higher temperatures were sometimes born deformed and unable to survive.     

Effects of aluminum,calcium and low pH on egg hatching and nymphal survival of Cloeon triangulifer McDunnough (Ephemeroptera: Baetidae)

Laboratory experiments were conducted to assess the effects of aluminum, calcium and low pH on egg hatching and nymphal survival of the mayfly Cloeon triangulifer. Percent successful hatch (living nymphs breaking free of the chorion) decreased and percent partial hatch (nymphs dying attached to the chorion) increased with increasing acidity (pH 7.5–3.0). Most hatches occurring below pH 5.0 were partial hatches. Decreased time of exposure to acidic waters increased percent successful hatch and decreased percent partial hatch. Time to first hatch was not affected by pH. Eggs were incubated in acidic waters (pH 4.0 and 5.5) with additions of calcium (10 and 100 mg l^–1) and aluminum (100 and 500 g l^–1). Aluminum decreased percent successful hatch and increased percent partial hatch and calcium increased both percent successful hatch and percent partial hatch. Time to first hatch was increased by both aluminum and calcium. The 96 h LC50 for small nymphs was pH 4.75. Addition of aluminum (100 and 500 µg l^–1) to acidic waters (pH 4.0 and 5.0) reduced nymphal mortality by 8–22%. Toxic effects of low pH on egg hatching and early nymphs may contribute to the absence of mayflies from acidified habitats.Contribution No. 1469 of the Maine Agricultural Experiment Station, University of Maine, Orono, Maine 04469 USA.Contribution No. 1469 of the Maine Agricultural Experiment Station, University of Maine, Orono, Maine 04469 USA.     

ECOLOGICAL HISTORY AND EVOLUTION IN A NOVEL ENVIRONMENT: HABITAT HETEROGENEITY AND INSECT ADAPTATION TO A NEW HOST PLANT

Environmental heterogeneity has often been implicated in the maintenance of genetic variation. However, previous research has not considered how environmental heterogeneity might affect the rate of adaptation to a novel environment. In this study, I used an insect-plant system to test the hypothesis that heterogeneous environments maintain more genetic variation in fitness components in a novel environment than do uniform environments. To manipulate recent ecological history, replicate populations of the dipteran leafminer Liriomyza trifolii were maintained for 20 generations in one of three treatments: a heterogeneous environment that contained five species of host plant, and two uniform environments that contained either a susceptible chrysanthemum or tomato. The hypothesis that greater genetic variance for survivorship and developmental time on a new host plant (a leafminer-resistant chrysanthemum) would be maintained in the heterogeneous treatment relative to the uniform environments was then tested with a sib-analysis and a natural selection experiment. Populations from the heterogeneous host plant treatment had no greater genetic variance in either larval survivorship or developmental time on the new host than did populations from either of the other treatments. Moreover, the rate of adaptation to the new host did not differ between the ecological history treatments, although the populations from the uniform chrysanthemum treatment had higher mean survivorship throughout the selection experiment. The estimates of the heritability of larval survivorship from the sib-analysis and selection experiment were quite similar. These results imply that ecologically realistic levels of environmental heterogeneity will not necessarily maintain more genetic variance than uniform environments when traits expressed in a particular novel environment are considered.