Thermoregulation in endothermic dung beetles (Coleoptera: Scarabaeidae): effect of body size and ecophysiological constraints in flight

We explore the physiological constraints of body temperature as related to body mass and ambient temperature during flight in endothermic dung beetles showing a mass-related breakpoint where species show strong vs. weak endothermy. We found two different strategies in the dung beetles prior to flight; larger beetles (>1.9 g) elevate and maintain their body temperature (T(b)) at levels well above ambient temperature (T(a)) whereas smaller beetles' (<1.9 g) T(b) tends to conform with T(a). Physiological constraints analysis revealed a constant maximum tolerated temperature (in flight) of 42 degrees C and a minimum temperature for flight of around 25 degrees C. These, with body mass, may play a role in thermal niche partitioning and geographical distribution patterns.     

Thermal investigations of a honey bee colony: thermoregulation of the hive during summer and winter and heat production of members of different bee castes

Summary The temperature at the centre, the periphery and the entrance of a honey bee colony (Apis mellifera carnica) was continuously determined during the summer season and the broodless time in winter. During the summer season the temperature in the brood nest averages 35.5°C with brief excursions up to 37.0°C and down to 33.8°C. Increasing environmental temperatures resulted in linear increases in the temperature of the hive entrance, its periphery and its centre. The temperature in the centre of an overwintering cluster is maintained at an average value of 21.3°C (min 12.0°C, max 33.5°C). With rising ambient temperatures the central temperature of a winter cluster drops whereas the peripheral temperature increases slightly. With decreasing external temperatures the peripheral temperature is lowered by a small amount while the cluster's centre temperature is raised. Linear relationships are observed between the central and the ambient temperature and between the central temperature and the temperature difference of the peripheral and the ambient temperatures. The slopes point to two minimum threshold values for the central (15°C) and the peripheral temperature (5°C) which should not be transgressed in an overwintering cluster. Microcalorimetric determinations of the heat production were performed on the three castes of the honey bee: workers, drones and queens of different ages. Among these groups single adult workers showed the highest heat production rates (209 mW·g^–1) with only neglectible fluctuations in the heat production rate. Juvenile workers exhibited a mean heat production rate of 142 mW·g^–1. The rate of heat production of adult workers is strongly dependent upon the number of bees together in a group. With more than 10 individuals weight-specific heat dissipation remains constant with increasing group sizes at a level approximately 1/17 that of an isolated bee. Differences are seen between the rates of virgin (117 mW·g^–1) and laying (102 mW·g^–1) queens. Laying queens showed less thermal fluctuations than virgin queens. High fluctuations in heat production rates are observed for drones. In both groups (fertile, juvenile) phases of high and extremely low activity succeed one another. The heat production of juvenile drones was 68 mW·g^–1, that of fertile drones 184 mW·g^–1 due to stronger locomotory activities.     

Thermoregulation in leopard tortoises in the Nama-Karoo: The importance of behaviour and core body temperatures

Despite being ectotherms, reptiles have an ability to thermoregulate which is enhanced by adopting a variety of behavioural mechanisms. Different behavioural postures, the use of retreat sites and selection of microhabitats enable reptiles to maintain their core body temperatures (T_b) above that of ambient temperatures (T_a) in winter or below T_a maximum in summer. This study describes the daily activity patterns of leopard tortoises (Stigmochelys pardalis) in relation to T_b and T_a, and the extent to which leopard tortoises can manipulate their T_b in response to seasonal changes in T_a. Ten and nine leopard tortoises were radio-tracked in 2002 and 2003, respectively and cloacal T_b and behaviours observed. Core T_bs were measured using Thermocron iButtons^™ surgically implanted into the body cavities of 4 and 5 adult telemetered tortoises for summer and winter 2003, respectively. There were seasonal differences in the extent to which certain behaviours were practiced and the time of day that these occurred. Leopard tortoises generally had unimodal activity patterns in winter and bimodal ones in summer. In winter tortoises were active at lower T_bs, and at lower T_a, than in summer. Tortoises maintained their core T_b well below T_a minimum profiles in summer and well above these in winter. Core T_b closely followed the increase in T_a minimum profiles in the mornings, however tortoises exhibited an extended thermal lag when T_a minimum profiles cooled overnight. By using different behavioural mechanisms in summer and winter, leopard tortoises maintained their core T_b at different levels compared with T_a minimum and maximum profiles. Consequently although they are ectotherms, they maintained their core T_b independent of T_a.     

