Asymmetric interspecific aggression in New Guinean songbirds that replace one another along an elevational gradient

A common pattern in tropical avifaunas is for closely related species to inhabit largely parapatric elevational distributions such that they replace one another along the elevational gradient. A long‐standing hypothesis for this pattern is that parapatry is maintained by interspecific interference competition mediated by interspecific aggression. However, empirical tests of this hypothesis remain scarce. We used reciprocal playback experiments to measure interspecific aggression in five species‐pairs of New Guinean passerine elevational replacements. We found evidence of interspecific aggression in three species‐pairs. In these three cases, interspecific aggression was asymmetric, with the lower elevation species more aggressive towards the upper elevation species than vice versa. Two patterns suggest that this interspecific aggression is a learned response to the presence of a heterospecific competitor rather than misdirected intraspecific aggression or an evolved response to a competitor. First, when present, interspecific aggression was always strongest at the upper elevation range margin of the lower elevation species (i.e. in the elevational zone in which the two species were found in close proximity and thus interacted with each other), and diminished over very short distances away from this zone. Secondly, the two species‐pairs that did not exhibit interspecific aggression had narrow ‘no man's land’ gaps between their elevational distributions such that heterospecifics did not encounter one another, possibly explaining the lack of interspecific aggression in these examples. Our results support the hypothesis that interspecific aggression is one factor influencing elevational limits in species‐pairs of New Guinean elevational replacements.     

The costs of avoiding matings in the dung fly Sepsis cynipsea

Mating is generally assumed to carry costs, particularly forfemales, which have to be traded off against each other andagainst the fitness benefits of mating. To understand any particularmating system and the evolution of sexual conflict, these costshave to be evaluated. Female dung flies, Sepsis cynipsea (Diptera:Sepsidae), typically attempt to dislodge mounted males by vigorousshaking. Such female reluctance to mate can only evolve if the cost of avoiding matings does not exceed the cost of copulation.We investigated female precopulatory costs of assessing andrejecting males in terms of increased predation, wing injuries,and (indirectly) energetics, all ultimately affecting mortality,and compared them to the costs of copulation assessed in thisand a companion study. Females housed with a male had lower survivorship than females housed with another female. This waslargely due to the costs of copulation rather than presumedenergetic costs of avoiding males, which were minor. Male harassmentaugmented female wing injuries, which accumulate with age inthe field and laboratory, but in laboratory experiments usingone common predator, wing injuries did not increase the susceptibility of S. cynipsea to predation, nor did their mating behavior perse. Instead, predation was highest and survivorship lowestin all-male groups, probably because males are more activein search of females and harass each other. Overall, the precopulatorycosts of mate assessment and rejection were low relative tothe costs of copulating, explaining female reluctance behavior in this and possibly other species.     

ECOLOGICAL HISTORY AND EVOLUTION IN A NOVEL ENVIRONMENT: HABITAT HETEROGENEITY AND INSECT ADAPTATION TO A NEW HOST PLANT

Environmental heterogeneity has often been implicated in the maintenance of genetic variation. However, previous research has not considered how environmental heterogeneity might affect the rate of adaptation to a novel environment. In this study, I used an insect-plant system to test the hypothesis that heterogeneous environments maintain more genetic variation in fitness components in a novel environment than do uniform environments. To manipulate recent ecological history, replicate populations of the dipteran leafminer Liriomyza trifolii were maintained for 20 generations in one of three treatments: a heterogeneous environment that contained five species of host plant, and two uniform environments that contained either a susceptible chrysanthemum or tomato. The hypothesis that greater genetic variance for survivorship and developmental time on a new host plant (a leafminer-resistant chrysanthemum) would be maintained in the heterogeneous treatment relative to the uniform environments was then tested with a sib-analysis and a natural selection experiment. Populations from the heterogeneous host plant treatment had no greater genetic variance in either larval survivorship or developmental time on the new host than did populations from either of the other treatments. Moreover, the rate of adaptation to the new host did not differ between the ecological history treatments, although the populations from the uniform chrysanthemum treatment had higher mean survivorship throughout the selection experiment. The estimates of the heritability of larval survivorship from the sib-analysis and selection experiment were quite similar. These results imply that ecologically realistic levels of environmental heterogeneity will not necessarily maintain more genetic variance than uniform environments when traits expressed in a particular novel environment are considered.     

