Thermal physiological ecology of Colias butterflies in flight

Summary As a comparison to the many studies of larger flying insects, we carried out an initial study of heat balance and thermal dependence of flight of a small butterfly (Colias) in a wind tunnel and in the wild.Unlike many larger, or facultatively endothermic insects, Colias do not regulate heat loss by altering hemolymph circulation between thorax and abdomen as a function of body temperature. During flight, thermal excess of the abdomen above ambient temperature is weakly but consistently coupled to that of the thorax. Total heat loss is best expressed as the sum of heat loss from the head and thorex combined plus heat loss from the abdomen because the whole body is not isothermal. Convective cooling is a simple linear function of the square root of air speed from 0.2 to 2.0 m/s in the wind tunnel. Solar heat flux is the main source of heat gain in flight, just as it is the exclusive source for warmup at rest. The balance of heat gain from sunlight versus heat loss from convection and radiation does not appear to change by more than a few percent between the wings-closed basking posture and the variable opening of wings in flight, although several aspects require further study. Heat generation by action of the flight muscles is small (on the order of 100 m W/g tissue) compared to values reported for other strongly flying insects. Colias appears to have only very limited capacity to modulate flight performance. Wing beat frequency varies from 12–19 Hz depending on body mass, air speed, and thoracic temperature. At suboptimal flight temperatures, wing beat frequency increases significantly with thoracic temperature and body mass but is independent of air speed. Within the reported thermal optimum of 35–39°C, wing beat frequency is negatively dependent on air speed at values above 1.5 m/s, but independent of mass and body temperature. Flight preference of butterflies in the wind tunnel is for air speeds of 0.5–1.5 m/s, and no flight occurs at or above 2.5 m/s. Voluntary flight initiation in the wild occurs only at air speeds 1.4 m/s.In the field, Colias fly just above the vegetation at body temperatures of 1–2°C greater than when basking at the top of the vegetation. These measurements are consistent with our findings on low heat gain from muscular activity during flight. Basking temperatures of butterflies sheltered from the wind within the vegetation were 1–2°C greater than flight temperatures at vegetation height.     

Evolutionary genetics of dorsal wing colour in Colias butterflies

The evolution of butterfly wing colouration is strongly affected by its multiple functions and by the correlated evolution of wing colour elements. Both factors may prevent local adaptation to ecological conditions. We investigated one aspect of wing colouration, the degree of dorsal wing melanization, in the butterfly Colias philodice eriphyle across an elevational gradient and its correlation with another aspect of wing colouration, ventral wing melanization. Dorsal wing melanization increased with elevation and these differences persisted in a common environment. Full-sibling analysis revealed high heritability for males but only intermediate heritability for females. The correlation between ventral and dorsal melanization showed significant elevational and sex-specific differences. In males the two traits were highly correlated, whereas in females the strength of the correlation decreased with increasing elevation. We conclude that uncoupling of ventral and dorsal melanization has evolved in females but not in males and discuss possible mechanisms underlying uncoupling.     

Thermoregulation and the determinants of heat transfer in Colias butterflies

Summary As a means of exploring behavioral and morphological adaptations for thermoregulation in Colias butterflies, convective heat transfer coefficients of real and model butterflies were measured in a wind tunnel as a function of wind speed and body orientation (yaw angle). Results are reported in terms of a dimensionless heat transfer coefficient (Nusselt number, Nu) and a dimensionless wind speed (Reynolds number, Re), for a wind speed range typical of that experienced by basking Colias in the field. The resultant Nusselt-Reynolds (Nu-Re) plots thus indicate the rates of heat transfer by forced convection as a function of wind speed for particular model geometries.For Reynolds numbers throughout the measured range, Nusselt numbers for C. eurytheme butterflies are consistently lower than those for long cylinders, and are independent of yaw angle. There is significant variation among individual butterflies in heat transfer coefficients throughout the Re range. Model butterflies without artificial fur have Nu-Re relations similar to those for cylinders. Heat transfer in these models depends upon yaw angle, with higher heat transfer at intermediate yaw angles (30–60°); these yaw effects increase with increasing Reynolds number. Models with artificial fur, like real Colias, have Nusselt numbers which are consistently lower than those for models without fur at given Reynolds numbers throughout the Re range. Unlike real Colias, however, the models with fur do show yaw angle effects similar to those for models without fur.The independence of heat loss from yaw angle for real Colias is consistent with field observations indicating no behavioral orientation to wind direction. The presence of fur on the models reduces heat loss but does not affect yaw dependence. The large individual variation in heat transfer coefficients among butterflies is probably due to differences in fur characteristics rather than to differences in wing morphology.Finally, a physical model of a butterfly was constructed which accurately simulates the body temperatures of basking Colias in the field for a variety of radiation and wind velocity conditions. The success of the butterfly simulator in mimicking Colias thermal characteristics confirms our preliminary understanding of the physical bases for and heat transfer mechanisms underlying thermoregulatory adaptations in these butterflies.     

