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.     

Thermoregulatory physiology of the carpenter bee,Xylocopa varipuncta

Summary The carpenter beesXylocopa varipuncta maintain thoracic temperatures of 33.0°C to 46.5°C during continuous free flight from 12°C to 40°C. Since the thoracic temperature excess is not constant (decreasing from 24°C at low air temperatures to 6°C at high) the bees are thermoregulating. We document physiological transfer of relatively large amounts of heat to the abdomen and to the head during pre-flight warm-up and during artificial thoracic heating. Most of the temperature increase of the head is due to passive conduction, while that of the abdomen is due to active physiological heat transfer despite a series of convolutions of the aorta in the petiole that anatomically conform to a counter-current heat exchanger. Although the thermoregulatory mechanisms during flight are far from clarified, our data suggest that thermoregulation involves a strong reliance on active convective cooling through increased flight speed.     

Influence of acclimation temperature on the shivering behavior of the butterfly Danaus plexippus (L.)

Summary Most of the monarch butterflies kept at 4–5° C for a few days shivered when released at a test temperature of 15–16° C, whereas fewer of the butterflies kept at 23–24° C did so. Cold-acclimated butterflies shivered more readily, as indicated by the length of the interval between release at the test temperature and the onset of shivering, and they shivered for longer periods of time. The effects of cold acclimation were reversible, but in only 1 out of 3 replicates was the warm acclimation clearly reversed. Cool animals shivered at room temperature, indicating that body temperature and not ambient temperature is important in releasing the behavior. It is suggested that the acclimation involves alteration in the central neurons controlling the activity of muscles involved in shivering.I thank Miss Janice Ruppert and Mr. C. J. Doughty for their valuable technical assistance. The co-operation of the administrators of New Brighton Beach State Park in permitting me to collect in the park is appreciated.     

The physiology of hypothermia in the black-capped Chickadee,Parus atricapillus

Summary The shivering, body temperature, and metabolic response to stable and decreasing ambient temperature were measured in winter acclimatized Black-capped Chickadees,Parus atricapillus. Shivering activity, measured by duration and amplitude of bursts, increased curvilinearly from thermoneutral temperatures of 27°C down to 0°C. This parabolic shivering response may be a major component of the curvilinear response of metabolism to decreasing ambient temperature.Birds exposed to 0°C exhibited metabolism 32–45% lower than predicted for a 12-g homeotherm and body temperatures 10°C below the pre-experimental nocturnal body temperature. This hypothermia was not the result of a breakdown in thermoregulation, but was a controlled effort serving to reduce overnight energy expenditure. It is suggested that (1) hypothermia was achieved by decreased shivering by pectoral muscles during exposure to decreasing ambient temperatures, (2) the rate of body temperature decline was moderated by intermittent and reduced bursts during the cooling period, and (3) body temperature was maintained at a particular level during exposure to a stable low ambient temperature by intense bursts lasting one to three minutes.The physiology of hypothermia in chickadees is similar to torpor; however, chickadees did not arouse to a normal diurnal body temperature in the laboratory, and their hypothermia was not induced by inanition or prolonged exposure to cold, as reported for other species capable of torpor.     

Genotypic and environmental effects on flight activity and oviposition in the Glanville fritillary butterfly

Adverse environmental conditions constrain active flight and thereby limit reproduction in most insects. Butterflies have evolved various adaptations in order to thermoregulate, allowing females to search for nectar and to oviposit under unfavorable thermal conditions. We studied experimentally and with observational data the effect of low ambient temperatures experienced in the morning on the timing of oviposition and clutch size in the Glanville fritillary butterfly (Melitaea cinxia). Comparisons were made between individuals with different forms of the gene Pgi, encoding the glycolytic enzyme phosphoglucose isomerase, since naturally segregating variation at Pgi is known to be correlated with flight metabolic rate, flight performance, and fecundity. Experiencing low temperature in the morning delayed the initiation of oviposition and decreased clutch size. We used a thermal image camera to measure the thoracic surface temperature of butterflies immediately after voluntary flight. Single nucleotide polymorphism at Pgi was associated with thoracic temperature at low ambient temperatures. This has consequences for reproduction because females that are able to fly at lower ambient temperatures generally initiate oviposition earlier in the afternoon, when the environmental conditions are most favorable and the average egg clutch size is generally largest. These results suggest that variation in physiological and molecular capacity to sustain active flight at low ambient temperature has significant fitness-related consequences in insects.     

