Swift laboratory thermal evolution of wing shape (but not size) in Drosophila subobscura and its relationship with chromosomal inversion polymorphism

Latitudinal clinal variation in wing size and shape has evolved in North American populations of Drosophila subobscura within about 20 years since colonization. While the size cline is consistent to that found in original European populations (and globally in other Drosophila species), different parts of the wing have evolved on the two continents. This clearly suggests that 'chance and necessity' are simultaneously playing their roles in the process of adaptation. We report here rapid and consistent thermal evolution of wing shape (but not size) that apparently is at odds with that suggestion. Three replicated populations of D. subobscura derived from an outbred stock at Puerto Montt (Chile) were kept at each of three temperatures (13, 18 and 22 degrees C) for 1 year and have diverged for 27 generations at most. We used the methods of geometric morphometrics to study wing shape variation in both females and males from the thermal stocks, and rates of genetic divergence for wing shape were found to be as fast or even faster than those previously estimated for wing size on a continental scale. These shape changes did not follow a neat linear trend with temperature, and are associated with localized shifts of particular landmarks with some differences between sexes. Wing shape variables were found to differ in response to male genetic constitution for polymorphic chromosomal inversions, which strongly suggests that changes in gene arrangement frequencies as a response to temperature underlie the correlated changes in wing shape because of gene-inversion linkage disequilibria. In fact, we also suggest that the shape cline in North America likely predated the size cline and is consistent with the quite different evolutionary rates between inversion and size clines. These findings cast strong doubts on the supposed 'unpredictability' of the geographical cline for wing traits in D. subobscura North American colonizing populations.     

Allometric and nonallometric components of Drosophila wing shape respond differently to developmental temperature

Phenotypic plasticity of wing size and shape of Drosophila simulans was analyzed across the entire range of viable developmental temperatures with Procrustes geometric morphometric method. In agreement with previous studies, size clearly decreases when temperature increases. Wing shape variation was decomposed into its allometric (24%) and nonallometric (76%) components, and both were shown to involve landmarks located throughout the entire wing blade. The allometric component basically revealed a progressive, monotonous variation along the temperature. Surprisingly, nonallometric shape changes were highly similar at both extremes of the thermal range, suggesting that stress, rather than temperature per se, is the key developmental factor affecting wing shape.     

Developmental time,body size and wing loading in Drosophila buzzatii from lowland and highland populations in Argentina

Genotype-by-temperature interaction is a necessary condition for adaptive evolution of fitness traits as a response to temperature. Several fitness-related traits (developmental time, pre-adult survival, thorax and wing lengths, and wing loading) were measured in laboratory-reared D. buzzatii from four populations sampled at different altitudes in north-western Argentina: a lowland population (407 m a.s.l.), two populations from intermediate altitude (780 to 950 m a.s.l.), and a highland population (2380 m a.s.l.). Temperature is the main climatic difference between the collection sites: lowland but not highland populations are exposed to physiologically high temperatures during both spring and summer in nature. Three growth temperatures (20, 25 and 30 degrees C) were used to test for population-by-temperature interactions. Both developmental time and pre-adult survival exhibit highly significant population-by-temperature interaction. Pre-adult survival at 30 degrees C is significantly higher in lowland than in highland populations, but not so at lower growth temperatures (20 and 25 degrees C). Both wing length and wing loading show no population-by-temperature interaction, indicating that these traits are not the direct targets of thermal adaptation in nature. Wing loading is higher in highland than in lowland populations, suggesting that flight performance is subject to stronger selection in the highland population. This hypothesis is consistent with ecological observations in both types of populations. There is no obvious among-population relationship between developmental time and body size, even though both traits are related within populations in a well-known trade-off. Overall, thermal adaptation is evident for developmental time and pre-adult survival but not for size-related traits.     