Influence of environmental and colony factors on the initial commodity choice of foragers of the stingless bee Plebeia tobagoensis (Hymenoptera, Meliponini)

Novice foragers of social bees have to decide what food commodity to collect when they start foraging for the first time. In this decision making process two types of factors are involved: internal factors (the response threshold) and external factors (environmental and colony conditions). In this study we will focus on the importance of two external factors, pollen storage level and information from experienced foragers about food availability in the field, on the initial commodity choice of foragers of the stingless bee species Plebeia tobagoensis. We also studied the effect of the initial choice of individuals on their subsequent foraging career. This study was performed in a closed greenhouse compartment, where food availability and colony condition could be controlled. Information on food availability in the field from experienced foragers and pollen storage level both greatly influenced the initial commodity choice of individuals, with more choices for the commodity communicated by experienced foragers or lacking in storage. The initial choice of foragers is of importance for their future foraging career, although a substantial proportion of foragers did switch between food commodities. Because of the ability of novice foragers to become flexibly distributed over foraging tasks, social bees are able to react to changes in their environment without directly having to decrease foraging effort devoted to other foraging tasks. This, in combination with individual flexibility during foraging careers makes it possible for colonies of P. tobagoensis to forage efficiently in an ever-changing environment. Received 7 November 2005; revised 12 January 2006; accepted 16 February 2006.     

Allometry of head and body size in Holocene foragers of the South African Cape

Opportunities to assess morphological allometry in small-bodied human populations are rare. The foragers of the Later Stone Age of the South African Cape are characteristically small-bodied. Previous studies have shown that during the period of ca. 3500 to 2000 years BP (uncalibrated (14) C dates), the regional population shows transient reduced stature, body mass, and cranial size, a pattern that has been tentatively tied to demographic pressure on resources. This study examines the relationships among cranial size (centroid size) and body size (femoral length, femoral head diameter, and bi-iliac breadth) during the second half of the Holocene (N = 62). Reduced major axis regression indicates negative allometry of cranial centroid size with body size. Residuals (from ordinary least squares regression of cranial centroid size on body size) are regressed on radiocarbon date to examine temporal changes in the relationship between cranial and body size. Cranial and pelvic sizes are most conserved through time, while more ancient skeletons possess shorter femora and smaller femoral heads. The relationship between cranial centroid size and femoral length shows larger and more variable residuals at more recent dates, indicating a greater or more variable disassociation between cranial size and stature relative to more ancient skeletons. A similar, but nonsignificant relationship exists between cranial size and bi-iliac breadth. These results provide insights into the use of aspects of body size and proportionality in the assessment of health in past populations.     

Effect of group size on the aggression strategy of an extirpating stingless bee, Trigona spinipes

Summary. Group aggression influences communication and defense strategies in many social insect communities. Such aggression plays a particularly significant role in the lives of stingless bees, important native Neotropical pollinators, in which the battle for food resources can be deadly and critical to colony survival. However, the effects of group size on individual aggression levels and the spatio-temporal aggression strategy of communal aggressors have not been fully explored. We therefore investigated how group size affects the aggression levels and the spatio-temporal attack strategy (which body parts, and the amount of time spent in attacking each part) in close combats between Trigona spinipes foragers and a natural competitor, Melipona rufiventris. In all trials, T. spinipes foragers competitively excluded M. rufiventris foragers from nearby feeders, exhibiting four levels of aggressive behavior ranging from threat displays to prolonged grappling and decapitation. Surprisingly, aggression levels and spatial strategy corresponded to the size of group attacks. Larger groups of attackers used individually lower aggression levels than small groups of attackers. Smaller groups also attacked appendages linked to escape (legs and wings) with greater frequency than larger groups, which focused on vital central body areas (abdomen, thorax, and head). Increased aggression corresponded to increased risks for attackers and the attacked. All combatants engaging at the highest level of aggression died (100% mortality). Thus the dominance style of T. spinipes may minimize attack risk and maximize victim harm with finely tuned hostility.Received 22 May 2004; revised 18 August 2004; accepted 26 October 2004.     

Microsatellite loci for the stingless bee Melipona rufiventris (Hymenoptera: Apidae)

Eight microsatellite primers were developed from ISSR (intersimple sequence repeats) markers for the stingless bee Melipona rufiventris. These primers were tested in 20 M. rufiventris workers, representing a single population from Minas Gerais state. The number of alleles per locus ranged from 2 to 5 (mean = 2.63) and the observed and expected heterozygosity values ranged from 0.00 to 0.44 (mean = 0.20) and from 0.05 to 0.68 (mean = 0.31), respectively. Several loci were also polymorphic in M. quadrifasciata, M. bicolor, M. mandacaia and Partamona helleri and should prove useful in population studies of other stingless bees.     

