Interspecific Studies Research Articles

Ontogenetic Scaling of Fore- and Hind Limb Posture in Wild Chacma Baboons (Papio hamadryas ursinus)

Large-scale interspecific studies of mammals ranging between 0.04–280 kg have shown that larger animals walk with more extended limb joints. Within a taxon or clade, however, the relationship between body size and joint posture is less straightforward. Factors that may affect the lack of congruence between broad and narrow phylogenetic analyses of limb kinematics include limited sampling of (1) ranges of body size, and/or (2) numbers of individuals. Unfortunately, both issues are inherent in laboratory-based or zoo locomotion research. In this study, we examined the relationship between body mass and elbow and knee joint angles (our proxies of fore- and hind limb posture, respectively) in a cross-sectional ontogenetic sample of wild chacma baboons (Papio hamadryas ursinus) habituated in the De Hoop Nature Reserve, South Africa. Videos were obtained from 33 individuals of known age (12 to ≥108 months) and body mass (2–29.5 kg) during walking trials. Results show that older, heavier baboons walk with significantly more extended knee joints but not elbow joints. This pattern is consistent when examining only males, but not within the female sample. Heavier, older baboons also display significantly less variation in their hind limb posture compared to lighter, young animals. Thus, within this ontogenetic sample of a single primate species spanning an order of magnitude in body mass, hind limb posture exhibited a postural scaling phenomenon while the forelimbs did not. These findings may further help explain 1) why younger mammals (including baboons) tend to have relatively stronger bones than adults, and 2) why humeri appear relatively weaker than femora (in at least baboons). Finally, this study demonstrates how field-acquired kinematics can help answer fundamental biomechanical questions usually addressed only in animal gait laboratories.

Molecular Authentication of Morus Folium Using Mitochondrial nad7 Intron 2 Region

Morus Folium (Sang-yeop in Korean) is one of the most important Oriental medicinal plants. In Korea, both M. alba and M. cathayana are regarded as the botanical sources for Morus Folium. In order to discriminate M. alba and M. cathayana from their adulterant, M. tricuspidata, mitochondrial NADH dehydrogenase subunit 7 (nad7) intron 2 region was targeted for molecular analysis with universal primers. DNA polymorphisms, including SNP sites, insertions, and deletions, were detected among these three species sequencing data. Based on these DNA polymorphisms, specific primers were designed for the three species respectively. Multiplex PCR was conducted for molecular authentication of M. alba, M. cathayana, and M. tricuspidata with specific primers. The present results indicate that it is possible to identify Morus Folium from its adulterant using mitochondrial nad7 intron 2 region. The established multiplex-PCR system was proved to be effective for identification of Morus Folium. The results indicate that mitochondrial introns can be used for inter-specific polymorphic study, and the described method can be applied for molecular identification of medicinal materials.

Evolutionary ecology of intraspecific brain size variation: a review.

The brain is a trait of central importance for organismal performance and fitness. To date, evolutionary studies of brain size variation have mainly utilized comparative methods applied at the level of species or higher taxa. However, these studies suffer from the difficulty of separating causality from correlation. In the other extreme, studies of brain plasticity have focused mainly on within-population patterns. Between these extremes lie interpopulational studies, focusing on brain size variation among populations of the same species that occupy different habitats or selective regimes. These studies form a rapidly growing field of investigations which can help us to understand brain evolution by providing a test bed for ideas born out of interspecific studies, as well as aid in uncovering the relative importance of genetic and environmental factors shaping variation in brain size and architecture. Aside from providing the first in depth review of published intraspecific studies of brain size variation, we discuss the prospects embedded with interpopulational studies of brain size variation. In particular, the following topics are identified as deserving further attention: (i) studies focusing on disentangling the contributions of genes, environment, and their interactions on brain variation within and among populations, (ii) studies applying quantitative genetic tools to evaluate the relative importance of genetic and environmental factors on brain features at different ontogenetic stages, (iii) apart from utilizing simple gross estimates of brain size, future studies could benefit from use of neuroanatomical, neurohistological, and/or molecular methods in characterizing variation in brain size and architecture.Evolution of brain size and architecture is a widely studied topic. However, the majority of studies are interspecific and comparative. Here we summarize the recently growing body of intraspecific studies based on population comparisons and outline the future potential in this approach.

Space-use scaling and home range overlap in primates.

