High Wing Loading Research Articles

Extremely large sweep amplitude enables high wing loading in giant hovering insects

Beetle Allomyrina dichotoma is one of the largest insects that performs many remarkable modes of locomotion, particularly hovering flight capability. In order to stay airborne, its flexible hindwings are flapped so as to work as a thrust generator. However, the wing loading of the beetle is relatively large (38.94 ± 3.73 N m−2) compared to those of other insects and hummingbirds, indicating that it is challenging for it to achieve flight. Here, we measured the hindwing morphology and kinematics of the beetle in order to discover its flight performance. Unlike many other insects, the beetle flaps its hindwings with an extremely large sweep amplitude of about 191.33 ± 6.12 deg at high flapping frequencies ranging from 36 to 41 Hz (mean wing tip speed ≈ 13.45 ± 0.58 m s−1). These capabilities enable the beetle to produce enough lift force to stay airborne with its bulky body (4–10 g). In order to investigate how the sweep amplitude affects the beetle’s flight efficiency, we utilized the unsteady blade element model to estimate the power requirements of the same vertical force production for various sweep amplitudes. The results indicate that the sweep amplitude as high as 190 deg is more beneficial for power requirements than the smaller amplitudes, which require higher frequencies resulting in higher inertial powers to produce the same vertical force. Thus, for this large beetle, high sweep amplitude may be a biological strategy for staying airborne. In addition, we thoroughly discussed the effects of input constraints on the outcome by investigating power loadings for variable sweep amplitudes at a constant vertical force, mean wing tip speed, and flapping frequency. Effect of wing surface area was also investigated and discussed to provide useful information for the development of an insect-inspired flapping-wing robot.

Bat sonar and wing morphology predict species vertical niche.

The use of echolocation allows insectivorous bats to access unique foraging niches by locating obstacles and prey with ultrasounds in complete darkness. To avoid interspecific competition, it is likely that sonar features and wing morphology co-evolved with species vertical distribution, but due to the technical difficulties of studying flight in the vertical dimension, this has never been demonstrated with empirical measurements. The authors equipped 48 wind masts with arrays of two microphones and located the vertical distribution of a community of 19 bat species and two species groups over their annual activity period (>8000 nights). The authors tested the correlation between the proportion of flights at height and the acoustic features of bat calls as well as their wing morphology. The authors found that call peak frequency and bandwidth are good predictors of bat use of the vertical space regardless of their acoustic strategies (i.e., gleaning, hawking, or detecting prey flutter). High wing aspect ratios and high wing loadings were associated with high proportions of time spent at height, confirming hypotheses from the literature.

Hymenoptera flight muscle mitochondrial function: Increasing metabolic power increases oxidative stress

Insect flight is a high intensity activity, but biomechanical and metabolic requirements may vary depending on life style and feeding mode. For example, bees generally feed on pollen and nectar, whereas wasps also actively hunt and scavenge heavy prey. These variations in metabolic demands may result in different capacities of metabolic pathways in flight muscle, and utilisation some of these pathways may come at a cost of increased free radical production. To examine how metabolic requirements and oxidative stress vary between species, we explored the variation in flight mechanics and metabolism of the honeybee (Apis mellifera), bumblebee (Bombus terrestris), and German wasp (Vespula germanica). Wing structures and flight muscle properties were compared alongside measures of oxygen flux and reactive oxygen species (ROS) production from permeabilised flight muscle. The wasp wing structure is best adapted for carrying heavy loads, with the highest wing aspect ratio, lowest wing loading, and highest flight muscle ratio. Bumblebees had the lowest wing aspect ratio and flight muscle ratio, and highest wing loading. Although wasps also had the highest rates of oxygen consumption during oxidative phosphorylation, oxygen consumption did not increase in the wasp muscle following chemical uncoupling, while it did for the two bee species. While mitochondrial glycerol 3-phosphate dehydrogenase (mGPDH) mediated oxygen flux was greatest in wasps, muscle fibres released greater amounts of ROS through this pathway. Overall, the wasp has maximised lifting capacities through varying wing and flight muscle mass and by maximising OXPHOS capacities, and this accompanies elevated ROS production.