Field data confirm the ability of a biophysical model to predict wild primate body temperature

In the face of climate change there is an urgent need to understand how animal performance is affected by environmental conditions. Biophysical models that use principles of heat and mass transfer can be used to explore how an animal's morphology, physiology, and behavior interact with its environment in terms of energy, mass and water balances to affect fitness and performance. We used Niche Mapper™ (NM) to build a vervet monkey (Chlorocebus pygerythrus) biophysical model and tested the model's ability to predict core body temperature (T_b) variation and thermal stress against T_b and behavioral data collected from wild vervets in South Africa. The mean observed T_b in both males and females was within 0.5 °C of NM's predicted T_bs for 91% of hours over the five-year study period. This is the first time that NM's T_b predictions have been validated against field data from a wild endotherm. Overall, these results provide confidence that NM can accurately predict thermal stress and can be used to provide insight into the thermoregulatory consequences of morphological (e.g., body size, shape, fur depth), physiological (e.g. T_b plasticity) and behavioral (e.g., huddling, resting, shade seeking) adaptations. Such an approach allows users to test hypotheses about how animals adapt to thermoregulatory challenges and make informed predictions about potential responses to environmental change such as climate change or habitat conversion. Importantly, NM's animal submodel is a general model that can be adapted to other species, requiring only basic information on an animal's morphology, physiology and behavior.     

Implications of variation in worker body size for the honey bee recruitment system

Honey bee scouts seek food from flowers, return to the colony, and may perform a dance used to recruit dance followers to the flowers. Variation in body size of workers may result in the communication of misinformation because some information acquired by the scout and signaled to recruits is affected by body size. I tested two predictions of this hypothesis. (1) Recruitment communication takes place between bees of similar size despite the withincolony size distribution. (2) There is an inverse relationship between the size variation of foraging honey bees (Apis mellifera)and the rate at which nectar is returned to the colony. A positive relationship was found between the size of a dancer and that of its dance followers, which together comprise a dance group. There was less variation in size within dance groups than among groups. These two factors effectively lower the difference in size between signal sender and signal receiver and may enhance the flow of accurate size-dependent information. Also, an inverse relationship between size variation and rate of nectar intake was detected in each of six colonies using partial correlation analysis. This may be due to communication of misinformation when size variance is higher. The relationship was statistically significant in two colonies and the combined results were significant. The results of the first study suggest the generally weak relationships found in this second study.     

Seasonal,spatial and diel partitioning of Acyrthosiphon pisum (Hemiptera: Aphididae) predators and predation in alfalfa fields

Predators are important natural enemies, often responsible for preventing pest population outbreaks of in many crops. Complementarity in resource use involves spatial or temporal segregation of predators, which can result in better biological control when several species of natural enemies share a prey. In this study, we investigated the seasonal, spatial and diel segregation of Acyrthosiphon pisum predators and its predation in alfalfa fields, by setting out cards with sentinel aphids, and making observations every 3 h for a 24 h period. A temporal and spatial segregation of predators was observed. Coccinellids were the most abundant predators, representing 51% of the total observations, followed by syrphid larvae. Coccinellids were also responsible for high levels of predation throughout the year, although the species responsible varied from spring to summer and autumn. On the other hand, syrphids were only found in spring and summer, while spiders only in autumn. Predator species also differed on their preferred sites for predation, with Heteropterans and syrphids found on the foliage, the spider Neomaso articeps only on the ground, and coccinellid and Anyphaenidae species on both sites. The two main predator groups also showed distinct diel patterns, with coccinellids observed only during day and syrphids only during night. This predatory activity corresponded with aphid predation, observing more predation in spring, on the foliage and during the day time. The proportion of predators observed preying on cards in the different seasons did not corresponded tightly with their field abundance, particularly in the case of coccinellids, which maintained high levels of predation in spite of great variations in its field abundance. Our results support the hypothesis of a spatio-temporal segregation of the predators associated with A. pisum in alfalfa, which might be beneficial for the outcome of biological control of this pest.     