The cost of reproduction in rosehip flies,Rhagoletis basiola: Eggs are time

Summary The hypothesis was tested that egg-laying by rosehip fruit flies,Rhagoletis basiola, would lead to reduced adult survivorship compared with oviposition-deprived flies, through experimental manipulation. Oviposition-deprived flies were assigned to two groups: mated and spinster. All three treatment groups were permitted to spend equal time and energy exploring and handling hosts but only the former group were allowed to deposit eggs (three/day). Results clearly showed that ovipositing flies suffered significantly higher mortality rates and that mated and spinster flies produced similar survivorship patterns. I consider these results both as a foraging problem (i.e. laying eggs in inferior hosts can cost both eggs and life) and as a more general life history problem.     

A New Approach that Eliminates Handling for Studying Aggression and the "Loser" Effect in Drosophila melanogaster

Aggressive behavior in Drosophila melanogaster is composed of the sequential expression of stereotypical behavioral patterns (for analysis see ¹). This complex behavior is influenced by genetic, hormonal and environmental factors. As in many organisms, previous fighting experience influences the fighting strategy of flies and the outcome of later contests: losing a fight increases the probability of losing later contests, revealing "loser" effects that likely involve learning and memory ^2-4. The learning and memory that accompanies expression of complex social behaviors like aggression, is sensitive to pre-test handling of animals ^5,6. Many experimental procedures are used in different laboratories to study aggression ^7-9, however, no routinely used protocol that excludes handling of flies is currently available. Here, we report a new behavioral apparatus that eliminates handling of flies, using instead their innate negative geotactic responses to move animals into or out of fighting chambers. In this protocol, small circular fight arenas containing a food cup are divided into two equal halves by a removable plastic slider prior to introduction of flies. Flies enter chambers from their home isolation vials via sliding chamber doors and geotaxis. Upon removal of plastic sliders, flies are free to interact. After specified time periods, flies are separated again by sliders for subsequent experimentation. All of this is done easily without handling of individual flies. This apparatus offers a novel approach to study aggression and the associated learning and memory, including the formation of "loser" effects in fly fights. In addition, this new general-purpose behavioral apparatus can be employed to study other social behaviors of flies and should, in general, be of interest for investigating experience-related changes in fundamental behavioral processes.     

How Food Controls Aggression in Drosophila

How animals use sensory information to weigh the risks vs. benefits of behavioral decisions remains poorly understood. Inter-male aggression is triggered when animals perceive both the presence of an appetitive resource, such as food or females, and of competing conspecific males. How such signals are detected and integrated to control the decision to fight is not clear. For instance, it is unclear whether food increases aggression directly, or as a secondary consequence of increased social interactions caused by attraction to food. Here we use the vinegar fly, Drosophila melanogaster, to investigate the manner by which food influences aggression. We show that food promotes aggression in flies, and that it does so independently of any effect on frequency of contact between males, increase in locomotor activity or general enhancement of social interactions. Importantly, the level of aggression depends on the absolute amount of food, rather than on its surface area or concentration. When food resources exceed a certain level, aggression is diminished, suggestive of reduced competition. Finally, we show that detection of sugar via Gr5a⁺ gustatory receptor neurons (GRNs) is necessary for food-promoted aggression. These data demonstrate that food exerts a specific effect to promote aggression in male flies, and that this effect is mediated, at least in part, by sweet-sensing GRNs.     

