种间专性协同进化关系对榕属分类的影响

榕与榕小蜂的专性共生系统为人们提供了研究生物协同进化规律和进化历史的完美范例,已成为国际上研究植物与昆虫相互作用的一个热点。这也给仅根据外部形态等传统方法进行的榕属经典分类带来了极大的影响和冲击。为了进一步研究榕属的系统与进化,介绍了该属分类学研究的历史、主要分类系统和存在的问题,并对今后榕属分类的发展趋势进行展望。     

The Influence of Previous Experience and Starvation on Aphid Feeding Behavior

It was hypothesized that (1) previous experience of aphids on a host plant leads to differences in their feeding behavior relative to aphids without previous experience on it and that (2) a change in the physiological state of the aphid modifies their experience-induced behavior. Using electronic recording, the feeding behavior of the aphid Sitobion fragariae (Walker) on wheat Triticum aestivum L. and oat Avena sativa L. was examined, comparing aphids with or without previous experience on a given host and with or without a period of starvation before assessing probing behavior. All comparisons were performed within a single aphid clone to minimize the effect of genetic variation. Feeding behavior on wheat was significantly affected by previous experience and starvation. The effect of previous experience interacted with the host plant where feeding behavior was tested. Aphids feeding on wheat following previous experience on wheat showed a longer time and a higher number of pathway activities and less time in waveform F (i.e., mechanical stylet work and penetration difficulties) than did aphids feeding on wheat after a previous experience on oat. No differences in the time from the beginning of the recording until the first salivation into the sieve elements were found. When aphids were subjected to a period of starvation, the time devoted to xylem ingestion increased compared with that of nonconstrained aphids. These results are discussed in terms of factors affecting foraging decisions.     

Social Cues Accelerate Reentrainment of Circadian Rhythms in Diurnal Female Octodon Degus (Rodentia-Octodontidae)

Previous studies paired diurnal Octodon degus undergoing/phase advances (phase-shifters) with those entrained to a light-dark (LD) cycle (donors). Results included opposite outcomes of male and female social cues on resynchronization following 6-h advances in females, but no effect of social cues on male resynchronization. The first experiment determined if social cues could influence resynchronization rates of circadian rhythms in male and female degus following a 6-h phase delay of the LD cycle. Female phase-shifters resynchronized temperature and activity rhythms 20-35% faster when housed with either entrained (donor) females or males compared with females housed alone. No significant differences in resynchronization rate for phase-shifting males existed between test conditions. This experiment extends the previous finding that females, but not males, respond strongly to donor cues to increase resynchronization rates in the presence of light. A second experiment determined that accelerated resynchronization rates of female phase-shifters housed with female donors were due to social cues directly affecting the circadian system rather than the result of social masking. On the day following resynchronization with or without a female donor present, phaseshifters were transferred individually to constant conditions (DD). The temperature and activity rhythms of female phase-shifters free-ran from the point at which resynchronization occurred for both the control and experimental females. Thus, social cues accelerate true reentrainment, not masking, of the circadian system in the presence of a LD cycle in female degus. Donor cues from females enhance reentrainment after advances and delays, but the effect of male donor cues is dependent on the direction of the phase shift.     

Social Cues Accelerate Reentrainment of Circadian Rhythms in Diurnal Female Octodon Degus (Rodentia-Octodontidae)

Previous studies paired diurnal Octodon degus undergoing/phase advances (phase-shifters) with those entrained to a light-dark (LD) cycle (donors). Results included opposite outcomes of male and female social cues on resynchronization following 6-h advances in females, but no effect of social cues on male resynchronization. The first experiment determined if social cues could influence resynchronization rates of circadian rhythms in male and female degus following a 6-h phase delay of the LD cycle. Female phase-shifters resynchronized temperature and activity rhythms 20–35% faster when housed with either entrained (donor) females or males compared with females housed alone. No significant differences in resynchronization rate for phase-shifting males existed between test conditions. This experiment extends the previous finding that females, but not males, respond strongly to donor cues to increase resynchronization rates in the presence of light. A second experiment determined that accelerated resynchronization rates of female phase-shifters housed with female donors were due to social cues directly affecting the circadian system rather than the result of social masking. On the day following resynchronization with or without a female donor present, phaseshifters were transferred individually to constant conditions (DD). The temperature and activity rhythms of female phase-shifters free-ran from the point at which resynchronization occurred for both the control and experimental females. Thus, social cues accelerate true reentrainment, not masking, of the circadian system in the presence of a LD cycle in female degus. Donor cues from females enhance reentrainment after advances and delays, but the effect of male donor cues is dependent on the direction of the phase shift.     