The evolution of body armor in mammals: plantigrade constraints of large body size

Predictions associated with opposing selection generating minimum variance in basal metabolic rate (BMR) in mammals at a constrained body mass (CBM; 358 g) were tested. The CBM is presumed to be associated with energetic constraints linked to predation and variable resources at intermediate sizes on a logarithmic mass scale. Opposing selection is thought to occur in response to energetic constraints associated with predation and unpredictable resources. As body size approaches and exceeds the CBM, mammals face increasing risks of predation and daily energy requirements. Fast running speeds may require high BMRs, but unpredictable and low resources may select for low BMRs, which also reduce foraging time and distances and thus predation risks. If these two selection forces oppose each other persistently, minimum BMR variance may result. However, extreme BMR outliers at and close to the CBM should be indicative of unbalanced selection and predator avoidance alternatives (escapers vs. defenders), and may therefore provide indirect support for opposing selection. It was confirmed that body armor in defenders evolves at and above the CBM, and armored mammals had significantly lower BMRs than their nonarmored counterparts. However, analyses comparing the BMR of escapers--the fastest nonarmored runners (Lagomorpha)--with similar-sized counterparts were inconclusive and were confounded by limb morphology associated with speed optimization. These analyses suggest that the risks and costs of predation and the speed limitations of the plantigrade foot may constrain the evolution of large body sizes in plantigrade mammals.     

Temporal partitioning and aggression among foragers: modeling the effects of stochasticity and individual state

In many natural systems, individuals compete with conspecificsand heterospecifics for food and in some cases, individualshave been observed to partition their foraging times or fightover food. In this study, I investigated when it is optimalfor a consumer to partition time and be aggressive. I formulatedan individual-based model of foraging and used game theoryto find evolutionarily stable strategies (ESSs) that maximizethe probability that consumers survive each day and acquiretheir daily food requirements. Consumers choose when to forageand when to behave aggressively during confrontations overfood. Consumers are each associated with a state variable,representing the amount of food eaten, and a dominance ranking, which describes how likely they are to forage and fight forfood. The ESS is sensitive to food abundance, consumer state,and the dominance ranking. When food is abundant, temporalpartitioning is often an ESS where the dominant consumer foragesfirst; however, partitioning is unlikely to be an ESS when food abundance is low. Fights over food are typically avoidedbut may be part of an ESS when food abundance is low, bothconsumers are hungry, or the time available for foraging eachday is drawing to a close. Because the ESS is sensitive toconsumer state, the stochastic nature of finding food often results in considerable variation in observed foraging dynamicsfrom one day to the next, even when consumers adopt the samestate-dependent strategy each day. Results are compared withempirical observations, and I discuss implications for consumercoexistence.     

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.     

Niche differentiation depends on body size in a cichlid fish: a model system of a community structured according to size regularities

1. Communities of different species are often structured according to niche differentiation associated with competitive interactions. We show that similar principles may apply on an ecological time-scale when individuals of a species having a wide size variation compete for resources, using the Lake Tanganyika cichlid Lobochilotes labiatus (5-30 cm). This species has a mouth especially adapted to suck up invertebrates from rock crevices. 2. Individuals defended feeding territories against similar-sized conspecifics, but not against different-sized ones. Thus, territories of similar-sized fish rarely overlapped, but up to a total of seven individuals (of seven size-ranks) had broadly overlapping territories with dissimilar-sized individuals. Comparison with expectation from the null model demonstrated clearly that observed size ratios between adjacent size rank were determined non-randomly regardless of sexual combinations. 3. Larger individuals took larger prey types of larger average size, but more importantly used wider rock crevices from which to suck food than smaller individuals. We calculated pairwise values of Schoener's index of diet overlap C(d) and the values of Levin's index of diet breadth B(d) (prey type and prey size) and the same for the width of the rock crevices used for foraging (C(r) and B(r)). C(d) remained high among all combinations of the seven ranks. In contrast, C(r) declined strongly in combinations of adjacent ranks (to 0.27), and was low or zero among further different size ranks. This shows that fish with overlapping territories divided the food resources largely through foraging site partitioning. Accordingly, B(d) did not depend on the size difference to the nearest two coinhabiting fish, whereas B(r) did. 4. We conclude that this L. labiatus community is structured non-randomly: body size-dependent effects on foraging site usage result in competition with, and territorial exclusion of, similar-sized individuals, but not of dissimilar-sized individuals that were accepted as coinhabitants. Accordingly, mean body size ratios (large/small) between two adjacent ranks were consistently approximately 1.28 [standard deviation (SD) = 0.07, n = 104], while approximately 1.34 from the null model (SD = 0.34, n = 10 400 simulations). We discuss our results as an example of Hutchinson's rule, applied originally to size ratios of different species.     