Sources of intraspecific variation in the hostplant seeking behavior of Colias butterflies

Summary Egg-laying females of the legume-feeding butterfly Colias philodice eriphyle were observed within a high-altitude study site during 1977 and 1978. Average oviposition selectivity showed two patterns of seasonal variation. First, second-brood females laid eggs more frequently after alighting upon hostplants than did first-brood females in both years. Second, a consistent decline in post-alighting oviposition probability near the end of the second brood corresponded with a dramatic decrease in the water content of hostplant foliage near the end of the dry summer season. In addition to seasonal variation in oviposition behavior, individual females landed and oviposited upon widely varying sets of legume hostplant species. By sampling hostplant abundances along the flightpaths of observed females, we show that individuals varied in their tendency to land upon different hostplants. Females tended to specialize upon one or two species, at least in the short term, and the pattern of oligophagy for the population as a whole is partly generated by variation in the host-seeking behavior of individual females.     

An AFLP-based interspecific linkage map of sympatric, hybridizing Colias butterflies

Colias eurytheme and C. philodice are sister species with broad sympatry in North America. They hybridize frequently and likely share a significant portion of their genomes through introgression. Both taxa have been ecologically well characterized and exploited to address a broad spectrum of evolutionary issues. Using AFLP markers, we constructed the first linkage map of Colias butterflies. The map is composed of 452 markers spanning 2541.7 cM distributed over 51 linkage groups (40 major groups and 11 small groups with 2-4 markers). Statistical tests indicate that these AFLP markers tend to cluster over the map, with the coefficient of variation of interval sizes being 1.236 (95% C.I. is 1.234-1.240). This nonrandom marker distribution can account for the nonequivalence between the number of linkage groups and the actual haploid chromosome number (N = 31). This study presents the initial step for further marker-assisted research on Colias butterflies, including QTL and introgression analyses. Further investigation of the genomes will help us understand better the roles of introgression and natural selection in the evolution of hybridizing species and devise more appropriate strategies to control these pests.     

Climate variability slows evolutionary responses of Colias butterflies to recent climate change

How does recent climate warming and climate variability alter fitness, phenotypic selection and evolution in natural populations? We combine biophysical, demographic and evolutionary models with recent climate data to address this question for the subalpine and alpine butterfly, Colias meadii, in the southern Rocky Mountains. We focus on predicting patterns of selection and evolution for a key thermoregulatory trait, melanin (solar absorptivity) on the posterior ventral hindwings, which affects patterns of body temperature, flight activity, adult and egg survival, and reproductive success in Colias. Both mean annual summer temperatures and thermal variability within summers have increased during the past 60 years at subalpine and alpine sites. At the subalpine site, predicted directional selection on wing absorptivity has shifted from generally positive (favouring increased wing melanin) to generally negative during the past 60 years, but there is substantial variation among years in the predicted magnitude and direction of selection and the optimal absorptivity. The predicted magnitude of directional selection at the alpine site declined during the past 60 years and varies substantially among years, but selection has generally been positive at this site. Predicted evolutionary responses to mean climate warming at the subalpine site since 1980 is small, because of the variability in selection and asymmetry of the fitness function. At both sites, the predicted effects of adaptive evolution on mean population fitness are much smaller than the fluctuations in mean fitness due to climate variability among years. Our analyses suggest that variation in climate within and among years may strongly limit evolutionary responses of ectotherms to mean climate warming in these habitats.     

Adaptive accuracy of temperature oviposition preferences in newts

Oviposition-site choice has profound fitness consequences for both a mother and her offspring. The adaptive significance of oviposition behaviour for both generations depends on two rarely considered assumptions: (1) the fit of maternal oviposition preferences with local phenotypic optimum (adaptive accuracy) and (2) the predictability of future conditions for developing offspring based on conditions at the time of oviposition. We examined both assumptions using temperature oviposition preferences (T _p,o) previously measured under laboratory conditions in the alpine newt, Ichthyosaura (formerly Triturus) alpestris. Analyses of temperature time series in the newt natural environment revealed, in agreement with oviposition-site choice of female newts, that T _p,o were closer to phenotypic optima at the water surface than at the maximal depth (bottom). Temperature time series in both depths contained a high proportion of predictable variation, though bottom thermal conditions were more predictable than those at the water surface. We concluded that female newts have to trade the adaptive accuracy of T _p,o for the predictability of future thermal conditions at the time of oviposition.     