Synchronous Crepuscular Flight of Female Asian Gypsy Moths: Relationships of Light Intensity and Ambient and Body Temperatures

Female gypsy moths (Lymantria dispar) of Asian heritage studied in central Siberia and Germany exhibit a highly synchronous flight at dusk, after light intensity falls to about 2 lux. This critical light intensity sets the timing of flight behaviors independent of ambient temperature. Flight follows several minutes of preflight wing fanning during which females in Germany and those from a laboratory colony (derived from Siberian stock) raised their thoracic temperatures to 32–33°C at ambient temperatures of 19–22°C. Thoracic temperature of females in free flight exceeded the air temperature (19–22°C) by approximately 11–13°C. The duration of wing fanning was strongly dependent on ambient temperature. In Germany, where ambient temperatures at dusk ranged between 21 and 25°C, females wing fanned for only 2.1 ± 0.2 (SE) min; in the much colder temperatures prevalent at dusk in Bellyk, central Siberia (11–13°C), females spent 11.2 ± 0.6 min in preflight wing fanning. The majority (80%) of mated and even virgin females initiated flight during the evening of the day they eclosed. However, in Bellyk, a small proportion (12%) of females wing fanned for an extended time but then stopped, whereas others (8%) never wing fanned and, therefore, did not take flight. Females also were capable of flight when disturbed during the daylight hours in Germany where the maximal temperature was high (27–30°C), but not in Siberia, where temperatures peaked at only 17–19°C. However, Siberian females were able to propel themselves off the tree on which they were perched by executing several vigorous wing flicks when approached by the predaceous tettigoniid, Tettigonia caudata.     

Effects of ambient temperature,humidity, and age on wing-beat frequency of Periplaneta species

Wing-beat frequency of 2-week-old male Periplaneta americana cockroaches was measured during tethered flight at ambient temperatures from 19 to 35°C and 50 and 95% r.h. Between 19 and 27°C the frequency increased at a nearly constant rate (0·6 Hz/deg at 50% r.h.; 0·7 Hz/deg at 95% r.h.), but the rate of increase decreased markedly between 27 and 35°C. This decline differed from results of past investigations of wing-beat frequency for this species. Wing-beat frequencies at 95% r.h. were significantly higher than those at 50% r.h. for ambient temperatures between 27 and 35°C. reductions in wing-beat frequency at high temperatures and differences in wing-beat frequency between 50 and 95% r.h. may reflect decreases in internal thoracic temperature resulting from cuticular water loss.     

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.     

Body surface temperatures of jerboas (Allactaga) in uniform thermal environments

Summary Body surface temperatures of threeAllactaga elater and oneA. hotsoni were measured by infrared radiography at ambient temperatures of 1° to 42°C. In each test the radiant temperature of environmental surfaces was the same as air temperature.At ambient temperatures of 40–42°C, the temperature of the entire body surface was close to ambient temperature. As ambient temperature was lowered toward 1°C, forehead and back temperatures became increasingly greater than ambient temperature (Fig. 3), indicating an increasing thermal flux across these parts of the body. Forehead and back temperatures were linear functions of ambient temperature below thermoneutrality and behaved as expected according to a model of thermal exchange developed here. The surface temperature of the extraordinarily large pinnae remained close to ambient temperature down to 10°C (Fig. 3), indicating that deep pinna temperature likely falls with decreasing ambient temperature and that the pinnae, despite their size, are not major sites of heat loss at low ambient temperatures.     