Food webs: Ordering species according to body size yields high degree of intervality

Food webs, the networks describing “who eats whom” in an ecosystem, are nearly interval, i.e. there is a way to order the species so that almost all the resources of each consumer are adjacent in the ordering. This feature has important consequences, as it means that the structure of food webs can be described using a single (or few) species' traits. Moreover, exploiting the quasi-intervality found in empirical webs can help build better models for food web structure. Here we investigate which species trait is a good proxy for ordering the species to produce quasi-interval orderings. We find that body size produces a significant degree of intervality in almost all food webs analyzed, although it does not match the maximum intervality for the networks. There is also a great variability between webs. Other orderings based on trophic levels produce a lower level of intervality. Finally, we extend the concept of intervality from predator-centered (in which resources are in intervals) to prey-centered (in which consumers are in intervals). In this case as well we find that body size yields a significant, but not maximal, level of intervality. These results show that body size is an important, although not perfect, trait that shapes species interactions in food webs. This has important implications for the formulation of simple models used to construct realistic representations of food webs.     

Some like it hot: body and weapon size affect thermoregulation in horned beetles

Body size and shape affect thermoregulatory properties of organisms, and in turn are believed to have shaped macroevolutionary patterns of morphological diversity across many taxa. However, it is less clear whether thermoregulation plays a role in shaping intraspecific morphological diversity such as sexual dimorphisms or the conditional expression of exaggerated secondary sexual traits. Here, we investigate individual thermoregulatory properties in two species of horned beetles that share similar ecologies and body size ranges, but differ substantially in degree of sexual and male dimorphism. We find that intraspecific variation in body size had an unexpectedly large effect on thermal preference behavior and the ability to passively regulate body temperature. Furthermore, we find that the presence or absence of exaggerated secondary sexual traits dramatically altered thermal preference behavior, consistent with a thermoregulatory cost of horn possession. Lastly, we show that the increase in surface area associated with the expression of enlarged horns is, by itself, insufficient to account for the radically altered thermoregulatory behavior observed in horn-bearing males, and discuss possible alternative, physiological explanations. These findings are among the first to link intra-and interspecific variation in body- and weapon size to thermal preferences within and between insect species.     

Chemical basis for inter-colonial aggression in the stingless bee Scaptotrigona bipunctata (Hymenoptera: Apidae)

Inter-colonial aggression was tested using three colonies of Scaptotrigona bipunctata in a natural setting when their nests were moved and by artificial contact between individuals. Examination of the cuticular lipids of individuals from two colonies kept under identical conditions showed clear differences in their cuticular hydrocarbon profiles. The cuticular lipids were a mixture of hydrocarbons (saturated and unsaturated alkanes and alkenes) within the range of C23-C29. The use of multivariate analysis (PCA and discriminant analysis) showed that seven of the identified surface compounds are enough to separate workers from colonies A and B from each other.     

Response of avian nectarivores to the flowering of Aloe marlothii : a nectar oasis during dry South African winters

In southern Africa, Aloe marlothii flowers during the dry winter season and offers copious dilute nectar to a variety of birds. Avian abundance and community composition were monitored at an A. marlothii forest at Suikerbosrand Nature Reserve, South Africa. Sampling occurred during two summer months (February–March) when no flowers were present, and six months (May–October) that spanned the winter flowering. We hypothesized that an influx of occasional nectarivores to the A. marlothii forest during flowering would lead to significant changes in the avian community. Overall bird abundance increased 2–3 fold at the peak of nectar availability (August). We recorded 38 bird species, of 83 species detected during transects, feeding on A. marlothii nectar; this diverse assemblage of birds belonged to 19 families, including Lybiidae, Coliidae, Pycnonotidae, Sylviidae, Cisticolidae, Muscicapidae, Sturnidae, Ploceidae and Fringillidae. Surprisingly, only two species of sunbird (Nectariniidae) were observed feeding on A. marlothii nectar, and both occurred in low abundance. We predicted that competition for nectar resources would be high, but few aggressive inter- and intra-specific interactions occurred between birds while feeding on inflorescences. During peak flowering, insect feeders (insectivores, omnivores, nectarivores) fed on nectar during the cold morning when insect activity was low, whilst non-insect feeders (frugivores and granivores) fed on nectar in the middle of the day. Our study highlights the importance of A. marlothii nectar as a seasonal food and water source for a diverse assemblage of occasional nectarivores.