栎旋木柄天牛取食与产卵选择性研究

栎旋木柄天牛Aphrodisium sauteri(Matsushita)是南方地区栎林的重要害虫之一。对三清山高海拔栎林中栎旋木柄天牛Aphrodisium sauteri Matsushita幼虫危害状况、成虫补充营养习性、成虫的产卵选择等情况进行了调查。结果表明,栎旋木柄天牛对寄主树木的危害具有明显的区域性,海拔、坡向、温湿度等对虫害发生有较大影响。主要危害小叶青冈(Cyclobalanopsis myrsinaefolia),轻度危害多脉青冈(C.multinervis)、甜槠(Castanopsis eryei)、木荷(Schima superb)等。野外调查和室内试验发现,成虫有取食液体物质的习性,不同营养液对成虫的取食次数具有显著影响。室内产卵选择试验,成虫在8种供试树种的自然枯枝疤痕凹陷处或枝皮裂缝间隙内产卵,不同树种上的产卵量存在显著差异。栎旋木柄天牛对枝条选择性的室内、外研究结果存在差异:室内试验表明,3种寄主树木的不同枝径水平上的产卵量没有显著差异。林间调查表明,栎旋木柄天牛产卵处的高度与小叶青冈胸径之间存在显著的线性相关,成虫更趋向于在枝径15～25mm、皮厚1～2mm的枝条产卵。对小叶青冈测量分析表明,枝条直径与枝皮厚度间存在着极显著的线性相关。     

Orange-derived stimuli regulating oviposition in the Mediterranean fruit fly

Abstract:  Some visual and olfactory host stimuli influencing oviposition of the Mediterranean fruit fly ( Ceratitis capitata Wiedemann) in orange fruit were investigated in a laboratory-reared strain of this species. Mated females of C. capitata were found to be attracted to the same extent by fragrant orange fruits and odourless sham oranges, while unmated females were notably less attracted than mated females by the above objects. Mated females laid significantly more eggs in orange-coloured than in whitish (optically neutral) spherical wax dummies (diameter, 9.0 cm) as well as in both orange-coloured and whitish wax dummies supplemented with high internal humidity (80–100%) compared with respective wax dummies supplied with low internal humidity (30–50%). The orange-like colour together with high internal humidity provides basic stimuli supporting adequate oviposition of C. capitata in oranges. The oviposition rate of female C. capitata was not significantly changed when graded dosages ranging from 0.004 to 19.6  μ l of orange peel oil per cm² were added to orange-coloured wax dummies, while oviposition was considerably subdued by addition of 3.9  μ l orange peel oil and completely disrupted by addition of 9.8 or 19.6  μ l of orange peel oil per cm² of whitish wax dummies. Oviposition of C. capitata in ripe orange fruit may thus be interpreted by the predilection of this tephritid species for an orange-coloured, glossy pericarp, being capable of counteracting the deterrent effect of the essential oil found in orange flavedo.     

Odour maps in the brain of butterflies with divergent host-plant preferences

Butterflies are believed to use mainly visual cues when searching for food and oviposition sites despite that their olfactory system is morphologically similar to their nocturnal relatives, the moths. The olfactory ability in butterflies has, however, not been thoroughly investigated. Therefore, we performed the first study of odour representation in the primary olfactory centre, the antennal lobes, of butterflies. Host plant range is highly variable within the butterfly family Nymphalidae, with extreme specialists and wide generalists found even among closely related species. Here we measured odour evoked Ca(2+) activity in the antennal lobes of two nymphalid species with diverging host plant preferences, the specialist Aglais urticae and the generalist Polygonia c-album. The butterflies responded with stimulus-specific combinations of activated glomeruli to single plant-related compounds and to extracts of host and non-host plants. In general, responses were similar between the species. However, the specialist A. urticae responded more specifically to its preferred host plant, stinging nettle, than P. c-album. In addition, we found a species-specific difference both in correlation between responses to two common green leaf volatiles and the sensitivity to these compounds. Our results indicate that these butterflies have the ability to detect and to discriminate between different plant-related odorants.     