Thermoregulation in four species of tropical solitary bees: the roles of size,sex and altitude

Body temperatures during free flight in the field, warm-up rates during pre-flight warm-up, and temperatures during tethered flight are measured for four tropical solitary bee species at three sites of differing altitude in Papua New Guinea. All four species are capable of endothermic preflight warm-up; three species give slopes of thoracic temperature on ambient temperature of significantly less than 1, indicating regulation of thoracic temperature. In the kleptoparasitic Coelioxys spp. (Megachilidae) and Thyreus quadrimaculatus (Anthophoridae), warm-up rates and thoracic temperatures in flight are low by comparison with the two provisioning species Creightonella frontalis (Megachilidae) and Amegilla sapiens (Anthophoridae). In both C. frontalis and A. sapiens thoracic temperatures correlate positively and significantly with both ambient temperature and body mass. In A. sapiens, body mass increases with altitude; this can be interpreted as a response to lower ambient temperatures at higher altitude, an example of Bergmann's rule. In both A. sapiens and C. frontalis populations at higher altitude have higher thoracic temperatures independent of differences of body mass, suggestive of additional morphological or physiological adaptation to lower ambient temperatures. In A. sapiens there is no qualitative difference in body temperatures between males and females after controlling for body mass, while male C. frontalis have significantly lower thoracic temperatures than females of the species. This difference between A. sapiens and C. frontalis is discussed with reference to variation in mating systems found in the Apoidea.Abbreviations C.R.I. Christensen Research Institute - P.N.G. Papua New Guinea - SFT stable flight temperature - T _a ambient air temperature - T _ab abdominal temperature - T _dif the temperature difference between thorax and abdomen - T _ex thoracic temperature excess - VFT voluntary flight temperature     

Responses and song pattern copying of Omega-type I-neurons in the cricket,Gryllus bimaculatus,at different prothoracic temperatures

Summary Omega-type I-neurons (ON/1) (Fig. 1A) were recorded intracellularly with the prothoracic ganglion kept at temperatures of either 8–9°, or 20–22° or 30–33 °C and the forelegs with the tympanal organs kept at ambient temperature (20–22 °C). The neurons were stimulated with synthetic calling songs (5 kHz carrier frequency) with syllable periods (SP in ms) varying between 20 and 100, presented at sound intensities between 40 and 80 dB SPL. The amplitude and duration of spikes as well as response latency decreased at higher temperatures (Figs. 1 B, 2, 6). At lower prothoracic temperatures (8–9 °C) the neuron's responses to songs with short SP (20 ms) failed to copy single syllables, or with moderate SP (40 ms) copied the syllable with low signal to noise ratio (Fig. 3). The auditory threshold of the ON/1 type neuron, when tested with the song model, was temperature-dependent. At 9° and 20 °C it was between 40 and 50 dB SPL and at 33 °C it was less than 40 dB SPL (Fig. 4). For each SP, the slope of the intensity-response function was positively correlated with temperature, however, at low prothoracic temperatures the slope was lower for songs with shorter SPs (Fig. 5). The poor copying of the syllabic structure of the songs with short SPs at low prothoracic temperatures finds a behavioral correlate because females when tested for phonotaxis on a walking compensator responded best to songs with longer SPs at a similar temperature.Abbreviations epsps excitatory postsynaptic potentials - ON/1 omega-type I-neuron - SP syllable period - SPL sound pressure level     

Thermal biology of flight in a butterfly: genotype,flight metabolism,and environmental conditions

Knowledge of the effects of thermal conditions on animal movement and dispersal is necessary for a mechanistic understanding of the consequences of climate change and habitat fragmentation. In particular, the flight of ectothermic insects such as small butterflies is greatly influenced by ambient temperature. Here, variation in body temperature during flight is investigated in an ecological model species, the Glanville fritillary butterfly (Melitaea cinxia). Attention is paid on the effects of flight metabolism, genotypes at candidate loci, and environmental conditions. Measurements were made under a natural range of conditions using infrared thermal imaging. Heating of flight muscles by flight metabolism has been presumed to be negligible in small butterflies. However, the results demonstrate that Glanville fritillary males with high flight metabolic rate maintain elevated body temperature better during flight than males with a low rate of flight metabolism. This effect is likely to have a significant influence on the dispersal performance and fitness of butterflies and demonstrates the possible importance of intraspecific physiological variation on dispersal in other similar ectothermic insects. The results also suggest that individuals having an advantage in low ambient temperatures can be susceptible to overheating at high temperatures. Further, tolerance of high temperatures may be important for flight performance, as indicated by an association of heat‐shock protein (Hsp70) genotype with flight metabolic rate and body temperature at takeoff. The dynamics of body temperature at flight and factors affecting it also differed significantly between female and male butterflies, indicating that thermal dynamics are governed by different mechanisms in the two sexes. This study contributes to knowledge about factors affecting intraspecific variation in dispersal‐related thermal performance in butterflies and other insects. Such information is needed for predictive models of the evolution of dispersal in the face of habitat fragmentation and climate change.     

