Daily biphasic behaviour in keratin-feeding desert trogid beetles in relation to climate

Abstract. 1. Patterns of daily activity of two ectothermic species of the trogid genus Omorgus were studied in the Kalahari of southern Africa in summer. The relationship between time and type of activity, and environmental temperature, relative humidity and light intensity, was investigated.
2. Beetle activity was biphasic, with peaks in activity at sunset and sunrise in both species.
3. Afternoon activity coincided with high temperatures and low humidity and morning activity with minimum daily temperature and maximum humidity.
4. Afternoon activity, which peaked at about 34°C and 25% r.h., consisted predominantly of complex breeding behaviour and dispersal. High temperatures are necessary for these activities.
5. In the morning feeding predominated. At this time temperature was about 27°C and r.h. above 60%. High r.h. is important because it restricts respiratory loss of water, improves food (moist hair and skin keratin) quality and may compensate for faecal water loss.
6. No evidence of competition between the two species was found.     

Effect of temperature on circadian rhythm controlling the crepuscular activity of the burying beetle Nicrophorus quadripunctatus Kraatz (Coleoptera: Silphidae)

Locomotor activity rhythm was examined at various temperatures under a 16 h light : 8 h dark photoperiod (LD 16:8) or LD 12:12 using adults of the burying beetle Nicrophorus quadripunctatus. At 20°C, the locomotor activity of the beetles showed a bimodal daily pattern with two peaks around lights on and lights off under both photoperiods. This bimodal activity rhythm persisted under constant darkness; therefore, the activity of adult N. quadripunctatus is controlled by a circadian clock. Adults showed a bimodal activity pattern for temperatures ranging from 15 to 25°C. The evening peak of the daily activity was earlier at lower temperatures. These findings suggest that in the field, N. quadripunctatus shows crepuscular activity, and is active earlier in the afternoon in cooler seasons. In this species, therefore, temperature appears to play an important role in the determination of daily activity patterns.     

Thermoregulatory trade-offs result from vegetation removal by a harvester ant

Abstract.  1. Analyses of ecological trade-offs help to explain how organisms balance competing demands. Harvester ants ( Pogonomyrmex occidentalis ), are conspicuous residents of shortgrass prairie in western North America; worker P. occidentalis actively clear all vegetation from the immediate vicinity of their large gravel mounds. This study is based on the prediction that vegetation clearing yields a thermal trade-off by increasing soil temperatures; during cool periods the resulting increase in soil temperature opens new time windows for activity, while during hot periods the soil temperature is more likely to exceed the maximum thermal tolerance for this species. To test the hypothesis that daily and seasonal trade-offs in ant activity result from vegetation removal, the effects of experimentally applied shade on activity patterns were measured.
2. Harvester ant activity correlated highly with ground temperature; experimental shading of ant mounds shifted daily activity patterns by lowering ground temperature. Shading in the morning significantly delayed the onset of ant activity by preventing solar warming of the mound. In contrast, mid-day experimental shading prevented elevation of ground temperatures to above 50 °C and allowed ants to remain active when lethally high temperatures would normally force them inside the mound.
3. A model derived from field data predicted surface ground temperature (and therefore ant activity) based on air temperature and solar radiation, under conditions of sun and shade. For each of six seasons modelled, shade removal yielded a net gain of activity time. These results indicate that vegetation removal by harvester ants produces an advantageous thermoregulatory effect by helping to maximise activity time.     

Critical thermal limits in Mediterranean ant species: trade-off between mortality risk and foraging performance

1. In Mediterranean ant communities, a close relationship has been found between activity rhythm in the period of maximum activity and position in the dominance hierarchy: subordinate species are active during the day, when conditions are more severe, while dominants are active during the afternoon and the night.
2. Results obtained in this study confirmed that the species foraging at higher temperatures were closer to their critical thermal limits than the species foraging at lower temperatures.
3. This enabled two extreme strategies of foraging in relation to temperature to be distinguished: (1) heat-intolerant ant species behaved as risk-averse species, foraging at temperatures very far from their critical thermal limits; and (2) heat-tolerant ant species behaved as risk-prone species, foraging very near their critical thermal limits and running a high heat mortality risk.
4. Heat-tolerant species benefited from this strategy by having better foraging performance at high temperatures.
5. This wide range of thermal niches may be one reason why Mediterranean ant faunas are so diverse in the face of limited diversity in vegetation and habitat structure: the daily range of temperature may be sufficiently great to meet the requirement both of heat-adapted and cold-adapted species as well as a spectrum of intermediate forms.     

