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.     

Thermoregulation and flying habits of the Japanese sulfur butterfly Colias erate (Lepidoptera: Pieridae) in an open habitat

This study evaluated the impact of the thermal environment on the flying behavior of male Japanese sulfur butterflies Colias erate searching for females in an open habitat. Thoracic temperature was monitored before and after flight. Mean thoracic temperature of butterflies immediately after landing was consistently higher than, but independent of, ambient temperature. Although ground speed of flying butterflies was different between flight types, air speed against the butterfly was similar across flight types. The excess of thoracic over ambient temperature was lower in flying butterflies than in basking ones, as predicted by a model. This difference appeared to be due to air current, which enhanced heat loss. In a laboratory study, newly eclosed male butterflies were placed under an incandescent lamp to measure their thoracic temperature at different air current speeds. The excess of thoracic over ambient temperature decreased as the speed of air currents increased. When the air current was similar to the air speed against flying butterflies in the field, a substantial decrease occurred in the operative thoracic temperature.     

The adaptive significance of alpine melanism in the butterfly Parnassius phoebus F. (Lepidoptera: Papilionidae)

Summary The adaptive significance of alpine melanism, the tendancy for insects to become darker with increased elevation and latitude, was investigated using the butterfly Parnassius phoebus. The effects on temperature dependent activity of five components of overall wing melanism, as well as size, were examined. The components of wing melanism examined were the transparency of the basal hindwing and distal fore-wing areas, the width of the black patch in the basal hind-wing area and the proportion of black to white scales in that area, and the proportion of the distal fore-wing covered by predominantly black scaling.The body temperature of dead specimens was correlated with air temperature, solar radiation, the width of the black patch at the base of the wings, and the proportion of black to white scales at the base of the wings. The minimum air temperatures and solar radiation levels required for initiation of flight did not vary with wing melanism of P. phoebus, in contrast to the results found for Colias butterflies by Roland (1982). However, under environmental conditions suitable for flight initiation, males with a higher proportion of black to white scales in the basal area of the hind-wing, and wider basal black patches, spent a greater proportion of time in flight at low air temperatures and low insolation. Increased basal wing melanism was also associated with increased movement of males within a population. In contrast, melanism in the distal area of the wings has no effect on activities which are dependant on body temperature. The amount of time spent feeding did not vary with differences in wing melanism. I suggest that in dorsal basking, slow-flying butterflies (Parnassius) basal wing color affects body temperature primarily during flight (rather than while basking), such that butterflies with darker wing bases cool down less rapidly because they absorb more solar radiation during flight.     

EVOLUTION AND COADAPTATION OF THERMOREGULATORY BEHAVIOR AND WING PIGMENTATION PATTERN IN PIERID BUTTERFLIES

This paper addresses the question of how the relationship between morphological structure and functional performance differs in related groups of organisms. I describe the relationship between a suite of phenotypic characters (behavioral posture and the pattern of wing pigmentation) and one function of these characters (thermoregulatory performance) for two groups of butterflies in the family Pieridae, focusing on how behavior and wing pattern interact to affect specific aspects of thermoregulation. Using both natural and experimentally created variation in wing-melanization patterns, I develop and test a series of predictions about the relations among thermoregulatory posture, melanization pattern, body temperature, and flight activity. Results show that increased melanization in different wing regions has positive, negative, or neutral effects in increasing body temperature of Pieris butterflies. The angle of the wings used during basking alters the relative importance of different modes of heat transfer and thereby determines the contribution of different dorsal wing regions to thermoregulation. Experimentally increased dorsal melanization can either increase or decrease the onset of flight activity and can directly alter thermoregulatory posture. For Pieris, dorsal melanization affects basking and flight, while ventral melanization primarily affects overheating. These results are used to generate a functional map relating melanization pattern to thermoregulatory performance in Pieris. Reflectance-basking posture, white background color, and melanization pattern represent coadapted characters in Pieris that interact to determine thermoregulatory performance. The differences in thermoregulatory posture and background color between pierid butterflies in the subfamilies Pierinae and Coliadinae have led to a reorganization and partial reversal of the thermoregulatory effects of melanization pattern. I suggest that this change in the physical mechanism of thermoregulatory adaption in pierids has qualitatively altered the nature of selection on wing-melanization pattern.     