Space use is an important aspect of animal ecology, yet our understanding is limited by a lack of synthesis between interspecific and intraspecific studies. We present analyses of a dataset of 286 estimates of home range overlap from 100 primate species, with comparable samples for other space-use traits. To the best of our knowledge, this represents the first multispecies study using overlap data estimated directly from field observations. We find that space-use traits in primates are only weakly related to body mass, reflecting their largely arboreal habits. Our results confirm a theory that home range overlap explains the differences in allometric scaling between population density and home range size. We then test a suite of hypotheses to explain home range overlap, both among and within species. We find that overlap is highest for larger-bodied species living in large home ranges at high population densities, where annual rainfall is low, and is higher for arboreal than terrestrial species. Most of these results are consistent with the economics of resource defence, although the predictions of one specific theory of home range overlap are not supported. We conclude that home range overlap is somewhat predictable, but the theoretical basis of animal space use remains patchy.

Pheomelanin-Based Plumage Coloration Predicts Survival Rates in Birds

Higher vertebrates synthesize two forms of melanin: eumelanin and pheomelanin. While the adaptive functions of eumelanin are diverse, those of pheomelanin, which is phototoxic and whose production consumes a key intracellular antioxidant (glutathione), are not clear apart from being involved in color patterns that confer concealment. The factors that have favored the evolution of pheomelanin thus remain a mystery, causing this pigment even to have been considered an "accident of nature." A recent hypothesis posits that pheomelanin has evolved because it represents an alternative mechanism to remove excess dietary cysteine, which can be toxic because of its oxidation. We tested for links between pheomelanin-based color and survival in both an intraspecific study of barn swallows Hirundo rustica and an interspecific study of 58 species of birds from North and Central America. As predicted on the basis that birds degrade excess dietary amino acids by transferring their amino group to uric acid synthesis, we found that under equal levels of uric acid in plasma, individuals or species with a higher intensity or greater proportion of plumage colored by pheomelanin (brown and chestnut coloration) had higher relative annual survival rates while controlling for the potentially confounding effects of age, sex, body size, and phylogenetic descent. Likewise, barn swallows with more intense pheomelanin-based coloration had higher prospects to survive the winter after controlling statistically for age, sex, body size, and level of uric acid. This supports the idea that pheomelanic traits evolve because of the removal of excess cysteine in nonstressful conditions, thus avoiding its toxic effects.

Number without language: comparative psychology and the evolution of numerical cognition

Number without language: comparative psychology and the evolution of numerical cognition

Where Are We Now? Bergmann’s Rule Sensu Lato in Insects

Bergmann's rule states that individuals of a species/clade at higher altitudes or latitudes will be larger than those at lower ones. A systemic review of the known literature on inter- and intraspecific variation in insect size along latitudinal or altitudinal clines was done to see how often such clines appeared and whether they reflected classwide, species-specific, or experimentally biased tendencies. Nearly even numbers of studies showed Bergmann clines and converse-Bergmann clines, where insects get smaller as latitude/altitude increases. In fact, the majority of studies suggested no clines at all. Small ranges may have obscured certain clines, while giant ranges may have introduced artifacts. Researchers examining interspecific patterns found clines less frequently than those examining intraspecific patterns because of variation among species within the clades, which renders interspecific studies unhelpful. Bergmann's rule does not apply to hexapods with nearly the same consistency as it does to endothermic vertebrates. The validity of Bergmann's rule for any group and range of insects is highly idiosyncratic and partially depends on the study design. I conclude that studies of Bergmann's rule should focus within species and look at widespread but contiguous populations to account for all sources of variation while minimizing error.

Seminal fluid proteins differ in abundance between genetic lineages of honeybees

Seminal fluid is transferred to the females' reproductive tract as part of the ejaculate and contains highly complex molecular machinery that is of central importance for male and female reproductive success. Interspecific studies suggest rapid evolutionary changes in the sequences of some seminal fluid proteins and also highlight the importance of specific seminal fluid proteins for sperm function and paternity success. Much less work has been conducted to study if variation in the steady-state abundance of seminal fluid proteins occurs within a species, which could provide a foundation for future selection to act upon. Here we used a unique breeding program of the honeybee Apis mellifera to provide evidence for quantified differences in seminal fluid protein abundances between three genetic lineages that have been bred for ~20 generations. We found the same subset of seminal fluid proteins to be present in all lines, but protein abundance or protein modification state varied significantly for 16% of the protein spots investigated. Protein spots with changed abundances were identified using mass spectrometry, with the abundance of a number documented from other species to be correlated with male fertility, reproductive success or immune-competence. We conclude that significant alterations in the abundance or modification state of specific proteins in seminal fluid can be linked to different genotypes in honeybees.