Body temperatures of Namib Desert tenebrionid beetles: their relationship in laboratory and field

Abstract The body temperatures of six apterous species of Namib Desert tenebrionid beetles were measured continuously with indwelling thermocouples under laboratory conditions and in the field. The range of body temperatures selected was within the upper half of their 'tolerated range', which we defined as the temperatures lying between measured critical thermal maximum and critical thermal minimum. In the field, individuals also maintained their body temperatures within the upper half of the 'tolerated range'. These beetles maintained higher body temperatures than those recorded for any other ectothermic insect. Three of the six species maintained lower body temperatures in the field than they selected in the laboratory. The other three species showed no significant difference between field and laboratory body temperatures. We conclude that these beetles are not forced by biotic or abiotic factors to adopt thermal niches which present them with physiological difficulties.     

Activity patterns, diets and interspecific interactions of coexisting spring and autumn breeding carabids: Carabusyaconinus and Leptocarabus kumagaii (Coleoptera, Carabidae)

Abstract. 1. Activity patterns, diets and interspecific interactions of two coexisting carabid beetles, Carabus yaconinus B. and Leptocarabus kumagaii K. et K., were studied in a lowland habitat in Japan.
2. Larvae of C. yaconinus appeared from May to October and fed on earthworms. Those of L. kumagaii appeared from October to the next May and fed on dipteran and lepidopteran larvae.
3. In the adult stages, feeding habits and daily activities of the two species were similar. The beetles fed mainly on earthworms and other animal materials, but various plant materials were also utilized.
4. Direct competition for food items was observed both intra- and interspecifically. Interspecific interactions occurred only in early summer, however, because of the difference in breeding seasons (i.e. spring for C. yaconinus and autumn for L. kumagaii).
5. The difference in seasonal life history patterns between these carabids is a possible mechanism of coexistence.     

The role of thermogenesis in the pollination biology of the Amazon waterlily Victoria amazonica

BACKGROUND AND AIMS: Several families of tropical plants have thermogenic flowers that show a 2-d protogynous sequence. Most are pollinated by large beetles that remain for the entire period in the flowers, where they compete for mates and feed. Active beetles require high body temperatures that they can achieve endogenously at great energy expense or attain passively and cheaply in a warm environment. Floral heating is therefore hypothesized to be a direct energy reward to endothermic beetles, in addition to its accepted role in enhancing scent production. METHODS: This study measures the pattern of floral heat production (as temperature in 20 flowers and respiration rates in five flowers) in Victoria amazonica at field sites in Guyana and correlates floral temperatures with body temperatures necessary for activity in visiting Cyclocephala hardyi beetles. KEY RESULTS: Thermogenesis occurred in a bimodal pattern, with peaks associated with the arrival and departure of beetles near sunset. Peak CO(2) production rates averaged 2.9 micromol s(-1), equivalent to a heat production of 1.4 W. Heat was generated mainly in the floral chamber on the first evening and by the stamen complex on the second. Mean chamber temperature remained between 29.3 and 34.7 degrees C during the first night, when ambient temperature was 23.5-25.2 degrees C. Beetles actively competed for mates and consumed stylar processes in the floral chamber, where their mean thoracic temperature was 33.2 degrees C. At the lower ambient temperatures outside of the flower, beetles capable of sustained flight had a similar mean temperature of 32.0 degrees C. CONCLUSIONS: Floral heating is not only associated with attraction, but continues throughout the night when beetles are active inside the flower and increases again when they leave. Floral chamber temperatures similar to activity temperatures of actively endothermic beetles imply that thermogenesis is an energy reward.     