Host-plant susceptibility to the carrot fly, Psila rosae. 3. The role of oviposition preferences and larval performance

The acceptability of various plant species to ovipositing carrot flies was weakly, but significantly correlated with the host's suitability for larval development. Both adult host-plant preferences and larval performance as determined in laboratory experiments explained a part of the variation in susceptibility among the various test plants observed in the field. Across the whole set of plant species examined, antixenosis contributed more substantially to resistance than antibiosis, while the reverse seemed to be true for carrot cultivars.     

Population structure of pierid butterflies IV. Genetic and physiological investment in offspring by male Colias

Summary Population structure encompasses all the rules by which a population's gametes come together, including genetic and physiological investment in offspring. We document female use of nutrients donated by males at mating, and complete sperm precedence, in Colias eurytheme Boisduval. The effect of these phenomena on the population structure of this species is discussed.     

Height preferences for oviposition of aphidophagousSyrphidae [Diptera]

Résumé La hauteur préférentielle de ponte a été étudiée chez plusieurs espèces deSyrphidae aphidophages. A cet effet, le nombre d'œufs pondus sur des fèveroles en pot (Vicia faba L.), infestées parAphis fabae Scop. et fixées à hauteur de 180 cm, 120 cm et 30 cm a été enregistré. Des hauteurs préférentielles bien précises ont été trouvées pour les différentes espèces, variant de celle deSyrphus triangulifer (Zett.) qui pond 69% des œufs à 180 cm et seulement 5% à 30 cm, à celle deLeucozona lucorum (L.) qui ne pond que 8% à 180 cm, mais 78% à 30 cm.S. ribesii (L.) préfère initialement 180 cm mais le pourcentage de pontes à 30 cm augmente en fonction de l'age. En ce qui concerne les fèveroles en cage, la couleur de la couche inférieure modifie la hauteur préférentielle de ponte.S. balteatus (Deg.), par exemple, pond de préférence à 30 cm lorsque le fond de la cage est recouvert d'herbe ou de papier vert mais ne montre pas de préférence particulière lorsque le sol est tout simplement de la terre ou est recouvert de tissu brun, argenté ou jaune. Il est possible que la couleur verte serve de point de repère visuel à partir duquel la femelle estime la hauteur de ponte.      

A mitochondrial-DNA-based phylogeny for some evolutionary-genetic model species of Colias butterflies (Lepidoptera, Pieridae)

We study the phylogenetic relationships among some North American Colias ("sulfur") butterflies, using mitochondrial gene sequences (ribosomal RNA, cytochrome oxidase I+II) totaling about 20% of the mitochondrial genome. We find that (1) the lowland species complex shows a branching order different from earlier views; (2) several montane and northern taxa may be more distinct than in earlier views; (3) one morphologically conservative Holarctic assemblage, C. hecla, is differentiated at the molecular-genetic level into at least three taxa which occupy distinct positions in the phylogeny and are sisters to diverse other taxa. These conclusions, constituting phylogenetic hypotheses, are supported by parsimony, maximum-likelihood, and Bayesian reconstruction algorithms. They are tested formally, by interior branch tests and paired-site tests, against alternative hypotheses derived from conventional species and subspecies naming combinations. In all cases our hypotheses are supported by these tests and the conventional alternatives are rejected. The "barcoding" subset of cytochrome oxidase I sequence identifies only some of the taxa supported by our full data set. Comparison of genetic divergence values among Colias taxa with those among related Pierid butterflies suggests that species radiations within Colias are comparatively younger. This emerging Colias phylogeny facilitates comparisons of genetic polymorphism and other adaptive mechanisms among taxa, thereby connecting micro- and macro-evolutionary processes.     

Mothers matter too: benefits of temperature oviposition preferences in newts

The maternal manipulation hypothesis states that ectothermic females modify thermal conditions during embryonic development to benefit their offspring (anticipatory maternal effect). However, the recent theory suggests that the ultimate currency of an adaptive maternal effect is female fitness that can be maximized also by decreasing mean fitness of individual offspring. We evaluated benefits of temperature oviposition preferences in Alpine newts (Ichthyosaura [formerly Triturus] alpestris) by comparing the thermal sensitivity of maternal and offspring traits across a range of preferred oviposition temperatures (12, 17, and 22°C) and by manipulating the egg-predation risk during oviposition in a laboratory thermal gradient (12-22°C). All traits showed varying responses to oviposition temperatures. Embryonic developmental rates increased with oviposition temperature, whereas hatchling size and swimming capacity showed the opposite pattern. Maternal oviposition and egg-predation rates were highest at the intermediate temperature. In the thermal gradient, females oviposited at the same temperature despite the presence of caged egg-predators, water beetles (Agabus bipustulatus). We conclude that female newts prefer a particular temperature for egg-deposition to maximize their oviposition performance rather than offspring fitness. The evolution of advanced reproductive modes, such as prolonged egg-retention and viviparity, may require, among others, the transition from selfish temperature preferences for ovipositon to the anticipatory maternal effect.     