Species With Greater Aerial Maneuverability Have Higher Frequency of Collisions With Aircraft: A Comparative Study

Antipredator responses may appear unsuccessful when animals are exposed to approaching vehicles, often resulting in mortality. Recent studies have addressed whether certain biological traits are associated with variation in collision risk with cars, but not with higher speed-vehicles like aircraft. Our goal was to establish the association between different species traits (i.e., body mass, eye size, brain size, wing loading, wing aspect ratio) and the frequency of bird collisions with aircraft (hereafter, bird strikes) using a comparative approach controlling for the effects of shared ancestry. We proposed directional predictions as to how each of the species traits would affect the frequency of bird strikes. Considering 39 bird species with all traits represented, the model containing wing loading had the best fit to account for the variance in bird strikes across species. In another model with 54 species exploring the fit to different polynomial models but considering only wing loading, we found that wing loading was negatively and linearly associated with the frequency of bird strikes. Counterintuitively, species with lower wing loading (hence, higher maneuverability) had a higher frequency of bird strikes. We discuss potential non-mutually exclusive explanations (e.g., high wing loading species fly faster, thus gaining some extra time to avoid the aircraft flight path; high wing loading species are hazed more intensively at airports, which could lower collisions, etc.). Ultimately, our findings uncovered that species with low wing loading get struck at a higher rate at airports, which reduces the safety risk for humans because these species tend not to cause damaging strikes, but the consequences of their potentially higher local mortality are unknown.

Experimental Increases in Foraging Costs Affect Pectoralis Muscle Mass and Myostatin Expression in Female, but Not Male, Zebra Finches (Taeniopygia guttata).

Skeletal muscle remodeling is an important component of phenotypic flexibility in birds and impacts organismal metabolism and performance, which could potentially influence fitness. One regulator of skeletal muscle remodeling is myostatin, an autocrine/paracrine muscle growth inhibitor that may be down-regulated under conditions promoting heavier muscle masses. In this study, we employed protocols requiring hovering while foraging to increase foraging costs and modify phenotypes of zebra finches (Taeniopygia guttata). We examined the effects of high-cost foraging (HF) on skeletal muscle masses and used real-time reverse-transcription polymerase chain reaction and Western blots to measure gene and protein expression of myostatin and its metalloproteinase activators tolloid-like proteases TLL-1 and TLL-2 in pectoralis muscle. Female finches average shorter wings and higher wing loading than males, so increased flight costs might be expected to disproportionately affect females. Indeed, HF female finches exhibited reduced total fat masses, increased pectoralis muscle masses, and lower myostatin protein levels than controls. Male finches showed no significant differences in pectoralis muscle masses or myostatin protein levels between HF and control birds. Myostatin, TLL-1, and TLL-2 messenger RNA (mRNA) expression remained stable between treatments for both female and male finches. Myostatin mRNA and protein levels showed variable directions of correlations with pectoralis mass residuals among treatments. Thus, these data offer only mixed support for a regulatory role for myostatin in mediating the flexibility of pectoralis muscle phenotypes of small birds.