Thermal ecology of Pieris butterflies (Lepidoptera: Pieridae): a new mechanism of behavioral thermoregulation

Summary I document a new mechanism for behavioral thermoregulation, not previously described in animals, called reflectance basking. This behavior, described here for Pieris butterflies, involves the use of the wings as solar reflectors that reflect solar radiation onto the body to increase body temperature. Results show that Pieris require thoracic (body) temperature. between 29° and 40° C in order to take off and fly, and achieve these elevated temperatures by basking. Diurnal patterns of population flight activity are closely correlated with patterns of body temperature during basking. Behavioral studies indicate that 1) Pieris orient to solar radiation, 2) they use thermoregulatory postures consistent with reflectance basking, and 3) they do not use the basking postures found in other Pierid butterflies (i.e., the Coliadinae). There are consistent differences in wing angles used in reflectance basking between Pieris in different subgenera. Results are discussed with respect to thermoregulation and wing color in other Pierid butterflies, and suggest that a re-evaluation of the functional significance of melanization in Pieris is needed.     

Foliage influences forced convection heat transfer in conifer branches and buds

Conifer foliage structures affect branch and bud temperature by altering the development and convective resistance of the thermal boundary layer. This paper examines foliage effects on forced convection in branches and buds of Picea glauca (Moench) Voss and Pinus contorta Dougl. Ex. Loud., two species that represent the range of variation in foliage structure among conifers. Forced convection is characterized by a power law relating Nusselt (heat transfer) and Reynolds (boundary layer development) numbers. Data were collected in a laminar flow wind tunnel for free stream velocities of 0.16-6.95 m s(-1). Scaling parameters were compared against literature values for silver cast branch replicas, a bed of real foliage, cylinders, and tube banks. Foliage structures reduced Nusselt numbers (heat transfer) relative to cylinders, which are typically used to approximate leafless branches and buds. Significantly different scaling relationships were observed for all foliage structures considered. Forced convection scaling relationships varied with foliage structure. The scaling relationships reported here account for variation within populations of branches and buds and can be used to characterize forced convection in a forest canopy.     

A novel method of behavioural thermoregulation in butterflies

Abstract The requirement for efficient thermoregulation has directed the coevolution of specialized morphological and behavioural traits in ectotherms. Adult butterflies exhibit three thermoregulatory mechanisms, termed dorsal, lateral and reflectance basking. In this study, we investigate a potential fourth mechanism whereby individuals perch with their wings fully spread and angled downwards such that the margins are appressed to the substrate. We find that mate‐locating male Hypolimnas bolina (L.) (Nymphalidae) adopt this posture when operational thoracic temperatures are lowest (less than approximately 34 °C). As thoracic temperature increases, males perch with wings increasingly closed and ultimately select shaded microhabitats. Using thermocouple‐implanted dead models, we show that appressed posture individuals warm faster than those adopting the conventional dorsal‐basking (horizontal wing) posture. This thermal advantage is not mitigated by shading of the outer 60–70% of the wing area, which suggests that – as with the conventional dorsal posture – only the basal wing surfaces contribute to heat gain via the absorption of solar irradiation. These investigations suggest that appression represents a novel extension of conventional dorsal basking behaviour in butterflies.     

Thermoregulatory significance of wing melanization in Pieris butterflies (Lepidoptera: Pieridae): physics,posture, and pattern

Summary Pieris butterflies use a novel behavioral posture for thermoregulation called reflectance basking, in which the wings are used as solar reflectors to reflect radiation to the body. As a means of exploring the thermoregulatory significance of wing melanization patterns, I examine the relation of basking posture and wing color pattern to body temperature. A mathematical model of the reflectance process predicts certain combinations of dorsal wing melanization pattern and basking posture that maximize body temperature. Laboratory experiments and field observations show that this model correctly predicts qualitative differences in the relation of body temperature to basking posture based on differences in the extent of dorsal melanization on the wing margins, both between species and between sexes within species of Pieris. This is the first demonstration in insects that coloration of the entire wing surface can affect thermoregulation. Model and experimental results suggest that, in certain wing regions, increased melanization can reduce body temperature in Pieris; this effect of melanization is exactly the opposite of that found in other Pierid butterflies that use their wings as solar absorbers. These results are discussed in terms of the evolution of wing melanization pattern and thermoregulatory behavior in butterflies.     