Experience Based Use of Landmark and Vector Based Orientation During Homing by the Ant Formica cunicularia (Hymenoptera: Formicidae)

Homing paths of Formica cunicularia foragers from an artificial food reward were analyzed on a familiar terrain and in displacement experiments on a familiar and an unfamiliar terrain. Foragers were tested either when relatively new to a foraging route (untrained group) or after a day’s experience with it (trained group). Untrained foragers followed direct homing paths to the nest site when tested in the familiar terrain but followed tortuous paths when displaced to the unfamiliar terrain. Trained foragers behaved similarly to untrained ones when tested from the food reward to the nest site in the familiar terrain but their behavior changed in displacements. Irrespective of the familiarity of the displacement site, these foragers followed paths taking them to the expected nest sites. The results showed that foragers did not rely on chemical cues for homing and revealed that untrained foragers disregarded path integration and were directed to the nest site when it is in their visual panorama. On the contrary, trained foragers may have relied on path integration on familiar and unfamiliar terrain. The results also demonstrated that experience greatly affected the preferential use of visual and vector based cues by foragers during homing.     

Influence of Conspecific Presence, Experience, and Host Quality on Oviposition Behavior and Clutch Size Determination in Anastrepha ludens (Diptera: Tephritidae)

Females of some insect species adjust the number of ovipositions and clutch size adaptively depending on conspecific density and probably experience. In a series of three experiments, we examined the effect of the presence of conspecifics, host quality, and oviposition experience on oviposition behavior and clutch size determination by females of the polyphagous fruit fly Anastrepha ludens (Diptera: Tephritidae). In the first experiment, we determined that grouped (eight females per cage) A. ludens females tended to visit and oviposit in more hosts than did solitary females probably as a result of stimulation by the presence of conspecifics. We also determined that females with previous oviposition experience visited and oviposited in more hosts than inexperienced ones. Importantly, when females were grouped, we observed significantly more landings on unoccupied hosts (i.e., devoid of flies) than on occupied ones (i.e., with at least one fly on it). However, oviposition experience, and not female density, was the most important factor affecting clutch size. Naive females deposited larger egg clutches than experienced ones. In the second experiment, we found that oviposition experience and host quality (i.e., clean fruit or fruit covered with a host marking pheromone [HMP] extract), influenced clutch size and the decision of females to defend or not defend the host. Clutch size and number of fights were greater on clean than on HMP-marked hosts. In the third experiment, we observed that host quality (i.e., size) played a significant role with regard to the number of female fights, host marking behavior, and clutch size. Specifically, females fought and dragged their aculeus longer on small- and medium-sized hosts than on large ones. But this behavior varied according to whether females were kept alone or grouped. Clutch size was greatest in the largest hosts. Considering all the above, we believe that the observed increase in ovipositional activity by grouped A. ludens females can be attributed to competition through mutual interference and not social facilitation as has been reported in other tephritid species.     

Foraging Efficacy of a Larval Parasitoid in a Cotton Patch: Influence of Chemical Cues and Learning

Plant-herbivore chemical signals and behavioral plasticity may enhance parasitoid host-foraging efficacy in the field; however, no studies have quantified the potential benefits from these factors under field-type conditions. The effect of plant-herbivore signals and learning on the foraging efficacy of Microplitis croceipes was quantified by directly observing and recording total and sequential duration of various foraging behaviors relative to 5 randomly placed herbivore-damaged and host-infested cotton plants and 20 undamaged and non-host-infested plants. Microplitis croceipes spent significantly more time searching (flying and antennation) on host infested versus uninfested plants. Antennation time was significantly and negatively correlated with successive host stings. Contrary to expectations of increased duration, flight time remained constant throughout the foraging bout, which may indicate that there was some learning associated with flight. These results suggest that plant-herbivore chemical signals and learning enhances the foraging efficacy of M. croceipes.     