Low oviposition stimuli reduce egg production in the pollen beetle Meligethes aeneus

Abstract. Gravid female pollen beetles, Meligethes aeneus (F.) (Coleoptera: Nitidulidae), were exposed at different intervals to oviposition sites that varied in acceptability.The egg load of dissected individuals which had been deprived of oviposition sites was not greater than individuals which had been frequently exposed to oviposition sites.However, the egg load of individuals which had been exposed to highly acceptable host plants was greater than those exposed to a host plant of low acceptability.Over the experimental period the total number of eggs which an individual produced was lower for those that were either deprived of an oviposition site or frequently exposed to a low-quality host plant.There was no evidence of either increased oviposition probability or clutch size as the period since the last oviposition increased.It is concluded that low host quality and low host encounter rate reduce the egg production of M.aeneus. It is further concluded that the suppressed egg production reduces the accumulation of eggs, so that physiological motivation for the insect to oviposit is not increased.If low oviposition site quality acts to reduce oogenesis, as found here, then the probability of egg deposition on low-quality species may not increase over time and the effect of antixenotic resistance may be enhanced.     

Host plant stimuli affecting oviposition behavior of the Eastern spruce budworm

The host selection of the female spruce budworm Choristoneura fumiferana (Clem.), Tortricidae, for oviposition is influenced both by the shape and the composition of the substrate. Needle extracts were bioassayed using artificial twigs. The active compounds are soluble in organic solvents and steam distillable. Synthetic D- and L- pinene occurring naturally in the extracts proved to be oviposition stimulants. The optical isomer L- pinene did not show any activity. Ablation experiments led to the conclusion that chemoreceptors for the host attractants and the stimulants are located on the antennae and probably also on the maxillary and labial palpi.

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Zusammenfassung Die Weibchen von Choristoneura fumiferana bevorzugen für die Eiablage ein nadelförmiges Substrat. Dabei spielen die mechanorezeptiven Sinneshaare auf den Ovipositor-Papillen vermutlich eine entscheidende Rolle. Petrolätherextrakte der Nadeloberflächen, Methanolextrakte und Wasserdampfdestillate ergaben eine erhöhte Eiablage auf damit behandelten künstlichen Nadelzweigen. Die aktiven Extraktfraktionen waren nur in organischen Lösungsmitteln löslich. Für die Attraktivität und stimulierende Wirkung der Extrakte müssen verschiedene Substanzen verantwortlich sein. - und -Pinen wurden mit Hilfe des Gaschromatographen in den aktiven Fraktionen nachgewiesen. Synthetisches D- und L- Pinen waren beide aktiv. Dagegen ergab L- Pinen bei den geprüften Konzentrationen keine erhöhte Eiablage. Mit Hilfe von Ablationsexperimenten (Amputation, Überziehen mit Paraffin) wurde versucht, die Chemorezeptoren für die attraktiven und stimulierenden Stoffe zu lokalisieren. Dabei ergab sich, dass die Geruchsrezeptoren auf den Antennen und vermutlich auch auf den Labial-und Maxillarpalpen für die Wirtswahl von Bedeutung zu sein scheinen.

     

The role of olfaction in canine food preferences

A bland diet with the odor of meat or air blown through it waspresented to Beagle dogs in daily two choice preference tests.At first, there was a 70% preference for the food that smelledlike meat but, by the third week, the preference had fallento 52%. These results indicated that, although dogs initiallyprefer food that smells like meat, odor must be paired withsome other property of meat, probably taste, for the preferencesto be sustained. Intact dogs significantly preferred lamb overhorsemeat 67±4%, pork over lamb 89±3%, pork overhorsemeat 89±3%, beef over lamb 84±2%, and beefover horsemeat 89±2% in two choice preference tests.Dogs made anosmic by intranasal infusion of zinc sulfate hadpreferences significantly less than those of intact dogs anddid not show a significant preference for one meat over anotherwith the exception of pork, which was preferred to lamb (61±4%).Anosmic dogs showed preferences similar to those of intact dogsfor a sucrose containing over a nonsucrose diet and for a horsemeatcontaining diet over a non-meat diet. These results indicatethat olfaction is important in canine food preferences whichinvolve discrimination between meats, but not for sweet versusnon-sweet and meat versus non-meat preferences.