IMPACT OF VARIABLE GEOMETRY MINIFLAPS ON SAILPLANE FLIGHT CHARACTERISTICS

Miniflaps (also known as mini-TED, active Gurnay flaps, etc.) located at the wing trailing edge enable to increase wing lift and reduce the aerodynamic drag. Variable geometry miniflaps (VGMF) elaborated at the Estonian Aviation Academy enable to expand the wing area up to 6.5%, when extended, and, at the same time, deflect 16.7 degrees downwards. The use of VGMF is especially promising for reducing the airspeed and sink speed of modern high wing loading sailplanes flying in thermals. The VGMFs were built in cooperation with the Lithuanian company JSC “Sportine Aviacija ir Ko”. They were fixed inside the sailplane LAK-17B trailing flaps. During the test flights, the VGMF effect on the sailplane’s sink speed was measured at different airspeeds and different flap positions. The flight parameters were recorded electronically and later calculated for standard atmosphere. The method of parallel flight was used for comparison. The results indicate that with the flaps position at + 9 degrees, the CAS 79.5 km/h sink speed decreased 0.775–0.555 m/s, i.e. 39.6%. The sailplane’s wing loading was 39.4 kg/m 2. With the flaps position at + 15 degrees, the optimal airspeed decreased, but the sink speed, compared to the previous, rose. Cl max increased with the VGMF 1.41–1.58 (i.e. 12%).

Wings of tropical finches: interspecific differences in shape are consistent with levels of mobility, but moult and feather fault patterns are more complex

ABSTRACTBirds’ wings reflect their life histories, suggesting evolutionary selection for wing shapes and moult strategies. Compared to sedentary species, long-distance migrants have narrower wings (for fast, efficient flight); they have fewer feather faults and avoid moulting flight feathers during migration (to optimise flight surface performance). It is unclear whether these patterns apply to species that fly short-intermediate distances, like tropical nomads. We compared wing shape, feather faulting, and flight-feather moult across five finch species from northern Australia with varying mobility: Pictorella Mannikins (Heteromunia pectoralis) and Gouldian Finches (Erythrura gouldiae) are highly mobile, nomadic at regional scales; Long-tailed Finches (Poephila acuticauda) and Double-barred Finches (Taeniopygia bichenovii) are sedentary with local nomadism; Crimson Finches (Neochmia phaeton) are sedentary. More mobile species had narrower wings and higher wing loading than relatively sedentary species, and less feather faulting. Variation in moult strategies was complex, but mobile species carried out moult fast, in a short time window, and moulted a single feather at a time. Unexpectedly, Gouldian Finch wings were more suited for fast efficient flight than Pictorella Mannikin wings, and yet they had more feather faulting. Differences in mobility may be a key dimension of niche separation that allows these species to co-exist.

Mixed effects of gating subterranean habitat on bats: A review

ABSTRACTCaves and abandoned mines, roosting environments for many bat species including subterranean obligates, are under human pressure (e.g., recreation) and pose threats to humans. Gating entrances of these subterranean features prevents human entry while protecting habitat for bats. However, bat response to these gates is not well understood and no comprehensive synthesis with species‐specific recommendations on protection measures for bat roosts exists. We reviewed literature on effects of gating subterranean habitat on bats and roost microclimate and found 22 studies and reports from North America, Australia, Europe, and Asia. Ten studies examined effects of gating on bat use, 9 described behavioral responses to gates, and 5 presented effects of gates on roost microclimate. Older gate designs (1950–1970), such as cement walls or iron doors, negatively affected bat use and roost microclimate, but responses to modern (1970s to present), bat‐compatible gate designs varied. Short‐term responses of bats to bat‐compatible gates were negative and included increases in energetically expensive flight behaviors. Although long‐term responses included a mix of population and species trends, we attribute these mixed responses in part to differences in flight agility among species. Bats with moderate to high agility (low wing loading, broad call bandwidths) adjust to gates, but species with low agility (high wing loading and narrow call bandwidths) may abandon sites after gating. Other factors including bat density in roosts and size of the cave or mine entrance also affect acceptance of gates and should be considered in gate design. Current gate designs are adequate for most species. Based on available literature, we recommend use of bat‐compatible gates with horizontal bar spacing of 14.6 cm, bar types that offer high structural rigidity in areas where vandalism is likely, and periodic monitoring of gates to ensure they remain in good condition. © 2017 The Wildlife Society.