Diel activity patterns of some field Carabidae

Abstract. 1. A 24 h time-sorting pitfall trap was used to collect Carabidae in a field in north-east England in 1974, 1976 and 1977.
2. Harpalus rufipes comprised 62% of the 5288 adult Carabidae caught. It was nocturnal, with an activity peak after midnight. The male activity pattern lagged behind that of females by about 1 h. The activity peak shifted to earlier in the night during September.
3. Activity curves are given for eighteen other common species. Large species were all nocturnal, but a third of the smaller species were active in the day. The numbers caught of twenty-eight less common species are tabulated and show the same trend. Overall, nearly 60% of all species caught were nocturnal and 20% diurnal; the remainder did not show a distinct pattern of day or night activity.
4. A total of eighty-two larvae of H.rufipes, Nebria brevicollis and Notiophilus biguttatus were caught, and showed similar activity patterns to those of the respective adult beetles.     

Seasonal changes of thermoregulatory efficiency in Djungarian hamsters

In their natural habitat, Djungarian hamsters are faced with dramatic seasonal changes. This requires various morphological and physiological adaptations allowing cope with harsh climate and food shortage, particularly in winter. These seasonal changes are controlled by the photoperiod and can be observed also in the laboratory at room temperature. The aim of the present study was to investigate if the efficiency of thermoregulation also depends on the photoperiod. For this reason, Djungarian hamsters were transferred to short-day conditions (SDC) with 8 h light and 16 h darkness. Two-thirds of the animals were classified as responders showing the typical seasonal changes – decrease of body mass, fur change, testes regression, vagina closing. The total activity per day did not change but, the nocturnal activity was spread over the longer dark time. The body temperature decreased, and the animals showed regular daily torpor. To investigate the thermoregulatory efficiency, body temperatures were correlated with motor activity. The obtained regression coefficients describe formally the effect of motor activity on body temperature, a measure for the efficiency of thermoregulation. In SDC, the coefficients were elevated, both during rest and activity, i.e. the same amount of activity did produce a larger increase in body temperature. Under field conditions, this might be an additional mechanism to compensate the bigger in winter heat loss. Also, the high coefficients may support the increase in body temperature at the end of a torpor phase by a bout of motor activity. The results show that, seasonal changes of thermoregulatory efficiency are an effective accessory way to cope with different temperatures in hamsters’ natural environment.     

Behavioral rhythms of the Japanese newts, Cynops pyrrhogaster, under a semi-natural condition

 Locomotor activity rhythms of the Japanese newt, Cynops pyrrhogaster, were recorded under a semi-natural condition using phototransistor systems. The daily activity rhythm showed a seasonal change: the locomotor activity was mainly diurnal (active during the daytime) from spring to early summer; mainly nocturnal (active during the night-time) from summer to autumn; and showed either a diurnal or nocturnal pattern, depending on the ambient temperature, in winter. To analyze the daily activity in detail, we observed the behavior of a group of newts (three males, three females) throughout 24 h. Four types of behavior (respiration, feeding, mating, and resting on the land) were observed. Each behavior had daily rhythms and showed a seasonal change. The behavior on land showed mainly a nocturnal or bimodal pattern (activity rhythms with two peaks) throughout the year and was more frequently observed in summer. Mating behavior also showed a seasonal change: high activity in spring, with peaks in the early morning and evening, but no activity in summer. Except in winter, feeding and respiratory behavior showed no seasonal changes in either activity period or frequency. Coupling between behavior and the clock seems to be weak in the Japanese newt because of indistinct daily rhythms and frequent phase changes of locomotor activity in water. Physical factors such as humidity and temperature seem to affect strongly the daily activity of the newts. Received: 21 April 1997 / Accepted: 1 September 1997     