Threshold for shivering in aerobic and anaerobic muscles in bantam cocks and incubating hens

Summary Electromyographic activity (EMG) from the musculus pectoralis (breast muscle), m. iliotibialis (thigh muscle) and m. gastrocnemius (leg muscle), cloacal temperature (Tb) and O₂ consumption were measured in bantam cocks (Gallus domesticus) exposed to different ambient temperatures (Ta). The same parameters were measured in bantam hens incubating eggs artificially thermoregulated to 40° and 25°C (Te).EMG activity appeared in thigh and leg muscles at Ta below 32°C (Tsh). This temperature probably represents the thermoneutral temperature (TNT) of the cock. EMG activity in breast muscles appeared at Ta below 20°C, or 4°C below the lower critical temperature (Tc).All muscles were quiet when the hen incubated 40°C egg at Ta=Tsh. When Te was abruptly changed to 25°C, EMG activity in the iliotibialis muscle appared 3 min before the activity in the pectoralis muscle. Tb dropped from 41.2° to 40.6°C in 14 min. When Te was returned to 40°C, the EMG activity in the pectoralis muscle disappeared almost at once, while the iliotibialis muscle was active until Tb returned to normal.Aerobic muscles seem to be responsible for shivering thermogenesis between Tc and Tsh, while anaerobic muscles are recruited at lower Ta or when the heat loss during incubation becomes severe.Abbreviations EMG electromyography - Ta ambient temperature - Tb cloacal temperature - Tc lower critical temperature - Te egg temperature - TNT thermoneutral temperature - Tsh shivering threshold temperature     

Honeybee waggle dances: the “energy hypothesis” and thermoregulatory behavior of foragers

Honeybees were trained to visit artificial feeding sites containing a 2 mol·1^-1 sucrose solution. To reach the feeder they either had to walk through 3 m of Teflon tube, or fly 20 m or 65 m and then walk through 3 m of tube. Only individuals that flew at least 65 m performed waggle dances. The distance indicated in these waggle dances, judged by the number of wagging movements per wagrun, was the same regardless of whether individuals had to run an additional 3 m of tube after flight or not. The energy needed during walking after flight was determined by measuring O₂ consumption. All individuals attempted to regulate their body temperatures between 36 and 42°C during walking and feeding (O₂ consumption=40l·min^-1 per bee). Calculations show that this walking through 3 m of tube requires as much energy as flying 128 m (difference between thoracic and ambient temperature=15°C). This energy expenditure was not reflected in the dances. The results do not support the hypothesis that honeybees estimate feeding site distances by measuring the energy required to reach a feeder.Abbreviations T_a ambient temperature - T _b body temperature - T _th thorax temperature     

Spring behavioral patterns of the apple blossom weevil

Individual pairs of overwintered adult apple blossom weevils, Anthonomus pomorum (L.), confined with apple twigs under different ambient temperatures in the laboratory and on apple trees in the field, were observed through day and night for their spring activities. Flight behavior in relation to ambient temperature was also investigated under laboratory conditions using flight stands. Both sexes displayed predominantly nocturnal behavior patterns in both the laboratory and the field. Feeding, crawling, and mating activities increased following sunset in the field or onset of scotophase in the laboratory while resting occurred most frequently during daylight hours. Results of the laboratory experiments showed that temperature affected significantly the activity patterns. The diel pattern of activities became less distinctive at higher temperatures (above 15°C), and total activities in crawling, feeding, and mating were suppressed significantly at lower temperatures (below 5°C). Over 97% of the test weevils initiated take-off response from flight stands at 20°C within the 30 min trial period; however, flight initiation rarely occurred at temperatures 12°C or below. Overall, results of the laboratory and field experiments indicate that A. pomorum is a remarkably cold-adapted insect with ability to crawl, feed, and mate at a few degrees above freezing, a physiological attribute necessary for the exploitation of early stages of apple bud development in the cold early spring.     