Temperature-related genetic changes in laboratory populations of Drosophila subobscura: evidence against simple climatic-based explanations for latitudinal clines

Parallel latitudinal clines to the long-standing ones in the original Palearctic populations have independently evolved at different rates for chromosomal polymorphism and body size in South and North American populations of Drosophila subobscura since colonization around 25 years ago. This strongly suggests that (micro) evolutionary changes are largely predictable, but the underlying mechanisms are unknown. The putative role of temperature per se was investigated by using three sets of populations at each of three temperatures (13 degrees , 18 degrees , and 22 degrees C) spanning much of the tolerable range for this species. We found a lower chromosomal diversity at the warmest temperature; a quick and consistent shift in gene arrangement frequencies in response to temperature; an evolutionary decrease in wing size, mediated by both cell area and cell number, at 18 degrees C; no relationship between wing size and those inversions involved in latitudinal clines; and a shortening of the basal length of longitudinal vein IV relative to its total length with increasing standard dose. The trends for chromosomal polymorphism and body size were generally inconsistent from simple climatic-based explanations of worldwide latitudinal patterns. The findings are discussed in the light of available information on D. subobscura and results from earlier thermal selection experiments with various Drosophila species.     

Growth temperature and phenotypic plasticity in two Drosophila sibling species: probable adaptive changes in flight capacities

In the sibling species Drosophila melanogaster and D. simulans, growth and development at constant temperatures, from 12 to 30 °C, resulted in extensive variations of adult size and flight parameters with significant differences between species. Changes in body weight, thorax length and wing length were nonlinear, with maximum values of each trait at lower temperatures for D. simulans than for its sibling species. By contrast, the wing/thorax ratio and the wing loading varied monotonically with growth temperature. These traits were negatively correlated, the wing/thorax ratio decreasing with growth temperature while the wing loading increased. Wing/thorax ratio, which is easier to measure, thus appears as a convenient predictor of wing loading. During tethered flight at the same ambient temperature, the wingbeat frequency changed linearly as a function of the wing moment of inertia. More interestingly, the beat rate was strongly correlated with the increase of wing loading at growth temperature above 13 °C. The likely adaptive significance of these morphometrical changes for flight efficiency is discussed.     

Developmental time and size-related traits in Drosophila buzzatii along an altitudinal gradient from Argentina

Clinal analysis for fitness-related traits provides a well-known approach to investigate adaptive evolution. Several fitness-related traits (developmental time, thorax length, wing length and wing loading) were measured at two laboratory generations (G7 and G33) of D. buzzatii from an altitudinal gradient from northwestern Argentina, where significant thermal differences persist. Developmental time (DT) was positively correlated with altitude of origin of population. Further, DT was negatively correlated with maximal mean temperature at the site of origin of population, and this thermal variable decreases with altitude. Wing loading tended to be larger in highland than in lowland populations, suggesting that flight performance is subject to stronger selection pressure in highland populations. Developmental time showed a significant increase with laboratory generation number. There was no significant correlation between developmental time and body size across populations along the altitudinal cline of DT. This result illustrates that developmental time and body size do not always evolve in the same direction, even though both traits are often positively and genetically correlated in a well-known tradeoff in Drosophila.     

Thermal evolution of pre-adult life history traits, geometric size and shape, and developmental stability in Drosophila subobscura

Replicated lines of Drosophila subobscura originating from a large outbred stock collected at the estimated Chilean epicentre (Puerto Montt) of the original New World invasion were allowed to evolve under controlled conditions of larval crowding for 3.5 years at three temperature levels (13, 18 and 22 degrees C). Several pre-adult life history traits (development time, survival and competitive ability), adult life history related traits (wing size, wing shape and wing-aspect ratio), and wing size and shape asymmetries were measured at the three temperatures. Cold-adapted (13 degrees C) populations evolved longer development times and showed lower survival at the highest developmental temperature. No divergence for wing size was detected following adaptation to temperature extremes (13 and 22 degrees C), in agreement with earlier observations, but wing shape changes were obvious as a result of both thermal adaptation and development at different temperatures. However, the evolutionary trends observed for the wing-aspect ratio were inconsistent with an adaptive hypothesis. There was some indication that wing shape asymmetry has evolutionarily increased in warm-adapted populations, which suggests that there is additive genetic variation for fluctuating asymmetry and that it can evolve under rapid environmental changes caused by thermal stress. Overall, our results cast strong doubts on the hypothesis that body size itself is the target of selection, and suggest that pre-adult life history traits are more closely related to thermal adaptation.     