Seasonal activity and unpredicted polar front migration of northern New Zealand Common Diving Petrels (Pelecanoides urinatrix)

ABSTRACTThe Common Diving Petrel (Pelecanoides urinatrix) is an abundant seabird species within New Zealand waters. Breeding in this species occurs from late July through to December and adults are thought to remain in New Zealand waters during non-breeding months. Yet no studies of the spatial ecology of the species within New Zealand have been undertaken to confirm this. We used geolocation loggers with seawater immersion probes to characterise the year-round distribution and activity of P. urinatrix from two colonies (Kauwahaia Island and Burgess Island) in northern New Zealand. Adult birds exhibited extended pre-laying occupation of breeding sites (~95 days) and throughout breeding remained within 300 km of their colony. Burgess Island birds returned near nightly to their burrows throughout the breeding season. During non-breeding birds made a rapid and unpredicted migration south-east to the Antarctic polar front (covering 3000–5000 km in 3–13 days) and exhibited a dramatic shift in behaviour by spending 95% of their time on (and under) the water surface. Given its already high wing loading, it appears that the primary moult reduced flight efficiency during this time. But by targeting highly productive oceanic regions, and the improved foraging opportunities therein, P. urinatrix may offset this reduced flight capacity.

Wing size but not wing shape is related to migratory behavior in a soaring bird

Both wing size and wing shape affect the flight abilities of birds. Intra and inter‐specific studies have revealed a pattern where high aspect ratio and low wing loading favour migratory behaviour. This, however, have not been studied in soaring migrants. We assessed the relationship between the wing size and shape and the characteristics of the migratory habits of the turkey vulture Cathartes aura, an obligate soaring migrant. We compared wing size and shape with migration strategy among three fully migratory, one partially migratory and one non‐migratory (resident) population distributed across the American continent. We calculated the aspect ratio and wing loading using wing tracings to characterize the wing morphology. We used satellite‐tracking data from the migratory populations to calculate distance, duration, speed and altitude during migration. Wing loading, but not aspect ratio, differed among the populations, segregating the resident population from the completely migratory ones. Unlike what has been reported in species using flapping flight during migration, the migratory flight parameters of turkey vultures were not related to the aspect ratio. By contrast, wing loading was related to most flight parameters. Birds with lower wing loading flew farther, faster, and higher during their longer journeys. Our results suggest that wing morphology in this soaring species enables lower‐cost flight, through low wing‐loading, and that differences in the relative sizes of wings may increase extra savings during migration. The possibility that wing shape is influenced by foraging as well as migratory flight is discussed. We conclude that flight efficiency may be improved through different morphological adaptations in birds with different flight mechanisms.

Residual Life Estimation of an Attach Angle for a Cargo Aircraft

Wing–fuselage attachment formally known as attach angle for cargo aircraft is an important component that connects the center wing to the fuselage. Owing to high wing loading, the edge and the center cracks on attach angle were detected and reported by users around the world. In this work, the critical edge and center crack lengths are computed using numerical techniques. The used procedure was first analytically validated for a simple case and then applied to original aircraft component. Based upon the critical crack lengths and crack-growth analysis, the remaining useful life of the attach angle has been estimated. Estimation of the remaining useful life for different parts of the aircraft at times become an essential requirement for aircraft operators, and the procedure adopted for this work can therefore be utilized for similar analysis of any aircraft part and would result in significant cost savings.

Flight Morphology, Compound Eye Structure and Dispersal in the Bog and the Cranberry Fritillary Butterflies: An Inter- and Intraspecific Comparison.