基于红外相机技术的白鹇日活动节律研究

2014年5月至2016年4月,利用红外相机技术,系统调查和分析了浙江古田山国家级自然保护区内白鹇(Lophura nycthemera)的性比、集群方式和日活动节律。调查期间共获得244个有效位点的数据,累计58 890个工作日,收集到33 276份白鹇的照片和视频,有效探测5 687次,统计出雄性成体(含亚成体)3 946只次,雌性4 179只次,雌雄性比为1.06︰1。记录到群体大小从2至12只不等,群的组成方式多样。日活动节律分析的结果显示,白鹇的日活动时间段为5:00~18:00时,于6:00时、11:00时和16:00时有3个活动高峰,其间于10:00时和14:00时出现两个活动的低谷。雌雄个体的日活动节律基本一致,但雄性个体相比雌性清晨的活动高峰延长1 h(雌性6:00~8:00时,雄性6:00~9:00时),而午后的静息低谷推后1 h(雌性14:00时,雄性15:00时)。日活动节律密度函数分析结果显示,白鹇的日活动节律存在季节性变化。白鹇在春季和冬季存在早晚两个明显的活动高峰,而夏季和秋季在6:00~18:00时都较为活跃。同时,相比其他季节,白鹇在冬季早晨开始活动的时间推迟,傍晚结束活动的时间提前。不同季节的比较表明,白鹇在冬季和春、夏季的日活动节律重叠程度最低。白鹇日活动节律在低海拔带(200~700 m)和高海拔带(700~1 200 m)上存在差异,低海拔带个体在夏季和冬季的傍晚比高海拔带个体活跃。本研究结果为保护区内白鹇的保护管理提供了基础的科学依据。     

Humidity responses and water balance of riparian species of Bembidiini (Col., Carabidae)

Abstract. 1. The humidity responses of Asaphidion pallipes (Duftschmid) and Bembidion spp. were studied at various temperatures in choice chambers.
2. Several species initially showed a hygropositive or indifferent response. Other species had a hygronegative reaction which later changed to a hygropositive one. The duration of the hygronegative reaction varied with species, age, temperature and relative humidity. B.litorale (Olivier) and B.semipunctatum (Donovan) change response when they have lost 8–9% of their initial body weight as water.
3. The survival time of species of Bembidion under dry conditions was short compared with that reported for many other carabid beetles. Newly emerged adults of B.argenteolum Ahrens that had been kept away from sand for 3 weeks had lower transpiration rates than any other species. Young adult Bembidion had lower transpiration rates than those in the breeding phase, larvae and dead adults. Transpiration rate appeared to be inversely proportional to the relative humidity whereas there was a non-linear increase in weight loss with increasing temperature.
4. The species gain most water by drinking and/or from food.
5. There was no correlation between humidity response and transpiration rate or ability to tolerate dry conditions of the species. There was, however, a correlation between behaviour/habitat affinity and humidity response. Diurnal species, living in sandy or silty, half shady-exposed habitats had an initial hygronegative response, whereas those which hide among gravel/stones, leaves or vegetation were indifferent or hygropositive.     

ADAPTATION AND CONSTRAINT IN THE EVOLUTION OF THE PHYSIOLOGY AND BEHAVIOR OF THE NAMIB DESERT TENEBRIONID BEETLE GENUS ONYMACRIS

A comparative phylogenetic approach was used to test the following adaptive hypotheses pertaining to the physiological abilities of the Namib desert tenebrionid beetle genus Onymacris to withstand the hot, dry desert environment: (1) Desert-interior species evolved longer legs (relative to body size) than beetles in the cooler coastal region to facilitate stilting, i.e., elevating their bodies out of the hot boundary layer of air close to the substrate. (2) Wax blooms on the exoskeleton, which reduce evaporative water loss, are more likely to evolve in desert-interior species than in coastal species. (3) The high costs of activity in the extreme climates select for perfect coadaptation of preferred body temperatures (i.e., optimal temperatures for activity) and those they achieve in the field. All three of these adaptive hypotheses were supported by the results of squared-change parsimony and independent-contrasts analyses. Additionally, a parsimony approach suggested that a novel means of obtaining water from periodic fogs, known as fog basking, has evolved independently on two occasions.     

Field studies of the temporal response patterns of male Diabrotica virgifera virgifera LeConte and D. barberi Smith and Lawrence to 8R-methyl-2R-decyl propanoate

ABSTRACT. Variations in the diel response of male northern and western corn rootworms attracted to a synthetic pheromone source were examined in southwestern Ontario. Activity was measured by the number of males caught at hourly intervals in sticky traps baited with 8 R -methyl-2 R -decyl propanoate. Northern corn rootworm males were attracted to the pheromone between 22.30 and 0330 hours and were unresponsive at all other times at temperatures above 15°C. Western corn rootworm males had a bimodal response with peaks in attraction from 08.30 to 12.30 hours and again from 15.30 to 18.30 hours. While the period of peak attraction for northern corn rootworm males was well defined, the pattern for western corn rootworm males was erratic and many beetles were caught throughout the day. Evening temperatures below 15°C suppressed the activity of both northern and western corn rootworms. While the nightly period of activity for the northern corn rootworms was shifted by low temperatures to the following morning, that of the western corn rootworm was not altered substantially. As a result, peaks in the response of northern and western corn rootworm males occurred simultaneously under these conditions, suggesting strongly that temporal separation of activity periods was not an important factor in maintaining sexual isolation for these species in Ontario.     