Causes of variation in wing loading among Drosophila species

We examined influences on wing and body size in 11 species (12 strains) of Drosophila. Six measures of wing length and width were closely correlated with wing area and suggested little variation in wing shape among the species. Among ten species wing loading, an important factor in flight costs and manoeuvrability, increased as body mass increased at a rate consistent with expectations from allometric scaling of wing area and body mass to body length. Intraspecific variation in wing loading showed similar relationships to body mass. Density and temperature during larval development influenced wing loading through general allometric relations of body size and wing area. Temperature during the pupal stage, but not during wing hardening after eclosion, influenced wing area independently of body size. Wing area increased as growth temperature decreased. Individuals reared at cooler temperatures thus compensated for a potential allometric increase in wing loading by differentially enlarging the wing area during pupal development.     

Morphological Change to Birds over 120 Years Is Not Explained by Thermal Adaptation to Climate Change

Changes in morphology have been postulated as one of the responses of animals to global warming, with increasing ambient temperatures leading to decreasing body size. However, the results of previous studies are inconsistent. Problems related to the analyses of trends in body size may be related to the short-term nature of data sets, to the selection of surrogates for body size, to the appropriate models for data analyses, and to the interpretation as morphology may change in response to ecological drivers other than climate and irrespective of size. Using generalized additive models, we analysed trends in three morphological traits of 4529 specimens of eleven bird species collected between 1889 and 2010 in southern Germany and adjacent areas. Changes and trends in morphology over time were not consistent when all species and traits were considered. Six of the eleven species displayed a significant association of tarsus length with time but the direction of the association varied. Wing length decreased in the majority of species but there were few significant trends in wing pointedness. Few of the traits were significantly associated with mean ambient temperatures. We argue that although there are significant changes in morphology over time there is no consistent trend for decreasing body size and therefore no support for the hypothesis of decreasing body size because of climate change. Non-consistent trends of change in surrogates for size within species indicate that fluctuations are influenced by factors other than temperature, and that not all surrogates may represent size appropriately. Future analyses should carefully select measures of body size and consider alternative hypotheses for change.     

Reaction norms across and genetic parameters at different temperatures for thorax and wing size traits in Drosophila aldrichi and D. buzzatii

Reaction norms across seven constant and one fluctuating temperature of development were measured for thorax length and several wing size traits for up to 10 isofemale lines of each of the cactophilic Drosophila species, D. aldrichi and D. buzzatii, originating from the same locality. Maximum thorax length was reached at different low to intermediate temperatures for the two species, whereas wing length was highest at the lowest temperature in both species. Various ratio parameters showed pronounced species differences. The reaction norm for the wing loading index (wing length/thorax length) decreased monotonically with temperature in both species, but was much steeper and spanned a wider range in D. aldrichi than in D. buzzatii, suggesting either that wing loading is not a good characterization of flight capacity or, more likely, that flight optimization does not occur in the same manner in both species. The vein ratio (distal length/proximal length of the third vein) increased with temperature in D. buzzatii but decreased in D. aldrichi. Wing development in the two species thus is very different, with the proximal part of the wing in D. buzzatii more closely allied to the thorax than to the distal part. Among line variation was significant for all traits in both species, and most pronounced for thorax length and the ratio parameters. Coefficients of variation were significantly different between the species for all traits, with those in D. aldrichi higher than in D. buzzatii. Genetic variance in plasticity was significant for all traits in D. buzzatii, but only for seven out of 12 in D. aldrichi. Additive genetic variances for all traits in both species were significantly larger than zero. Genetic correlations between thorax length and several wing length parameters, and between these and wing area, were positive and generally significant in both species. The genetic correlation between the distal and the proximal length of the third vein was not significantly different from zero in D. aldrichi, but negative and significant in D. buzzatii. Heritabilites varied significantly among temperatures for almost all traits in both species. Phenotypic variances were generally higher in D. aldrichi than in D. buzzatii, and commonly highest at the extreme temperatures in the former species. At the high temperature the genetic variances also were usually highest in D. aldrichi. The data clearly suggest that the process of thermal adaptation is species specific and caution against generalizations based on the study of single species.     

Latitudinal variation in wild populations of Drosophila melanogaster: heritabilities and reaction norms

Large amounts of genetic variation for wing length and wing area were demonstrated both within and between Drosophila melanogaster populations along a latitudinal gradient in South America. Wing length and wing area showed a strong positive correlation with latitude in both wild flies and laboratory-raised descendants. Large population differences were observed for heritability and coefficient of variation of these two traits, whereas relatively small population differences were found for development time, viability, pupal mortality, sex ratio and their norms of reaction to four developmental temperatures. No clear-cut latitudinal clines were established for these life-history characters. These results are discussed in the light of Bergmann's Rule and the relation between larval development and adult body size.     