Understanding dispersal is of prime importance in conservation and population biology. Individual traits related to motion and navigation during dispersal may differ: (1) among species differing in habitat distribution, which in turn, may lead to interspecific differences in the potential for and costs of dispersal, (2) among populations of a species that experiences different levels of habitat fragmentation; (3) among individuals differing in their dispersal strategy and (4) between the sexes due to sexual differences in behaviour and dispersal tendencies. In butterflies, the visual system plays a central role in dispersal, but exactly how the visual system is related to dispersal has received far less attention than flight morphology. We studied two butterfly species to explore the relationships between flight and eye morphology, and dispersal. We predicted interspecific, intraspecific and intersexual differences for both flight and eye morphology relative to i) species-specific habitat distribution, ii) variation in dispersal strategy within each species and iii) behavioural differences between sexes. However, we did not investigate for potential population differences. We found: (1) sexual differences that presumably reflect different demands on both male and female visual and flight systems, (2) a higher wing loading (i.e. a proxy for flight performance), larger eyes and larger facet sizes in the frontal and lateral region of the eye (i.e. better navigation capacities) in the species inhabiting naturally fragmented habitat compared to the species inhabiting rather continuous habitat, and (3) larger facets in the frontal region in dispersers compared to residents within a species. Hence, dispersers may have similar locomotory capacity but potentially better navigation capacity. Dispersal ecology and evolution have attracted much attention, but there are still significant gaps in our understanding of the mechanisms of dispersal. Unfortunately, for many species we lack detailed information on the role of behavioural, morphological and physiological traits for dispersal. Our novel study supports the existence of inter- and intra-specific evolutionary responses in both motion and navigation capacities (i.e. flight and eye morphology) linked to dispersal.

Dispersal Polymorphisms in Invasive Fire Ants.

In the Found or Fly (FoF) hypothesis ant queens experience reproduction-dispersal tradeoffs such that queens with heavier abdomens are better at founding colonies but are worse flyers. We tested predictions of FoF in two globally invasive fire ants, Solenopsis geminata (Fabricius, 1804) and S. invicta (Buren, 1972). Colonies of these species may produce two different monogyne queen types—claustral queens with heavy abdomens that found colonies independently, and parasitic queens with small abdomens that enter conspecific nests. Claustral and parasitic queens were similarly sized, but the abdomens of claustral queens weighed twice as much as those of their parasitic counterparts. Their heavier abdomens adversely impacted morphological predictors of flight ability, resulting in 32–38% lower flight muscle ratios, 55–63% higher wing loading, and 32–33% higher abdomen drag. In lab experiments maximum flight durations in claustral S. invicta queens decreased by about 18 minutes for every milligram of abdomen mass. Combining our results into a simple fitness tradeoff model, we calculated that an average parasitic S. invicta queen could produce only 1/3 as many worker offspring as a claustral queen, but could fly 4 times as long and have a 17- to 36-fold larger potential colonization area. Investigations of dispersal polymorphisms and their associated tradeoffs promises to shed light on range expansions in invasive species, the evolution of alternative reproductive strategies, and the selective forces driving the recurrent evolution of parasitism in ants.

Postnatal growth and age estimation in Scotophilus kuhlii.

Adequate postnatal growth is important for young bats to develop skilled sensory and locomotor abilities, which are highly associated with their survival once independent. This study investigated the postnatal growth and development of Scotophilus kuhlii in captivity. An empirical growth curve was established, and the postnatal growth rate was quantified to derive an age-predictive equation. By further controlling the fostering conditions of twins, the differences in the development patterns between pups that received maternal care or were hand-reared were analyzed to determine whether the latter developed in the same manner as their maternally reared counterparts. Our results indicate that both forearm length and body mass increased rapidly and linearly during the first 4 weeks, after which the growth rate gradually decreased to reach a stable level. The first flight occurred at an average age of 39 days with a mean forearm length and body mass of 92.07% and 70.52% of maternal size, respectively. The developmental pattern of hand-reared pups, although similar to that of their maternally reared twin siblings, displayed a slightly faster growth rate in the 4th and 5th weeks. The heavier body mass of hand-reared pups during the pre-fledging period may cause higher wing loading, potentially influencing the flight performance and survival of the bats once independent.