Untested assumptions about within-species sample size and missing data in interspecific studies

Phylogenetic comparative studies rely on species-specific data that often contain missing values and/or differ in sample size among species. These phenomena may violate statistical assumptions about the non-random variance component in sampling effort. A major reason why this assumption is often not fulfilled is because the probability of being sampled (i.e., being captured or observed) may depend on species-specific characteristics. Here, we test this assumption by using information on within-species sample sizes and missing data from five independent comparative datasets of European birds. First, we show that the two estimates of data availability (missing values and within-species sample size) are positively correlated and are associated with research effort in general (the number of papers published). Second, we demonstrate biologically meaningful relationships between data availability and phenotypic traits. For example, population size, risk-taking, and habitat specialization independently predicted within-species sample size. The key determinants of missing data were population size and distribution range. However, data availability was not structured by phylogenetic relationships. These results indicate that the accuracy of sampling is repeatable and distributed non-randomly among species, as several species-specific attributes determined the probability of observation. Therefore, data availability seems to be a species-specific trait that can be shaped by ecology, life history, and behavior. Such relationships raise issues about non-random sampling, which requires attention in comparative studies.

No evidence for the ‘expensive‐tissue hypothesis’ from an intraspecific study in a highly variable species

The 'expensive-tissue hypothesis' states that investment in one metabolically costly tissue necessitates decreased investment in other tissues and has been one of the keystone concepts used in studying the evolution of metabolically expensive tissues. The trade-offs expected under this hypothesis have been investigated in comparative studies in a number of clades, yet support for the hypothesis is mixed. Nevertheless, the expensive-tissue hypothesis has been used to explain everything from the evolution of the human brain to patterns of reproductive investment in bats. The ambiguous support for the hypothesis may be due to interspecific differences in selection, which could lead to spurious results both positive and negative. To control for this, we conduct a study of trade-offs within a single species, Thalassoma bifasciatum, a coral reef fish that exhibits more intraspecific variation in a single tissue (testes) than is seen across many of the clades previously analysed in studies of tissue investment. This constitutes a robust test of the constraints posited under the expensive-tissue hypothesis that is not affected by many of the factors that may confound interspecific studies. However, we find no evidence of trade-offs between investment in testes and investment in liver or brain, which are typically considered to be metabolically expensive. Our results demonstrate that the frequent rejection of the expensive-tissue hypothesis may not be an artefact of interspecific differences in selection and suggests that organisms may be capable of compensating for substantial changes in tissue investment without sacrificing mass in other expensive tissues.

Intraspecific Correlations of Basal and Maximal Metabolic Rates in Birds and the Aerobic Capacity Model for the Evolution of Endothermy

The underlying assumption of the aerobic capacity model for the evolution of endothermy is that basal (BMR) and maximal aerobic metabolic rates are phenotypically linked. However, because BMR is largely a function of central organs whereas maximal metabolic output is largely a function of skeletal muscles, the mechanistic underpinnings for their linkage are not obvious. Interspecific studies in birds generally support a phenotypic correlation between BMR and maximal metabolic output. If the aerobic capacity model is valid, these phenotypic correlations should also extend to intraspecific comparisons. We measured BMR, Msum (maximum thermoregulatory metabolic rate) and MMR (maximum exercise metabolic rate in a hop-flutter chamber) in winter for dark-eyed juncos (Junco hyemalis), American goldfinches (Carduelis tristis; Msum and MMR only), and black-capped chickadees (Poecile atricapillus; BMR and Msum only) and examined correlations among these variables. We also measured BMR and Msum in individual house sparrows (Passer domesticus) in both summer, winter and spring. For both raw metabolic rates and residuals from allometric regressions, BMR was not significantly correlated with either Msum or MMR in juncos. Moreover, no significant correlation between Msum and MMR or their mass-independent residuals occurred for juncos or goldfinches. Raw BMR and Msum were significantly positively correlated for black-capped chickadees and house sparrows, but mass-independent residuals of BMR and Msum were not. These data suggest that central organ and exercise organ metabolic levels are not inextricably linked and that muscular capacities for exercise and shivering do not necessarily vary in tandem in individual birds. Why intraspecific and interspecific avian studies show differing results and the significance of these differences to the aerobic capacity model are unknown, and resolution of these questions will require additional studies of potential mechanistic links between minimal and maximal metabolic output.