LIFE ON THE EDGE: STRESS SURVIVAL ADAPTATIONS IN SOUTHERN LIMIT MACROCYSTIS PYRIFERA POPULATIONS

In spite of subtropical temperatures often occurring at the Macrocystis pyrifera distributional southern limit in the Northern Hemisphere in Baja California, Mexico, the population has been successful at surviving and recovering from disturbance. To explore the physiological characteristics of southern limit M. pyrifera , microscopic and macroscopic phases of southern limit plants were tested for stress tolerance and compared to northern plant responses in three experiments. In spore development experiments, under constant temperature and nitrate stress treatments, spores from the southern-limit population survived higher temperatures, had greater survivorship, and were able to complete their life cycle at higher temperatures and lower nitrate concentrations than northern population spores. During stress-recovery treatments, where all microscopic stages were tested for recovery after stress shock exposure (temperature, nitrate and light stress), 96% of all haploid treatments showed some regrowth during recovery; haploid survival was not significantly affected by spore population origin, light, or nitrate. However, only 26% of diploid treatments showed regrowth and diploid survival was significantly affected by spore population origin; only the southern limit spores recovered after stress treatment exposure. In juvenile transplant experiments, under all conditions, juvenile plants originating from the southern-limit had higher tissue nitrogen content, greater frond addition rates, and higher photosynthetic efficiency than northern plants, particularly at higher temperatures. The physiological differences observed for the southern limit plants versus those from the north contributes to our understanding of how the southern limit plants can survive in an environment that might otherwise be considered extreme for the species.     

Effects of Bacillus thuringiensis var. israelensis on Target and Nontarget Organisms: a Review of Laboratory and Field Experiments

Since the discovery of Bacillus thuringiensis var. israelensis (Bti) in 1976, extensive literature has proved its efficacy to control mosquitoes and black flies, of which many species are known as important vectors of diseases or simply as pests of humans and animals. Since 1978, Bti has been used in many countries on all continents and numerous studies have been made on target mosquitoes and black flies, as well as nontarget organisms (NTO). This review analyses the results of 75 studies on these organisms covering approximately 125 families, 300 genera and 400 species. Different factors such as species, instar, feeding behaviour and environmental parameters (larval density, water temperature, suspended matter etc.) may drastically affect the efficacy of the Bti products. This is addressed in detail by reviewing the main factors affecting mosquitoes as well as black flies. The results of a wide range of laboratory and field experiments using different target and nontarget species, various preparations and formulations of Bti and different biotic or abiotic factors are present in the literature, making the data difficult to compare on a common basis. Our analysis shows that, under different application conditions, the effects of Bti on target and nontarget organisms may be hard to predict. Although Bti has been proclaimed to be relatively highly specific, some studies show that some NTO are affected either by single or repeated Bti treatments. Present use against black flies seems ecologically acceptable. High frequencies of application and/or overdosages against mosquitoes may result in some persistence of the toxin crystals and ultimately this may have adverse effects on the food web. A long-term study (published in 1998) in mosquito habitats has shown that intensive Bti treatments over three years did in fact produce an impact on the food web in wetlands. This raises questions, for the first time, on Bti environmental specificity. The importance of this impact is discussed and the alternatives for practical pest control are considered. Some modifications of Bti use against mosquitoes, guided by research, is probably the best of these alternatives.     

Multiple mating reduces male survivorship but not ejaculate size in the polygamous insect Stenomacra marginella (Heteroptera: Largidae)

In polygamous species, successful males should be able to inseminate multiple females, to defeat sperm from previous males, to avoid sperm displacement by other males, and to induce females to use his sperm during fertilization. Since resources are limited, adaptations to perform any of these functions may conflict with each other (and with other life-history traits) and trade-offs are expected to evolve. We studied if males of the polygamous true bug Stenomacra marginella face a trade-off between multiple mating and survivorship, by comparing the survivorship of virgin and multiply mated males. We also looked for physiological costs of ejaculate production by examining ejaculate production in consecutive matings in multiple mated males. Multiply mated males were able to produce ejaculates of similar size in up to six consecutive copulations but they had decreased survivorship in comparison with virgin males. There was no difference in survivorship between males mated three and six consecutive times, suggesting that the negative relation between survivorship and number of copulations is not linear. The decrease in survivorship seems to be a cost of mating and ejaculate production. This cost could favor the evolution of prudence in the allocation of resources to ejaculate production (e.g., cryptic male choice).     