Movements of marsh-dwelling invertebrates

SUMMARY. 1. Invertebrate movement was monitored in a parkland marsh in central Alberta, Canada, for 24 h periods every 2 weeks during the ice-free season. Stationary plankton nets captured organisms moving towards or away from shore.
2. Movement occurred throughout the sampling programme, particularly during the warm summer months. Most organisms were night active and exhibited directional movements. Hyalella azteca (Saussure) and Caenis simulans McDunnough constituted most of the catch.
3. Although there was a significant positive seasonal relationship between maximum water temperature and total numbers of animals captured. no physical factor appeared to control movement on a daily basis.     

The effect of a seasonal time constraint on development time, body size, condition, and morph determination in the horned beetle Allomyrina dichotoma L. (Coleoptera: Scarabaeidae)

Abstract.  1. In horned beetles selection favours males that adjust their investment in horn development in relation to cues that predict adult body size. Here it is shown that in the Japanese horned beetle, Allomyrina dichotoma . There is a significant discontinuity in the horn length body size allometry. This can be described as a linear relationship that is shifted towards an increased horn length to body length ratio in males with horns longer than 16 mm.
2. Larval nutrition explains morph determination in A. dichotoma . However, unlike other species, variation in larval nutrition was the result of a seasonal time constraint that limits the time available for feeding prior to the onset of winter diapause.
3. Even when eggs were reared with an ad libitum food supply, minor morphs were still observed. Individuals that were oviposited later in the season had less time to feed, shorter development times, eclosed as smaller individuals and, in the case of males, were more likely to be hornless. Major morphs, minor morphs, and females all reduced their body size in response to seasonal time constraints in the same way. However, males that were laid later in the season had faster development times than females laid at the same time, but showed no reduction in their size relative to females, suggesting seasonal time constraints increase growth rates in males but not in females.
4. No evidence was found that seasonal time constraints resulted in a reduction of size-corrected fat reserves at eclosion, or that minor morphs gained any developmental advantage by reducing investment in horn length.     

Responses to desiccation in four Coleopterans from sub-Antarctic South Georgia

RATES OF WATER LOSS WERE DETERMINED FOR FOUR COLEOPTERA SPECIES: the herbivores Hydromedion sparsutum, Perimylops antarcticus (Family Perimylopidae) and the carnivores Trechisibus antarcticus, Oopterus soledadinus (Family Carabidae) collected during summer from a range of terrestrial habitats at South Georgia. A recording microbalance enabled measurement of individual weight loss with time in <5% r.h. at 10, 20, 30 and 35 degrees C. Adults of T. antarcticus had significantly higher rates of water loss than any other species over all temperatures. Individuals of both herbivores exhibited the slowest water loss rates under the experimental conditions. Within species, rates at 10 or 20 degrees C were slower than at the higher temperatures. Adult P. antarcticus had significantly greater amounts of body water than adult H. sparsutum for each of the four temperatures. Within species and life-stages of both herbivores, body water contents after drying at 10 degrees C were significantly lower than individuals dried at 30-35 degrees C, but no such differences were observed for the carnivores. At each temperature, rates of water loss were negatively correlated with initial live weight in all four species, but this was not the case within species or between adults and larvae. Maximum survival times during desiccation declined as temperature increased, but did not differ between species at 10 degrees C. Over 30-35 degrees C, survival times of both herbivores were significantly longer than either of the carnivores. Smaller insects (e.g. the carabids) had faster rates of water loss than the larger perimylopids under the same environmental conditions. The latter had greater resistance to desiccation than the former. It is suggested that the larger body water content of P. antarcticus enables it to resist desiccation more than the other three species, which correlates with its ecological distribution. Differences in water contents after drying individuals at low and high temperatures may be caused either by the water binding properties of cells and tissues or by reduction in energy stores in order to maintain metabolism at lower environmental temperatures causing a body weight loss. Whilst both herbivores show some physiological adaptations to drying conditions, it is suggested that the two carnivorous beetles may have adapted behaviourally to the South Georgia environment.     