Takeoff temperatures in Melitaea cinxia butterflies from latitudinal and elevational range limits: a potential adaptation to solar irradiance

1. This study provides evidence that a heliophilic butterfly, the Glanville fritillary (Melitaea cinxia) has adapted differently to environmental variation across latitudes and elevations. 2. In cool air, basking M. cinxia orient themselves perpendicular to the sun's rays to gain heat and take off. During flight, solar heating is reduced because orientation perpendicular to the sun is no longer possible and convective cooling occurs. Consequently, M. cinxia have been shown to suffer net heat loss in flight, even in full sunshine. When flight duration is restricted in this way, the takeoff temperature becomes an important thermal adaptation. 3. Using a thermal imaging camera, takeoff temperatures were measured in experimental butterflies. Butterflies from the northern range limit in Finland took flight at slightly hotter temperatures than butterflies from the southern limit in Spain, and much hotter than butterflies from the elevational limit (1900–2300 m) in the French Alps. Butterflies from low‐elevation populations in southern France also took off much hotter than did the nearby Alpine population. 4. These results suggest that the influence of elevation is different from that of latitude in more respects than ambient temperature. Values of solar irradiance in the butterflies' flight season in each region show that insects from the coolest habitats, Finland and the Alps, experienced similar solar irradiance during basking, but that Finns experienced much lower irradiance in flight. This difference may have favored Finnish butterflies evolving higher takeoff temperatures than Alpine butterflies that also flew in cool air but benefited from more intense radiant energy after takeoff.     

Melanism and diel activity of alpine Colias (Lepidoptera: Pieridae)

Summary The hypothesis that increased melanism provides a benefit in prolonging diel activity through more efficient absorption of solar radiation was tested in the field on a population of alpine Colias sulphur butterflies. A marked increase in the duration of flight and feeding behaviour existed for melanistic individuals when compared to lighter individuals under cool temperatures and low intensity solar radiation. More melanistic butterflies moved longer distance per day, and emigrated from the population at a faster rate. At high temperature and high radiant load lighter coloured individuals appeared more active. This is the first field demonstration of the advantage of melanism for increasing activity of ectotherms in cold environments.     

Behavioural thermoregulation and the relative roles of convection and radiation in a basking butterfly

Poikilothermic animals are often reliant on behavioural thermoregulation to elevate core-body temperature above the temperature of their surroundings. Butterflies are able to do this by altering body posture and location while basking, however the specific mechanisms that achieve such regulation vary among species. The role of the wings has been particularly difficult to describe, with uncertainty surrounding whether they are positioned to reduce convective heat loss or to maximise heat gained through radiation. Characterisation of the extent to which these processes affect core-body temperature will provide insights into the way in which a species׳ thermal sensitivity and morphological traits have evolved. We conducted field and laboratory measurements to assess how basking posture affects the core-body temperature of an Australian butterfly, the common brown (Heteronympha merope). We show that, with wings held open, heat lost through convection is reduced while heat gained through radiation is simultaneously maximised. These responses have been incorporated into a biophysical model that accurately predicts the core-body temperature of basking specimens in the field, providing a powerful tool to explore how climate constrains the distribution and abundance of basking butterflies.     

The role of chemotactile stimuli in the oviposition preferences of Colias butterflies

Summary The legume foodplants of Colias butterflies possess specific chemical components which stimulate oviposition in females of the genus. Different legumes provide different degrees of stimulation, as demonstrated by close correlation between field observations and laboratory experiments with a new behavioral assay. Several generalizations emerge from this study. 1) Chemical preferences for various foodplants are under genetic control in C. meadii, and are at least partly independent of an individual's previous exposure to different legumes. 2) In some cases, chemical cues alone do not allow females to discriminate between species. Long-range search cues may be used to enhance discrimination in such instances. 3) Lupinus, a legume which is not usually utilized by Colias, stimulates oviposition in the laboratory, indicating its chemical affinity with other Leguminosae. The behavioral assay described should prove useful for the future identification of specific oviposition stimulants and deterrents. It is stressed, however, that chemotactile cues are involved only in the final step of oviposition, and that understanding foodplant choice in nature will require in-depth investigation into the mechanics of individual search processes.     