The effects of tail autotomy on survivorship and body growth of Uta stansburiana under conditions of high mortality

We examined the effects of tail autotomy on survivorship and body growth of both adult and juvenile Uta stansburiana by directly manipulating tail condition. Tail loss decreased neither survivorship nor rate of body growth for individuals in two natural populations. Lack of an influence of tail loss on survivorship in these two populations may be the result of high mortality. Under high mortality any differential effects of tail loss will be lower than in populations facing lower mortality. Growth experiments in the laboratory demonstrated that, under conditions of minimal environmental variation and social interactions, there is no tradeoff between body growth and tail regeneration as has been suggested for other species of lizards. One possible reason for this difference is that U. stansburiana does not use the tail as a storage organ for lipids. The original and regenerated tails are composed mainly of protein. In general, any differential body growth between tailed and tailless individuals may be due to social interactions and not a diversion of limited energy into tail regeneration.     

Single and multigenerational responses of body mass to atmospheric oxygen concentrations in Drosophila melanogaster : evidence for roles of plasticity and evolution

Greater oxygen availability has been hypothesized to be important in allowing the evolution of larger invertebrates during the Earth’s history, and across aquatic environments. We tested for evolutionary and developmental responses of adult body size of Drosophila melanogaster to hypoxia and hyperoxia. Individually reared flies were smaller in hypoxia, but hyperoxia had no effect. In each of three oxygen treatments (hypoxia, normoxia or hyperoxia) we reared three replicate lines of flies for seven generations, followed by four generations in normoxia. In hypoxia, responses were due primarily to developmental plasticity, as average body size fell in one generation and returned to control values after one to two generations of normoxia. In hyperoxia, flies evolved larger body sizes. Maximal fly mass was reached during the first generation of return from hyperoxia to normoxia. Our results suggest that higher oxygen levels could cause invertebrate species to evolve larger average sizes, rather than simply permitting evolution of giant species.     

Short-day increases in aggression are inversely related to circulating testosterone concentrations in male Siberian hamsters (Phodopus sungorus)

Many nontropical rodent species display seasonal changes in both physiology and behavior that occur primarily in response to changes in photoperiod. Short-day reductions in reproduction are due, in part, to reductions in gonadal steroid hormones. In addition, gonadal steroids, primarily testosterone (T), have been implicated in aggression in many mammalian species. Some species, however, display increased aggression in short days despite basal circulating concentrations of T. The goal of the present studies was to test the effects of photoperiod on aggression in male Siberian hamsters (Phodopus sungorus) and to determine the role of T in mediating photoperiodic changes in aggression. In Experiment 1, hamsters were housed in long and short days for either 10 or 20 weeks and aggression was determined using a resident-intruder model. Hamsters housed in short days for 10 weeks underwent gonadal regression and displayed increased aggression compared to long-day-housed animals. Prolonged maintenance in short days (i.e., 20 weeks), however, led to gonadal recrudescence and reduced aggression. In Experiment 2, hamsters were housed in long and short days for 10 weeks. Half of the short-day-housed animals were implanted with capsules containing T whereas the remaining animals received empty capsules. In addition, half of the long-day-housed animals were castrated whereas the remaining animals received sham surgeries. Short-day control hamsters displayed increased aggression compared to either castrated or intact long-day-housed animals. Short-day-housed T treated hamsters, however, did not differ in aggression from long-day-housed animals. Collectively, these results confirm previous findings of increased aggression in short-day-housed hamsters and suggest that short-day-induced increases in aggression are inversely related to gonadal steroid hormones.     