Interactions between size and temperature influence fecundity and longevity of a tortricid moth,Zeiraphera canadensis

Females of Zeiraphera canadensis Mut. & Free., the spruce bud moth, were reared in the laboratory at constant and alternating temperatures, and in an outdoor insectary, to (1) determine the effects of temperature, age and size on several reproductive parameters and, (2) to test the hypothesis that body size-temperature interactions influence longevity and realized fecundity. Egg maturation was linearly related to age and large moths developed eggs at a higher rate than small ones. Mcan lifetime oviposition rate reached a maximum and remained stable at temperatures 20° C while the mean lifetime rate of egg maturation increased linearly with temperature, indicating that higher temperatures adversely affect oviposition. The production of nonviable eggs increased with age but also with temperature, suggesting high temperature (25° C) reduces egg quality and/or hinders fertilization. The realized fecundity and longevity of females reared under an alternating temperature regime (mean 20° C) was significantly less than that of females reared at constant 20° C. Similar realized fecundity, longevity and mean lifetime oviposition rates for females reared at temperatures alternating between 10 and 25° C (mean 20° C) and those at constant 25° C reflected the inability of females to recover from elevated diurnal temperatures. Longevity was positively related to female body size at constant 15 and 20° C but the relationships were negative for moths exposed to diurnal temperatures equal to or exceeding 25° C. Due to the reduced longevity of large moths at high temperatures, linear regressions between size and realized fecundity were only significant at constant temperatures 20° C. At higher temperatures, the size-fecundity relationship became curvilinear as a result of the diminished reproductive output of large individuals. Reduced fecundity and longevity of large females at high temperatures may have been due to elevated internal temperatures of large-bodied moths. Large females in a controlled-environment chamber maintained at 25° C developed an internal temperature excess (i.e. temperature above ambient) of nearly 2° C while small-bodied females exceeded ambient by only 0.3° C. However, when held at 20° C, the temperature excess of large-bodied moths was much less than 1° C and small-bodied females did not differ from ambient. Such interactions between temperature and body size suggest that there should be stabilizing selection toward moderate-sized individuals and may explain the absence of size-related effects on fecundity and longevity previously reported for several other lepidopterans.     

Effects of temperature on properties of flight neurons in the locust

High ambient temperatures increase the wing-beat frequency in flying locusts, Locusta migratoria. We investigated parameters of circuit and cellular properties of flight motoneurons at temperatures permissive for flight (20–40 °C). As the thoracic temperature increased motoneuronal conduction velocity increased from an average of 4.40 m/s at 25 °C to 6.73 m/s at 35 °C, and the membrane time constant decreased from 11.45 ms to 7.52 ms. These property changes may increase locust wing-beat frequency by affecting the temporal summation of inputs to flight neurons in the central circuitry. Increases in thoracic temperature from 25–35 °C also resulted in a hyperpolarization of the resting membrane potentials of flight motoneurons from an average of-41.1 mV to -47.5 mV, and a decrease of input resistances from an average of 3.45 M to 2.00 M. Temperature affected the measured input resistance both by affecting membrane properties, and by altering synaptic input. We suggest that the increase in conduction velocity Q₁₀=1.53) and the decrease of membrane time constant (Q₁₀=0.62) would more than account for the wing-beat frequency increase (Q₁₀=1.15). Hyperpolarization of the resting membrane potential (Q₁₀=1.18) and reduction in input resistance (Q₁₀=0.54) may be involved in automatic compensation of temperature effects.Abbreviations ANOVA analysis of variance - CPG central pattern generator - DL dorsal longitudinal muscles - EMG electromyographic - MN motoneuron - PSP post synaptic potential - Q₁₀ temperature coefficient - RMP resting membrane potential - S.D. standard deviation - SR stretch receptor     

Response characteristics of cold cell on the antenna ofLocusta migratoria L.