Oxygen transfer in Streptomyces fermentation broths

The oxygen transfer coefficient has been investigated in S. noursci and S. lavendulae fermentation broths obtained from fermentors of different operating volumes (61., 30001., 20,0001.). Fermentors had K_La_s values ranging from 1.0 to 17.0 min^?1, calculated from sulphite oxidation rates. The dynamic measurement of the volumetric oxygen transfer coefficient. (K_La) has been performed in the different fermenting systems. As the fermentation progressed, especially in the first stages, K_La values have decreased in both fermentations and in each system of fermentors. In order to characterise the whole fermenting system an average K_La_s was calculated from the obtained K_La values. The average K_La grew with increasing K_La_s values and ranged from 0.03 to 3.72 min^?l. Some factors possibly having an influence on the, change of K_La have been studied. The oxygen transfer coefficients of the broths have been measured in falling films and ranged from 0.05 to 0.4 cm min^?1. The flow conditions have been characterized by Reynolds numbers of broths varying between 1.0 and 60.0. The average thickness of the falling films have been measured and plotted against Reynolds number. The Re⁺ which is the breaking-point of the plot increased as the fermentation proceeded. In the region of Re⁺ the values of the oxygen transfer coefficient increased rapidly. An approximate correlation could be established between the Re⁺ and the physical properties of fermentation broth.     

Predation,thermoregulation, and wing color in pierid butterflies

Summary This paper explores two hypotheses about the relationships among predation, thermoregulation, and wing color in butterflies: First, that butterflies are susceptible to predation during thermally marginal periods (e.g., cool weather) when effective thermoregulation and flight are not possible; second, that Pieris butterflies are relatively unpalatable to visual predators, supporting the idea that the white wing pigment of Pieris represents aposematic coloration. Field experiments with Pieris and Colias in 1984 and 1985 demonstrate that substantial predation may occur during the morning period before butterflies are able to actively fly. Circumstantial evidence is presented to suggest that at least some of the predation is by small, cursorial mammals. Feeding experiments in the field using Grey Jays as predators indicate that Pieris napi and P. occidentalis are less palatable than other sympatric butterflies, including confamial Colias alexandra. These and previous results suggest that Pieris are edible but less preferred as prey by birds, and that the degree of palatibility may vary among Pieris species. The relatively low palatability of these Pieris is consistent with the hypothesis that their white pigmentation represents aposematic coloration; however, the cues by which potential bird predators might discriminate against Pieris have not been established.     

An evaluation of heat exchange in small birds

Summary Equations are derived that quantify the component thermal resistances to heat transfer in small birds and account explicity for the effects of variation in these resistances. Heat transfer theory is used to quantify external resistances, and an experiment was conducted to estimate body resistance (r _b, plumage plus tissues) as a function of external temperature and wind speed. The value ofr _b decreased with wind speed, and decreased as air temperature approached 0°C. Heat transfer from small birds is shown to be relatively independent of external resistances and mainly dependent onr _b. The predictions of the new theoretical equations are shown to agree well with existing empirical data.     

Gleitflug des FlugbeutlersPetaurus breviceps papuanus

Summary Aerodynamic parameters are determined inPetarums from cine films of gliding flight which partly show an exact side view of gliding flight and landing approach. The outstretched gliding skin does not show any flutter movements. Its relative camber is about 0,1, its position of maximal camber, related to total length, is about of 0,4. Reynolds number is approximately 10⁵. Gliding angles are high, i.e. 20° to 30° during gliding to touch down and up to 60° during aerodynamic braking just before touch down. Gliding numbers are accordingly high (bad). Aerodynamic angles of attack are high (45° to 60°), lying higher than the value of 35° at which the smallest (best) gliding numbers of 0,53 are indicated (gliding angle 28°) by aPetaurus preparation polar, determined in a wind tunnel.Petaurus is able to turn rolls and to fly curves by specific coordinations of flying skin, legs and tail, some of which are described and compared with the 1974 measurements on wind tunnel models.     