Fitness effects of sex ratio response to host quality and size in the parasitoid wasp Spalangia cameroni

The parasitoid wasp Spalangia cameroni oviposited a greaterproportion of daughters in stable fly pupae than in house flypupae, even when I controlled for stable flies being smallerthan house flies. Sex ratio manipulation in response to hostquality has been modeled as being adaptive through an effectof host quality on the size and hence offspring production ofdaughters. 5. cameronis response to host species may insteadbe adaptive through an effect on larval survivorship, the developmenttime of daughters, and the size of sons. There was greater survivalof daughters than sons on stable flies. Controlling for hostsize, I found that development time of daughters was about 2%less on stable flies than on house flies. The decrease in developmenttime corresponds to a 2% increase in fitness as estimated byr, the intrinsic rate of increase, and is equivalent to abouta 9% increase in offspring production. Sons were about 2% largerfrom house flies than stable flies, which may increase offspringproduction by up to 3%. Host species had no consistent effecton size of daughters or development time of sons. In additionto the response to host species, mothers oviposited a greaterproportion of daughters in larger stable fly hosts. Whetherthis behavior is adaptive is unclear. Although offspring werelarger when they developed on larger stable flies, the rateof increase was less for daughters dian for sons. Effects ofstable fly size on offspring development time were negligible.     

Adrenal hormones mediate melatonin-induced increases in aggression in male Siberian hamsters (Phodopus sungorus)

Among the suite of seasonal adaptations displayed by nontropical rodents, some species demonstrate increased territorial aggression in short compared with long day lengths despite basal levels of testosterone. The precise physiological mechanisms mediating seasonal changes in aggression, however, remain largely unknown. The goal of the present study was to examine the role of melatonin, as well as adrenal hormones, in the regulation of seasonal aggression in male Siberian hamsters (Phodopus sungorus). In Experiment 1, male Siberian hamsters received either daily (s.c.) injections of melatonin (15 microg/day) or saline 2 h before lights out for 10 consecutive days. In Experiment 2, hamsters received adrenal demedullations (ADMEDx), whereas in Experiment 3 animals received adrenalectomies (ADx); control animals in both experiments received sham surgeries. Animals in both experiments subsequently received daily injections of melatonin or vehicle as in Experiment 1. Animals in all experiments were tested using a resident-intruder model of aggression. In Experiment 1, exogenous melatonin treatment increased aggression compared with control hamsters. In Experiment 2, ADMEDx had no effect on melatonin-induced aggression. In Experiment 3, the melatonin-induced increase in aggression was significantly attenuated by ADx. Collectively, the results of the present study demonstrate that short day-like patterns of melatonin increase aggression in male Siberian hamsters and suggest that increased aggression is due, in part, to changes in adrenocortical steroids.     

Testing a typology system of running waters for conservation planning in Hungary

The zebra mussel (Dreissena polymorpha) and its congener the quagga mussel (Dreissena rostriformis bugensis) are both invaders in freshwater, but have very different invasion histories, with zebra mussels attaining substantially faster rates of spread at virtually all spatial scales. However, in waterbodies where they co-occur, D. r. bugensis can displace D. polymorpha. To determine if the mechanisms for this displacement are associated with different survival and growth, we kept mussels in flow-through tanks for 289 days with two temperature regimes that mimicked the natural surface water (littoral zone) and hypolimnion conditions of Lake Erie. For the littoral zone regime, we used water directly from the surface of Lake Erie (range 4–25°C, average 11.9 ± 0.6°C). For the profundal zone treatment, Lake Erie surface water was chilled to about 6°C (range 5–8°C, average 6.2 ± 0.6°C) for the full duration of the experiment. For each of these temperature regimes, we used three replicate tanks with only zebra mussels present and three replicate tanks with only quagga mussels (150 ind./tank each), and three replicate tanks with both species (75 ind./tank of each species). Quagga mussels had higher survivorship and grew more than zebra mussels in all treatments. For both species, the size of the mussel entering the winter was critical for survivorship. Larger mussels had a higher survival over the winter in all treatments. For both species, there was a survivorship and growth tradeoff. In the warmer littoral zone treatment both species had higher growth, but lower survival than in the colder profundal zone treatment. Surprisingly, although quagga mussels outperformed zebra mussels, zebra mussel survivorship was better when they were faced with competition by quagga mussels than with just intraspecific competition. In addition, quagga mussels suffered size-specific mortality during the growing season only when facing interspecific competition with zebra mussels. Further experiments are needed to determine the possible mechanisms for these interspecific effects.     