Body size affects individual winter foraging strategies of thick-billed murres in the Bering Sea.

Foraging and migration often require different energetic and movement strategies. Though not readily apparent, constraints during one phase might influence the foraging strategies observed in another. For marine birds that fly and dive, body size constraints likely present a trade-off between foraging ability and migration as smaller bodies reduce flight costs, whereas larger bodies are advantageous for diving deeper. This study examines individual wintering strategies of deep diving thick-billed murres (Uria lomvia) breeding at three colonies in the south-eastern Bering Sea: St Paul, St George and Bogoslof. These colonies, arranged north to south, are located such that breeding birds forage in a gradient from shelf to deep-water habitats. We used geolocation time-depth recorders and stable isotopes from feathers to determine differences in foraging behaviour and diet of murres during three non-breeding periods, 2008-2011. Body size was quantified by a principal component analysis (wing, culmen, head+bill and tarsus length). A hierarchical cluster analysis identified winter foraging strategies based on individual movement, diving behaviour and diet (inferred from stable isotopes). Structural body size differed by breeding island. Larger birds from St Paul had higher wing loading than smaller birds from St George. Larger birds, mainly from St Paul, dove to deeper depths, spent more time in the Bering Sea, and likely consumed higher trophic-level prey in late winter. Three winter foraging strategies were identified. The main strategy, employed by small birds from all three breeding colonies in the first 2 years, was characterized by high residency areas in the North Pacific south of the Aleutians and nocturnal diving. In contrast, 31% of birds from St Paul remained in the Bering Sea and foraged mainly during the day, apparently feeding on higher trophic-level prey. Throat feather stable isotopes indicated that individuals exhibited flexibility in the use of this colony-specific foraging strategy. The third strategy only occurred in 2010/2011, when birds dove more and deeper, suggesting limited prey resources. Foraging strategies partitioned with respect to annual differences, presumably in response to shifts in distribution of prey, and were linked to body size. The presence of a colony-specific wintering strategy suggests the potential for overwinter survival differences between these populations.

Performances of a small hypersonic airplane (HyPlane)

In the present work a preliminary performance study regarding a small hypersonic airplane named HyPlane is presented. It is designed for point-to-point medium range hypersonic trip, within the “urgent business travel” market segment. The vehicle is also consistent with long duration suborbital space tourism missions, in the frame of the Space Renaissance (SR) Italia Space Tourism Program. The design of such a hypersonic airplane is based on the concept of integrating available technologies developed for aeronautical and space atmospheric re-entry systems.The vehicle, characterized by high aerodynamic efficiency and low wing loading and powered by Turbine Based Combined Cycle (TBCC) engines plus a throtteable Rocket, is able to produce a reduced sonic boom and to perform Horizontal Takeoff and Horizontal Landing (HTHL) on runways.Aerodynamic and propulsive performances for the different flight regimes encountered during the missions are studied. Aerodynamic heating effects are analyzed, in order to identify suitable structures and materials design to sustain the hypersonic flight conditions. Different flight paths are also investigated, including hypersonic cruise and suborbital parabolic trajectories, which provide Space tourists with the opportunity of long duration missions, offering short and repeated periods of low-gravity, in the high stratosphere where a large view of the Earth is ensured.

Mobility explains the response of aerial insectivorous bats to anthropogenic habitat change in the Neotropics