Adaptation to different climates results in divergent phenotypic plasticity of wing size and shape in an invasive drosophilid

The phenotypic plasticity of wing size and wing shape of Zaprionus indianus was investigated in relation to growth temperature (17°C to 31°C) in two natural populations living under different climates, equatorial and subtropical. The two populations were clearly distinguished not only by their wing size (the populations from the colder climate being bigger in size), but also by the shape of the response curves to growth temperature i.e., their reaction norms. In this respect, the temperature at which the size of the wing was maximum was about 3°C higher in the equatorial population. Such a difference in size plasticity is already found in two other nonclosely related species, might be a general evolutionary pattern in drosophilids. Wing shape was investigated by calculating an ellipse included into the wing blade, then by considering the ratio of the two axes, and also by analysing the angular position of 10 wing-vein landmarks. For an overall shape index (ratio of the two axes of the ellipse), a regular and almost linear increase was observed with increasing temperature i.e., a more round shape at high temperatures. Wing shape was also analysed by considering the variations of the various angles according to temperature. A diversity of response curves was observed, revealing either a monotonous increase or decrease with increasing temperature, and sometimes a bell shape curve. An interesting conclusion is that, in most cases, a significant difference was observed between the two populations, and the difference was more pronounced at low temperatures. These angular variations are difficult to interpret in an evolutionary context. More comparative studies should be undertaken before reaching some general conclusions.     

Quantitative-genetic analysis of wing form and bilateral asymmetry in isochromosomal lines ofDrosophila subobscura using Procrustes methods

Fluctuating asymmetry (FA) is often used as a measure of underlying developmental instability (DI), motivated by the idea that morphological variance is maladaptive. Whether or not DI has evolutionary potential is a highly disputed topic, marred by methodological problems and fuzzy prejudices. We report here some results from an ongoing study of the effects of karyotype, homozygosity and temperature on wing form and bilateral asymmetry using isochromosomal lines ofDrosophila subobscura. Our approach uses the recently developed methodologies in geometric morphometrics to analyse shape configurations of landmarks within the standard statistical framework employed in studies of bilateral asymmetries, and we have extended these methods to partition the individual variation and the variation in asymmetries into genetic and environmental causal components. The analyses revealed temperaturedependent expression of genetic variation for wing size and wing shape, directional asymmetry (DA) of wing size, increased asymmetries at suboptimal temperature, and a transition from FA to DA in males as a result of increase in the rearing temperature. No genetic variation was generally detected for FA in our samples, but these are preliminary results because no crosses between lines were carried out and, therefore, the contribution of dominance was not taken into account. In addition, only a subset of the standing genetic variation was represented in the experiments.     

Population densities in Drosophila obscura Fallén and D. subobscura Collin

Abstract. 1. The population densities of Drosophila obscura and D. subobscura in mixed deciduous woodland were investigated by multiple recapture techniques, marking flies with micronized fluorescent dusts.
2. Approximate densities per 100 m² were as follows: September 1974: D.subobscura males 5, D.subobscura females 31. April 1975: D.obscura males 1, D.obscura females 2; D.subobscura males 0.3, D.subobscura females 1. June 1975: D.subobscura males 6, D.subobscura females 9.
3. Actual estimates have been compared using relative estimates derived from a reanalysis of Shorrocks's (1975) data.
4. Throughout most of the year D.subobscura was numerically dominant, rising to a peak of 750/100 m² in the winter; but in late spring D.obscura was dominant, having reduced winter mortality by diapausing.
5. D.obscura 's density was comparable with those of its Nearctic relatives; D.subobscura 's was considerably higher.
6. The effects of previous workers' approximations on the estimation of population parameters have been examined, and possible theoretical consequences have been indicated.     

Increasing winter temperatures explain body size decrease in wintering bird populations of Northern Europe—But response patterns vary along the spatioclimatic gradient

Aim

Recent evidence has shown changes in body size and shape of individuals, which are suggested to be a result of global warming caused by climate change. Here, we explored the spatiotemporal changes in wing length and body mass of 24 wintering bird species in Northern Europe and how these relate to temperature anomaly.

Location

Finland and Sweden, Europe.

Time Period

50 years, 1970 to 2020.