Partitioning variance in a physiological trait: desiccation resistance in keratin beetles (Coleoptera, Trogidae)

1. Although variation in physiological traits forms the substance of evolutionary physiology, the way that this variation is partitioned among different hierarchical levels (e.g. population, species, genus) is not well known.
2. In this study variance partitioning is examined in body size, water content, lipid content, rate of water loss, maximum tolerable water loss and survival time at four levels (individual, population, species, genus) in southern African keratin beetles.
3. It is found that most variance in body size, and the physiological traits that are strongly influenced by body size (water and lipid content, maximum tolerable water loss, rate of water loss), is partitioned at the generic level (50–70%), then at the species level (20–50%) and finally at the population (1–9%) and individual (2–18%) levels.
4. On the other hand, variance in survival time, and variance in rate of maximum water loss once the effects of body size have been taken into account, are partitioned mostly at the species level (40–70%), whereas maximum tolerable water loss and lipid and water content show greatest variance at the individual level (63–75%). This is largely a consequence of differences in the extent of scaling of these traits.
5. The results suggest that where possible the effects of phylogeny should be controlled for when examining desiccation resistance in insects using comparative methods, but that confidence in conclusions from previous studies, which have used mass-specific data, is not unwarranted.     

The utilization of patchy thermal microhabitats by the ectothermic insect predator, Cicindela sexguttata

Abstract. 1. Tiger beetles ( Cicindela ) of open habitats have served as model ectotherms in studies of the dependence of activity and habitat utilization on temperature. Potential departures from the cicindelid model were investigated in Cicindela sexguttata , a species inhabiting forests where thermal resources are patchy and ephemeral.
2. Body temperatures (T_b) were determined by inserting thermocouples into beetles immediately after observing specific behaviours in the field. Cicindela sexguttata elevated T_b by basking, foraged at a preferred T_b of 32.8 °C, and stilted, sun-faced, or sought shade when T_b exceeded 35 °C. Although these behaviours were typical of Cicindela , their set points were lower than those of species in more open habitats.
3. Illuminated substrates were utilized for basking and foraging. Beetles were dispersed throughout the forest floor in early spring, but became aggregated in light gaps when the canopy leafed out. Operative temperatures of thermal models indicated that beetles were unable to maintain the preferred T_b in shade and would not be able to maintain a preferred T_b in light gaps during the autumn, when adults are normally in diapause.
4. Beetles were confined to foraging in light gaps once the canopy was closed. Foraging rate and searching speed were independent of T_b, in contrast to other cicindelids. Adults rarely searched for prey, but ambushed small arthropods that alighted in the light gap. Dependence on patches of illumination as thermal resources may increase prey capture, intraspecific encounters, and risk of predation, and preclude foraging in the autumn when Cicindela species of open habitats are also active.     

Effects of foraging mode and season on the energetics of the Marine Iguana, Amblyrhynchus cristatus

1. Marine Iguanas ( Amblyrhynchus cristatus ) inhabiting the rocky shores of the Galápagos Islands apply two foraging strategies, intertidal and subtidal foraging, in a seasonal climate. Effects of both foraging strategy and seasonality on the daily energy expenditure (DEE) were measured using doubly labelled water.
2. Difference in foraging mode did not result in significant differences in DEE.
3. On Santa Fé the DEE in the warm season was significantly higher than in the cool season (67·8 ± 21·8 kJ kg^–0·8 day^–1 vs 38·0 kJ kg^–0·8 day^–1). This difference can be explained by body temperature. A model estimate of the body temperature was used to predict monthly DEE figures, giving a year round budget. On average a 1-kg iguana would need only 47 kJ day^–1, or 17 mJ year ^–1. This is lower than previous estimates in which body temperatures were not taken into account.
4. The water flux of the Marine Iguana increases with increasing foraging time. The linear rise per minute foraging is roughly two times as high for subtidally foraging animals as for intertidal foragers.