Seasonal stage differences overwhelm environmental and individual factors as determinants of energy expenditure in free-ranging red squirrels

Summary 1. Despite the central importance of the rate of energy expenditure in the lives of animals, the major drivers of within-species variation in energy expenditure remain uncertain, largely because most intraspecific studies focus on one or only a few potential determinants of expenditure. 2. Here, we examine the determinants of daily energy expenditure (DEE) in free-ranging female North American red squirrels (Tamiasciurus hudsonicus Erxleben) occupying a highly seasonal environment. By relating variation in 260 measurements of DEE from 176 individuals to key sources of seasonal (reproductive and foraging stages), environmental (resources and air temperature) and individual (body mass and individual identity) variation, our comprehensive analysis examines the relative importance of DEE predictors that have been more commonly examined in isolation. 3. Red squirrels demonstrated extensive variation in DEE with 5th (177 kJ per day) and 95th (660 kJ per day) percentile DEE levels that would correspond to mammals on an interspecific scale ranging in mass from 148 to 1120 g. 4. Seasonal stage differences accounted for most variation in DEE, with high expenditure during lactation and autumn hoarding, and very low expenditure during winter. Contrary to interspecific studies, energy expenditure increased with increasing ambient temperature and it was weakly related to body mass in all seasons except for winter. High resource availability was associated with reduced energy expenditure in winter, but elevated expenditure during lactation and hoarding. 5. Collectively, these results highlight substantial intraspecific variation in energy expenditure, most of which can be explained by a combination of seasonal stages and environmental conditions, and fundamental differences in the importance and direction of determinants of energy expenditure when examined at the intra- versus the interspecific level.

Understanding trait interactions and their impacts on growth in Scots pine branches across Europe

Summary1. Plants exhibit a wide variety in traits at different organizational levels. Intraspecific and interspecific studies have potential to demonstrate functional relationships and trade‐offs amongst traits, with potential consequences for growth. However, the distinction between the correlative and functional nature of trait covariation presents a challenge because traits interact in complex ways.2. We present an intraspecific study on Scots pine branches and use functional multi‐trait concepts to organize and understand trait interactions and their impacts on growth. Branch‐level traits were assessed for 97 branches from 12 Scots pine sites across Europe.3. To test alternative hypotheses on cause–effect relationships between anatomical traits, hydraulic traits and branch growth, we measured for each branch: the tracheid hydraulic diameter, double cell wall thickness, cell lumen span area, wood density, cavitation vulnerability, wood‐specific hydraulic conductivity, the leaf area to sapwood area ratio and branch growth. We used mixed linear effect models and path models to show how anatomical traits determine hydraulic traits and, in turn, how those traits influence growth.4. Tracheid hydraulic diameter was the best predictor of cavitation vulnerability (R2 = 0·09 explained by path model) and specific conductivity (R2 = 0·19) amongst anatomical traits. Leaf area to sapwood area ratio had the strongest direct effect on branch growth (R2 = 0·19) and was positively associated with the tracheid hydraulic diameter (R2 = 0·22). A number of bivariate correlations between traits could be explained by these functional relationships amongst traits.5. The plasticity in tracheid hydraulic diameter (10.0–15.1 μm) and leaf area to sapwood area ratio (600–6051 cm2 cm−2) and the maintenance of a minimum leaf water potential (between −2 and −2·5 MPa) appear to drive the anatomical and hydraulic traits of Scots pine across Europe. These properties are major drivers of the functional trait network underlying the growth variation amongst pine branches and thus possibly contribute to the ecological success of pines at a local and continental scale.

Grafting pea, faba bean, and lentil to improve pulse crop breeding

Gurusamy, V., Warkentin, T. D. and Vandenberg, A. 2012. Grafting pea, faba bean, and lentil to improve pulse crop breeding. Can. J. Plant Sci. 92: 31–38. In vivo grafting experiments were conducted to determine their potential for improving pulse crop breeding techniques for pea, faba bean and lentil. Four scion×five rootstock genotype combinations were used for pea. Survival of grafted pea scions was not affected by genotype of scions or rootstocks, even for wild subspecies. Some scion-rootstock combinations resulted in reduced flowering time for scions. Total seed production of rootstock regrowth plus grafted scion was greater than for ungrafted controls or rooted cuttings. For faba bean, four scion×four rootstock combinations showed scion-rootstock interaction affected percent survival and flowering time of grafted scions. In vivo grafting of lentil scions to faba bean rootstocks is reported for the first time. Two lentil genotypes were grafted onto four faba bean rootstocks. The effects of lentil scion and faba bean rootstock genotype were significant for percent survival, but not for seed production. Percent scion survival was 85% for pea, 56% for faba bean, and 55% for lentil scions on faba bean rootstocks. In vivo grafting techniques can help to maximize the size of F2populations for breeding and genetic studies. Intergeneric in vivo grafting of lentil onto faba bean rootstock could be useful for interspecific hybridization studies for lentil.