Major Taxa Studied

Birds, 24 species.

Methods

We used site-specific, long-term winter ringing data containing wing length and body mass measurements from across Sweden and Finland for 24 bird species. We modelled wing length and body mass change over time, in relation to the spatioclimatic gradient and as response to temperature anomalies (of [i] the same winter as the ringing took place, [ii] the previous winter and [iii] the previous spring) by accounting for phylogenetic relatedness between species and their species-specific responses to each predictor of interest.

Results

We show that across all species, body size has decreased since the 1970s, with a negative relationship between wing length and temperature anomalies of previous winters, suggesting carry-over effects likely linked to body size-related survival or dispersal. Body mass was negatively related to the temperature anomaly of the same winter, indicating more immediate effects related to reduced fat reserves during mild winters.

Main Conclusions

Our results highlight a climate-driven decrease in body size across several species and its association with positive anomalies in winter temperature in the high latitudes. However, the responses are not spatially uniform and there is considerable species-specific variation, emphasizing the importance of conducting multispecies studies when investigating responses to climate change. The mechanisms of decreasing wing length and body mass seem to differ and underline the immediate and carry-over effects of temperature warming during the nonbreeding season.     

LATITUDINAL VARIATION OF WING:THORAX SIZE RATIO AND WING-ASPECT RATIO IN DROSOPHILA MELANOGASTER

In dipterans, the wing-beat frequency, and, hence, the lift generated, increases linearly with ambient temperature. If flight performance is an important target of natural selection, higher wing:thorax size ratio and wing-aspect ratio should be favored at low temperatures because they increase the lift for a given body weight. We investigated this hypothesis by examining wing: thorax size ratio and wing-aspect ratio in Drosophila melanogaster collected from wild populations along a latitudinal gradient and in their descendants reared under standard laboratory conditions. In a subset of lines, we also studied the phenotypic plasticity of these traits in response to temperature. To examine whether the latitudinal trends in wing:thorax size ratio and wing-aspect ratio could have resulted from a correlated response to latitudinal selection on wing area, we investigated the correlated responses of these characters in lines artificially selected for wing area. In both the geographic and the artificially selected lines, wing:thorax size ratio and wing-aspect ratio decreased in response to increasing temperature during development. Phenotypic plasticity for either trait did not vary among latitudinal lines or selective regimes. Wing:thorax size ratio and wing-aspect ratio increased significantly with latitude in field-collected flies. The cline in wing:thorax size ratio had a genetic component, but the cline in wing-aspect ratio did not. Artificial selection for increased wing area led to a statistically insignificant correlated increase in wing:thorax size ratio and a decrease in wing-aspect ratio. Our observations are consistent with the hypotheses that high wing-thorax size ratio and wing aspect ratio are per se selectively advantageous at low temperatures.     

Direct and correlated responses to artificial selection on developmental time and wing length in Drosophila buzzatii

Abstract.— Developmental time and body size are two positively correlated traits closely related to fitness in many organisms including Drosophila . Previous work suggested that these two traits are involved in a trade-off that may result from a negative genetic correlation between their effects on pre-adult and adult fitness. Here, we examine the evolution of developmental time and body size (indexed by wing length) under artificial selection applied to one or both traits in replicated D. buzzatii populations. Directional changes in both developmental time and wing length indicate the presence of substantial additive genetic variance for both traits. The strongest response to selection for fast development was found in lines selected simultaneously to reduce both developmental time and wing length, probably as an expected consequence of a synergistic effect of indirect selection. When selection was applied in the direction opposite to the putative genetic correlation, that is, large wing length but fast development, no responses were observed for developmental time. Lines selected to reduce both wing length and developmental time diverged slightly faster from the control than lines selected to increase wing length and reduce developmental time. However, wing length did not diverge from the control in lines selected only for fast development. These results suggest a complex genetic basis of the correlation between developmental time and wing length, but are generally consistent with the hypothesis that both traits are related in a trade-off. However, we found that this trade-off may disappear under uncrowded conditions, with fast-developing lines exhibiting a higher pre-adult viability than other lines when tested at high larval density.     