Habitat loss due to anthropogenic activities is a critical threat to biodiversity. Understanding which factors determine species persistence in degraded environments is essential for conservation management. Here, we investigated how aerial insectivorous bats in the Neotropics, an ecologically important, but understudied group of vertebrates, are affected by deforestation and urbanization. We conducted a standardized acoustic survey in four habitat types in 14 areas across Panama (500km×260km), a scale hitherto unprecedented for the Neotropics, and assessed occupancy in 13 aerial insectivorous bat species, accounting for species- and habitat-specific detectability. In addition, we used wing measures to derive a proxy for flight efficiency and flight speed, as an indication of species-specific mobility. Results show that detectability does not only vary strongly among species but also, within a species, depending on the habitat: for nine species intra-specific detectability varied >10%. Model estimates for occupancy revealed that aerial insectivorous bats are highly heterogeneous in their response to deforestation and urbanization. Canopy height and canopy cover predicted occupancy best for most species. Relating occupancy to mobility we found that less mobile species with broad wings (high wing loading and low aspect ratio) decrease occupancy in deforested and urban areas, while more mobile species with narrow wings increase in these habitats. We suggest that this pattern applies also to species not analyzed in this study. Our study exemplifies how morphological information can help define conservation priorities when information on occurrence and distribution is limited.

Leucocyte profiles of Arctic marine birds: correlates of migration and breeding phenology.

Most Arctic marine birds are migratory, wintering south of the limit of annual pack ice and returning north each year for the physiologically stressful breeding season. The Arctic environment is changing rapidly due to global warming and anthropogenic activities, which may influence the timing of breeding in relation to arrival times following migration, as well as providing additional stressors (e.g. disturbance from ships) to which birds may respond. During stressful parts of their annual cycle, such as breeding, birds may reallocate resources so that they have increased heterophil-to-lymphocyte ratios in their white blood cell (leucocyte) profiles. We analysed leucocyte profiles of nine species of marine birds to establish reference ranges for these species in advance of future Arctic change. Leucocyte profiles tended to cluster among taxonomic groups across studies, suggesting that reference values for a particular group can be established, and within species there was evidence that birds from colonies that had to migrate farther had higher heterophil-to-lymphocyte ratios during incubation than those that did not have to travel as far, particularly for species with high wing loading.

Correlates of dispersal extent predict the degree of population genetic structuring in bats

Dispersal is essential for maintaining demographic and genetic connectivity. For bats, correlates of dispersal extent such as morphology and movement dynamics are reported as having an influence on population genetic structure although these traits exhibit co-variance which has not been previously examined. We used a principal components framework with phylogenetically independent contrasts to compare five dispersal extent predictors (wing loading, aspect ratio, geographic range size, migratory status and median latitude) with population genetic structure among bats. We found that high wing loading values and migration negatively correlate with genetic structure after accounting for co-variance. These findings suggest that bats that can achieve higher flight speeds and migrate seasonally have higher gene flow and resultant genetic connectivity relative to bats that fly slower and do not migrate. These results represent a step towards understanding factors that shaped the genetic structure of bat populations.

Costs and constraints in aphid‐ant mutualism

AbstractWhile many studies have demonstrated that ants provide beneficial services to aphids, Bristow (Ant‐plant interactions, Oxford University Press, Oxford, 104–119, 1991) first questioned why so few aphid species are ant‐attended. Phylogenetic trees have demonstrated multiple gains and loss of ant‐attendance in the course of aphid‐ant interactions, implying that mutualisms easily form and dissolve. Several studies have reported the factors that influence the formation and maintenance of aphid‐ant interactions. Examples include the physiological costs of ant attendance, competition for mutualistic ants, ant predation on aphids, the influence of host plants, and parasitoid wasps. Recent physiological techniques have also revealed the chemical component of aphid‐ant mutualisms. The honeydew of ant‐attended aphids contains melezitose (a trisaccharide), which has an important role in aphid‐ant interactions. Studies of cuticular hydrocarbons on aphids and ants have clarified the underlying mechanisms of ant predation on aphids. Attending ants also reduce aphid dispersal ability, causing the formation of fragmented aphid populations with low genetic diversity in each population. The reduced aphid dispersal could be partly explained by higher wing loading and reduction of flight apparatus due to ant attendance. Whether ant attendance is associated with the range of host plants of aphids or genetic variation in microorganism in aphids remain to be explored.