The antenna of Drosophila as a 'love song' receptor

ABSTRACT. Behavioural experiments have demonstrated that the songs of Drosophila species are involved in courtship and aggression, and suggest that the songs are perceived via Johnston's organ. Direct electrophysiological evidence is here presented to show that the antenna of Drosphila is indeed capable of responding to the range of sounds produced by the flies.     

Developmental instability as phenodeviance in a secondary sexual trait increases sharply with thermal stress

Abstract We test for effects of thermal stress applied to pupal flies from Noumea (New Caledonia) and Taipei (Taiwan) on developmental instability (DI) in the male sex comb of Drosophila bipectinata, as well as on pre‐adult survivorship and adult body size. The temperature treatments were Low (25 °C), High (29 °C) and Variable (18 h at 29 °C, 6 h at 34 °C). Although the Variable treatment reduced survivorship and body size, absolute comb size and fluctuating asymmetry generally were invariant across treatments. In contrast, comb phenodeviance increased with stress in both populations. Phenodeviance in one comb segment (C2) increased sharply with stress, whereas phenodeviance in a second major segment (C1) also increased with stress but only in Noumea flies. A major conclusion is that phenodeviations induced in a secondary sexual trait reflect the developmental environment that also damages fitness components, a foundation stone of the hypothesis that expressions of DI reveal phenotypic quality in sexual selection.     

Effects of irradiation on Mediterranean fruit flies (Diptera: Tephritidae): emergence,survivorship, lure attraction,and mating competition

Irradiation of puparia in Mediterranean fruit fly, Ceratitis capitata (Wiedemann), sterile insect release programs can negatively affect adult fly performance. Emergence, survivorship, lure attraction, and mating competition tests were performed on irradiated and unirradiated Mediterranean fruit flies in Hawaii. Unirradiated flies of the Vienna-7 (tsl) strain had higher emergence, flight ability, and survivorship compared with irradiated flies. In general, unirradiated flies were more responsive to trimedlure, but this effect was not consistent for all strains at every age. Laboratory strains, of both unirradiated and irradiated flies, responded to trimedlure at a younger age than wild flies, which may be a result of inadvertent selection for decreased development time in laboratory-reared flies. Mating competition tests with irradiated and unirradiated flies showed no significant differences. Costs associated with the irradiation process and the development of alternative control techniques are discussed.     

Studying aggression in Drosophila (fruit flies)

Aggression is an innate behavior that evolved in the framework of defending or obtaining resources. This complex social behavior is influenced by genetic, hormonal and environmental factors. In many organisms, aggression is critical to survival but controlling and suppressing aggression in distinct contexts also has become increasingly important. In recent years, invertebrates have become increasingly useful as model systems for investigating the genetic and systems biological basis of complex social behavior. This is in part due to the diverse repertoire of behaviors exhibited by these organisms. In the accompanying video, we outline a method for analyzing aggression in Drosophila whose design encompasses important eco-ethological constraints. Details include steps for: making a fighting chamber; isolating and painting flies; adding flies to the fight chamber; and video taping fights. This approach is currently being used to identify candidate genes important in aggression and in elaborating the neuronal circuitry that underlies the output of aggression and other social behaviors.     

Virgin females compete for mates in the male lekking species Ceratitis capitata

Abstract. Aggressive behaviour occurring in intrasexual competition is an important trait for animal fitness. Although female intrasexual aggression is reported in several insect species, little is known about female competition and aggressive interactions in polygynous male lekking species. The interactions of female Mediterranean fruit flies, Ceratitis capitata (a male lekking species), with other females and mating pairs under laboratory conditions are investigated. Mature, unmated (virgin) females are aggressive against each other and against mating pairs, whereas immature females are not. Female aggression against other females decreases dramatically after mating; however, mated females maintain aggression against mating pairs. In addition, higher intrasexual aggression rates are observed for mature, virgin females than for virgin males of the same age. The results show that female aggressiveness is virginity related, suggesting female competition for mates. These findings have important implications for understanding the physiological aspects of a complex social behaviour such as aggression and should stimulate further research on female agonistic behaviour in male lekking mating systems.