Summary The dendritic outer segment of the cell which is most likely the cold unit in the poreless coeloconic sensilla onLocusta migratoria antennae, has finger-like projections up to 1.5 m long and 0.13 m thick (Fig. 1). This unit responds to constant temperature, to slowly changing temperature and to step changes. Under stationary conditions impulse frequency attained 35 imp/s. Between 14 °C and 41 °C the higher frequencies were associated with the higher temperatures (Fig. 5). In this range the differential sensitivity is positive but not large: + 0.8 (imp/s)/°C. Its resolving power for steady temperature is 4.7 °C.Downward step changes produced by shifting between airstreams at different temperatures yield far higher frequencies (Figs. 2, 3). Step amplitudes were between –0.1 °C and –12 °C; the conditioning temperature from which the steps were initiated, was between 16 °C and 33 °C. Frequency peaked during the first 50 ms after stimulus onset (Fig. 2) and reached its highest values (310–340 imp/s) at initial temperatures above 30 °C and steps larger than –10 °C (Fig. 4). The mean differential sensitivity from 23 curves was –19 (imp/s)/°C and the resolving power 0.6 °C.During slowly changing temperature the impulse frequency was governed by two parameters simultaneously: ambient temperature and its rate of change. Rates were between 0.001 °C/s or less, and 0.03 °C/s in either direction. Frequency was higher during slow cooling at a given temperature than during slow warming (Fig. 6). The average differential sensitivity to the rate of change was –210 (imp/s)/(°C/s). Further, the larger responses to cooling developed at lower ambient temperatures (differential sensitivity: –1.0 (imp/s)/°C). It is to be noted that this sign is negative, in contrast to the sign for differential sensitivity to constant temperature and also for the influence of initial temperature on the response to downward step changes.Abbreviations b Slope of characteristic curve, differential sensitivity - F impulse frequency in imp/s - imp/s impulses/s - P _w partial pressure of water vapor in torr - r correlation coefficient - T temperature in °C - T T-step - x resolving power in °C     

Patterns of daily flight activity in onitine dung beetles (Scarabaeinae: Onitini)

Different species of African dung beetles emerge from the soil at characteristic times of the day to fly and colonize the freshly-deposited dung of mammalian herbivores. Onitine dung beetles in their natural habitat displayed one of five distinctive daily flight behaviours: dusk crepuscular (Onitis alexis Klug, O. caffer Boheman, O. fulgidus Klug, O. tortuosus Houston, O. vanderkelleni Lansberge, O. westermanni Lansberge); dusk/dawn crepuscular (O. pecuarius Lansberge and O. viridulus Boheman); dusk/dawn crepuscular and nocturnal (O. aygulus (Fabricius), O. mendax Gillet, O. uncinatus Klug); late afternoon-dusk and dawn-early morning [Heteronitis castelnaui (Harold)]; or diurnal flight activity [O. belial (Fabricius), O. ion (Olivier)]. These diagnostic daily flight behaviours span a light intensity range of over 6 orders of magnitude and have been retained in selected species introduced into Australia. Ambient light intensity appears to be the primary determinant of the daily flight period in onitine dung beetles. Because the dung of mobile herbivores is rapidly exploited by onitine species for feeding and breeding purposes, different flight behaviours result in a spatial and temporal partitioning of species in the local dung beetle community. The timing of flight may contribute to, or lead to avoidance of, competition between species which may ultimately affect colonization success. Many onitines show a strong preference for dung of specific herbivores, which may further reduce interspecific competition. All crepuscular-nocturnal species examined raised their thoracic temperatures endothermically to between 35°C and 40°C before the onset of flight. In O. aygulus the thoracic temperature excess was as large as 19.3°C. The thermal threshold below which the frequency of flight onsets drops off rapidly is about 12°C for O. aygulus and 17°C for O. alexis and O. pecuarius. Radiant loss of body heat during cool nights and dawns may explain why smaller species (<0.4 g body weight), in particular, are adapted behaviourally so that they fly only during the day or early dusk.