Wing morphology,winter ecology,and fecundity selection: evidence for sex-dependence in barn swallows (<Emphasis Type="Italic">Hirundo rustica</Emphasis>)

Variation in wing morphology results from the combination of diverse selection pressures. Wing feather morphology within species varies with sex and ontogenetic effects, and also with ecological factors. Yet, the direction of causation for the wing morphology–ecology association remains to be elucidated. Under the ‘ecology-dependence’ hypothesis, wing morphology covaries with ecological conditions, because the latter affect feather molt. Alternatively, the ‘habitat choice’ hypothesis posits that individuals with different wing morphology choose different habitats because of the habitat-dependent advantages of a specific wing morphology. We tested these competing hypotheses in the migratory, aerially insectivorous barn swallow (Hirundo rustica). We quantified wing morphology (isometric size, pointedness, and convexity) on the same individuals during consecutive breeding seasons (i.e., before and after molt in sub-Saharan wintering areas) and located wintering areas using light-level geolocators. Wing pointedness of females but not males during 1 year negatively correlated with vegetation vigor (gauged by the Normalized Difference Vegetation Index; NDVI) in the African area where individuals spent the next winter. Partial least-squares path modelling showed that the association between wing morphology and NDVI was sex-dependent. Conversely, NDVI during wintering did not predict wing morphology in the next breeding season. Because wing morphology can have carry-over effects on subsequent performance, we investigated selection on wing traits and found strong positive fecundity selection on wing size of females. Our results suggest that female barn swallows choose their wintering habitat depending on their wing morphology. In addition, directional fecundity selection operates on females, suggesting sex-dependence of current selection on the flight apparatus.     

Variation of life-history and morphometrical traits in Drosophila buzzatii and Drosophila simulans collected along an altitudinal gradient from a Canary island

Variation in three life‐history traits (developmental time, preadult viability and daily female productivity) and five morphometrical traits (thorax length, wing length, wing width, wing/thorax ratio and wing‐aspect ratio) was studied at three developmental temperatures (20, 25 and 30 °C) in Drosophila buzzatii and Drosophila simulans collected on the island of La Gomera (Canary Archipelago). The flies originated from five closely situated localities, representing different altitudes (from 20 to 886 m above sea level) and a range of climatic conditions. We found statistically significant population effects for all traits in D. buzzatii and for most of the traits in D. simulans. Although no correlations of trait values with altitude were detected, geographical patterns for three life‐history traits and body size in D. buzzatii indicated that short‐range geographical variation in this species could be maintained by local climatic selection. Five of eight traits showed population‐by‐temperature interactions either in D. buzzatii or in D. simulans, but in all cases except wing width in D. buzzatii this could not be interpreted as adaptive responses to thermal conditions in the localities. The range of plastic changes across temperatures for particular traits differed between species, indicating a possibility for different levels of environmental stress experienced by the natural populations. The reaction norm curves and the response of within‐population variability to thermal treatments suggested better adaptations to higher and lower temperatures for D. buzzatii and D. simulans, respectively. The levels of among‐population differentiation depended on developmental temperature, implying environmental effects on the expression of the genetic variance. At 20 and 25 °C, interpopulation variability in D. buzzatii was higher than in D. simulans, while at 30 °C the opposite trend was observed. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 84 , 119–136.     

Symmetry breaking in interspecific Drosophila hybrids is not due to developmental noise

Hybrids from crosses of different species have been reported to display decreased developmental stability when compared to their pure species, which is conventionally attributed to a breakdown of coadapted gene complexes. Drosophila subobscura and its close relative D. madeirensis were hybridized in the laboratory to test the hypothesis that genuine fluctuating asymmetry, measured as the within-individual variance between right and left wings that results from random perturbations in development, would significantly increase after interspecific hybridization. When sires of D. subobscura were mated to heterospecific females following a hybrid half-sib breeding design, F1 hybrid females showed a large bilateral asymmetry with a substantial proportion of individuals having an asymmetric index larger than 5% of total wing size. Such an anomaly, however, cannot be plainly explained by an increase of developmental instability in hybrids but is the result of some aberrant developmental processes. Our findings suggest that interspecific hybrids are as able as their parents to buffer developmental noise, notwithstanding the fact that their proper bilateral development can be harshly compromised. Together with the low correspondence between the co-variation structures of the interindividual genetic components and the within-individual ones from a Procrustes analysis, our data also suggest that the underlying processes that control (genetic) canalization and developmental stability do not share a common mechanism. We argue that the conventional account of decreased developmental stability in interspecific hybrids needs to be reappraised.