DNA barcodes identify Central Asian Colias butterflies (Lepidoptera,Pieridae)

A majority of the known Colias species (Lepidoptera: Pieridae, Coliadinae) occur in the mountainous regions of Central-Asia, vast areas that are hard to access, rendering the knowledge of many species limited due to the lack of extensive sampling. Two gene regions, the mitochondrial COI ‘barcode’ region and the nuclear ribosomal protein RpS2 gene region were used for exploring the utility of these DNA markers for species identification. A comprehensive sampling of COI barcodes for Central Asian Colias butterflies showed that the barcodes facilitated identification of most of the included species. Phylogenetic reconstruction based on parsimony and Neighbour-Joining recovered most species as monophyletic entities. For the RpS2 gene region species-specific sequences were registered for some of the included Colias spp. Nevertheless, this gene region was not deemed useful as additional molecular ‘barcode’. A parsimony analysis of the combined COI and RpS2 data did not support the current subgeneric classification based on morphological characteristics.     

Thermoregulation in resting and active polar bears

Summary Polar bears (Ursus maritimus) regulate their body temperatures both physiologically and behaviourally proportional to their level of activity while within the thermoneutral zone. Core temperatures (T _c=36.9±0.5°C at rest) varied with the 4^th power of walking speed for the two subadult (220 kg) bears tested, whereas subcutaneous temperatures (T _sk=35.3±2.2°C at rest) were closely correlated withT _c but also varied with wind speed (v _a) and ambient temperature (T _a). Radiative fur temperatures (T _r) were closely correlated withT _a and negligibly withT _sk. Predictive equations for these temperature relationships were derived by regression analysis. Maximum rates of heat storage (S _max) were above that predicted from the literature implying that the polar bear is an energetically costly walker. Radiative heat losses of a resting polar bear amount to between 36–67% of the metabolism and assuming a respiratory heat loss of 7–10%, convective heat losses (by difference) would thus range from 33–64%. When walking, the cooling of the fur surface by forced convection and the pendulum effect of the moving legs of the bear lead to estimated convective heat losses on the order of 75% of the heat production while radiative losses are reduced to 13–22%. Increasing wind speeds enhance further this reciprocal effect.     

Free Convection in a Parallelogrammic Porous Cavity Filled with a Nanofluid Using Tiwari and Das’ Nanofluid Model

The free convection heat transfer of Cu-water nanofluids in a parallelogrammic enclosure filled with porous media is numerically analyzed. The bottom and top of the enclosure are insulated while the sidewalls are subject to limited temperature difference. The Darcy flow and the Tiwari and Das’ nanofluid models are considered. The governing dimensionless partial differential equations are numerically solved using a finite difference code. The results are reported for isotherms and streamlines as well as Nusselt number as a function of the volume fraction of nanoparticles, porosity, types of the porous matrix, inclination angle, aspect ratio and different Rayleigh numbers. It is found that the presence of the nanoparticles inside the enclosure deteriorates the heat transfer rate, which is caused due to the increase of dynamic viscosity by the presence of nanoparticles. Therefore, in applications in which the nanofluids are used for their advantages, such as enhanced dielectric properties or antibacterial properties, more caution for the heat transfer design of the enclosure is necessary.     

Gleitflug des FlugbeutlersPetaurus breviceps papuanus (Thomas)

Summary Experiments with models made on thin flat plates were made in front of a wind tunnel in order to determine the relationship between their coefficients of lift and drag (Fig. 2) and the angle of attack as well as their aspects of flow (Fig. 8). The following results were achieved. When the plates were covered on one side with the fur from aPetaurus' flight skin, coefficients of lift at average angles of attack were improved, the critical angle of attack was increased and the characteristics of flow separation were flattened. However, these positive effects were only achieved by physiological orientation of the fur (i.e. placed on the upper surface with fur lying backwards; Fig. 3). The gliding number is improved at very high angles of attack only (Fig. 6). Thus the fur of aPetaurus acts as a lift generator within high (critical) angles of attack, during gliding flight (Nachtigall, 1979). Other natural and synthetic furs show a qualitatively similar, but quantitatively less distinct effect (Fig. 7). The aerodynamic efficiency of a fur coating is due to the boundary layer effects provided by the individual hairs which seem to act as miniature back-flow breaks (Fig. 8,9). The bionic transferability of this effect to technical wings is discussed.

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Mit Unterstützung der Deutschen Forschungsgemeinschaft

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Für melßtechnische Mitarbeit danke ich Frfiulein Hedwig Reichel und Herrn Rainer Grosch. Frau Lore Dinnendahl danke ich für Literaturhinweise.