“Nullius in Bergmann” or the pluralistic approach to ecogeographical rules: a reply to Watt et al. (2010)

Bergmann's rule is one of the best known empirical generalizations in biogeography and remains a topic of much interest and debate. Watt et al. claimed in 2010 that the only definition for the rule should be the one devised by Bergmann in the 19th‐century. Based on direct translations from the original German manuscript, they concluded that tests of the rule should be restricted to interspecific studies of body size variation in endotherms. Furthermore, they suggested that Bergmann's heat conservation mechanism is an integral part of the rule and, hence, a simple falsificationist test of this mechanism might be enough to validate the rule. Here I advocate on a pluralistic approach to study ecogeographical rules, in general, and Bergmann's rule in particular. Our perceptions on the status and validity of laws and rules depend on the narrowness of the epistemological scope we adopt. Laws can have exceptions, do not necessarily have to be explanatory and may not be predictive. Also, we should differentiate between correlative and causal laws. Bergmann's rule is a correlative law, not a causal law, as Watt et al. implicitly assumed. It is a broad generalization with no inherent mechanism, and subject to the scrutiny of empirical investigation. Because of the ecological and evolutionary contingencies there will hardly be any law in ecology that is universally true. We have to consider each of these contingencies and study Bergmann's rule in a diversity of systems, organisms and levels of biological organization to gain further insight into the processes underlying the geographic variation in body size. On the basis of the empirical evidence to date, we cannot entirely dismiss a thermoregulatory mechanism to explain body size clines in both ectotherms and endotherms and support a food availability mechanism instead, as Watt et al. suggested. Even so, a unifying explanation for Bergmann's rule still remains elusive.

Intraspecific divergence in the lateral line system in the nine‐spined stickleback (Pungitius pungitius)

The mechanosensory lateral line system of fishes is an important organ system conveying information crucial to individual fitness. Yet, our knowledge of lateral line diversity is almost exclusively based on interspecific studies, whereas intraspecific variability and possible population divergence have remained largely unexplored. We investigated lateral line system variability in four marine and five pond populations of nine-spined stickleback (Pungitius pungitius). We found significant differences in neuromast number between pond and marine fish. In particular, three of seventeen lateral line regions (viz. caudal peduncle superficial neuromasts; canal neuromasts from the anterior trunk and caudal peduncle) showed strong divergence between habitats. Similar results were obtained with laboratory-reared individuals from a subset of populations, suggesting that the patterns found in nature likely have a genetic basis. Interestingly, we also found habitat-dependent population divergence in neuromast variability, with pond populations showing greater heterogeneity than marine populations, although only in wild-caught fish. A comparison of neutral genetic (F(ST)) and phenotypic (P(ST)) differentiation suggested that natural selection is likely associated with habitat-dependent divergence in neuromast counts. Hence, the results align with the conclusion that the mechanosensory lateral line system divergence among marine and pond nine-spined sticklebacks is adaptive.

Genome-Wide Survey of Natural Selection on Functional, Structural, and Network Properties of Polymorphic Sites in Saccharomyces paradoxus

To characterize the genetic basis of phenotypic evolution, numerous studies have identified individual genes that have likely evolved under natural selection. However, phenotypic changes may represent the cumulative effect of similar evolutionary forces acting on functionally related groups of genes. Phylogenetic analyses of divergent yeast species have identified functional groups of genes that have evolved at significantly different rates, suggestive of differential selection on the functional properties. However, due to environmental heterogeneity over long evolutionary timescales, selection operating within a single lineage may be dramatically different, and it is not detectable via interspecific comparisons alone. Moreover, interspecific studies typically quantify selection on protein-coding regions using the D(n)/D(s) ratio, which cannot be extended easily to study selection on noncoding regions or synonymous sites. The population genetic-based analysis of selection operating within a single lineage ameliorates these limitations. We investigated selection on several properties associated with genes, promoters, or polymorphic sites, by analyzing the derived allele frequency spectrum of single nucleotide polymorphisms (SNPs) in 28 strains of Saccharomyces paradoxus. We found evidence for significant differential selection between many functionally relevant categories of SNPs, underscoring the utility of function-centric approaches for discovering signatures of natural selection. When comparable, our findings are largely consistent with previous studies based on interspecific comparisons, with one notable exception: our study finds that mutations from an ancient amino acid to a relatively new amino acid are selectively disfavored, whereas interspecific comparisons have found selection against ancient amino acids. Several of our findings have not been addressed through prior interspecific studies: we find that synonymous mutations from preferred to unpreferred codons are selected against and that synonymous SNPs in the linker regions of proteins are relatively less constrained than those within protein domains. We present the first global survey of selection acting on various functional properties in S. paradoxus. We found that selection pressures previously detected over long evolutionary timescales have also shaped the evolution of S. paradoxus. Importantly, we also make novel discoveries untenable via conventional interspecific analyses.

Assessing tiger–prey interactions in Sumatran rainforests

Little is known about interactions between the critically endangered Sumatran tiger Panthera tigris sumatrae and its prey because of the difficulties associated with detecting these species. In this study, we quantify temporal overlap between the Sumatran tiger and five of its presumed prey species from four study areas comprising disturbed lowland to primary submontane forest. Data from 126 camera traps over 8984 camera days were used to estimate species activity patterns and, in turn, their overlap through the coefficient Δ (ranging from 0 to 1, i.e. no overlap to complete overlap). A newly developed statistical technique was applied to determine confidence intervals associated with respective overlap, which is important, as such measures of precision are usually not estimated in these types of study. Strong temporal overlap was found between tiger and muntjac Muntiacus muntjac (Δ=0.80, 95%CI=0.71–0.84) and tiger and sambar Cervus unicolor (Δ=0.81, 0.55–0.85), with the latter illustrating the importance of measuring precision. According to the foraging theory, Sumatran tigers should focus on expending lower levels of energy searching for and then capturing larger bodied prey that present the least risk. Hence, surprisingly, there was little overlap between the crepuscular tiger and the largest-bodied prey species available, the nocturnal tapir Tapirus indicus (0.52, 0.44–0.60), suggesting that it is not a principal prey species. This study provides the first insights into Sumatran tiger–prey temporal interactions. The ability to estimate overlap statistics with measures of precision has obvious and wide benefits for other predator–prey and interspecific competition studies.

Effect of Malathion on Larval Competition Between Aedes albopictus and Aedes atropalpus (Diptera: Culicidae)

Aedes albopictus (Skuse) and Aedes atropalpus (Coquillett) (Diptera: Culicidae) are container-dwelling mosquito species that are well established in the eastern United States. Interspecific larval competition studies have shown Ae. albopictus to be a superior competitor over many species. A laboratory experiment was conducted in artificial containers to evaluate the effects of malathion on larval interactions between Ae. albopictus and Ae. atropalpus. The survivorship of Ae. albopictus increased with increasing Ae. atropalpus densities in control but decreased with increasing Ae. atropalpus densities in the presence of malathion. Alternatively, Ae. atropalpus survivorship did not differ between control and malathion treatments. Developmental times were not affected by interspecific competition in both treatments for either species. These results show that malathion could facilitate coexistence between Ae. albopictus and Ae. atropalpus. This demonstrates how sublethal concentrations of malathion (and perhaps other pesticides with similar modes of action) can enable an inferior competitor to coexist in the same habitat with a superior competitor. This is the first report of synergistic survival of a weaker mosquito competitor in the presence of a pesticide due to condition-specific competition.

Hovering guards of the stingless bee Tetragonisca angustula increase colony defensive perimeter as shown by intra- and inter-specific comparisons

Social insects need to defend their nest against robbery, parasitism and predation. The stingless bee Tetragonisca angustula is unique in that it has guards that hover near the nest entrance in addition to guards that stand at the entrance. We tested both the general hypothesis that hovering guards increase the effectiveness with which flying intruders are detected and the specific hypothesis that hovering guards improve the detection of workers of the obligate robber bee, Lestrimellita limao. In an intraspecific study comparing colonies, we found a strong positive relationship between the number of hovering guards and the distance at which a dummy robber bee or L. limao worker, experimentally moved towards the nest entrance, was detected. These results were mirrored in an interspecific study showing that four species of stingless bees with similar population colonies but which lacked hovering guards, detected L. limao only at the nest entrance, in contrast to T. angustula. In addition, we found that a greater number of attacks by guards occurred when dummies were impregnated with citral, a major component of L. limao mandibular gland odour. Our results support the hypothesis that T. angustula hovering guards increase the detection perimeter for flying intruders, especially L. limao.