Age and Sex-Based Differences in the Use of Prey Sensory Cues in Wolf Spiders (Araneae: Lycosidae)

Differences in foraging patterns mediated by sensory cues were examined between adult and juvenile male and female wolf spiders (Schizocosa rovneri; Lycosidae). Patch residence time for thirty-one spiders were tested among juveniles and adults in artificial foraging patches. Patches varied in sensory information provided by live prey (crickets) as follows: visual stimuli alone; vibratory stimuli alone; visual and vibratory stimuli together; and control (no stimuli). Spiders moved between patches for one hour, but could not feed. Adult Schizocosa rovneri use primarily visual information to determine patch residence time, but juveniles use vibratory cues as well. Significant age and sex-based differences in the use of sensory cues suggest that observed divergent foraging strategies are partly due to the use of different perceptual cues in prey detection.     

Orientation Cage and Release Experiments with Migratory Wheatears (Oenanthe oenanthe) in Scandinavia and Greenland: The Importance of Visual Cues

Migratory orientation of Scandinavian and Greenland wheatears was recorded during the autumn migration periods of 1988 and 1989. Orientation cage tests were conducted under clear sunset skies, to investigate the importance of different visible sky sections on orientation performance. In addition, wheatears were released under clear starry skies and under total overcast to examine the orientation of free-flying birds. The following results were obtained:

• 1 Wheatears tested with a restricted visible sky section (90° centered around zenith) in orientation cages, showed a mean orientation towards geographic W/geomagnetic NW (Greenland) and towards geographic and magnetic WNW-NW (Sweden). These mean directions are clearly inconsistent with the expected autumn migration directions, SW-SSW in Scandinavia and SE in Greenland, as revealed by ringing recoveries for the two populations.
• 2 When the birds were allowed a much more extensive view of the sky, almost down to the horizon (above 10° elevation), Scandinavian wheatears chose headings in agreement with ringing data. Greenland birds were not significantly oriented.
• 3 Release experiments under clear starry skies resulted in mean vanishing directions in good agreement with ringing data from both sites. Greenland wheatears released under total overcast showed a similar orientation as under clear skies, indicating that a view of the stars may not be of crucial importance for selecting a seasonally accurate migratory direction.

The results suggest that an unobstructed view of the sky, including visual cues low over the horizon, is important, possibly in combination with geomagnetic cues, for the orientation of migratory naive wheatears. Furthermore, the birds showed remarkably similar orientation responses in Greenland and Scandinavia, respectively, indicating that they use basically the same orientation system, despite considerable differences in visual and geomagnetic orientation premises at the two different geographic and magnetic latitudes.     

Effects of Acute and Chronic Exposure to Predatory Cues on Spatial Learning Capabilities in the Subterranean Rodent Ctenomys talarum (Rodentia: Ctenomyidae)

Spatial learning, the cognitive ability that allows animals to efficiently locate food, partners or avoid predators, can be affected by various factors such as stress. Stressors represent a threat to the animal's homeostasis and trigger a set of physiological and behavioral changes known as stress response. The herbivorous rodent Ctenomys talarum (tuco‐tuco) inhabits underground complex gallery systems. Dispersal as well as food collection occurs on the surface, where they are exposed to terrestrial and aerial predators. We evaluated the effect of a natural stress factor, predation, on spatial learning of C. talarum. Additionally, we collected data on neutrophil/lymphocyte ratio, hematocrit, blood glucose, and plasma cortisol to assess physiological levels of stress. The acute exposure to direct cues indicating the presence of a predator (immobilization + cat urine) increased the time needed to reach the goal in a longitudinal labyrinth. The number of errors also increased albeit the difference was only marginally significant. The chronic exposure to predator cues (immobilization, cat urine, and cat fur) presented in an unpredictable order significantly increased both time and errors committed by tuco‐tucos during learning trials in the labyrinth. Results show the strong impact of predatory stress on the spatial abilities of C. talarum, a key cognitive process involved in most of their vital activities. They also highlight the importance of investigating wild species as regards the development of antipredatory defensive behaviors that contribute both to the avoidance of deleterious consequences of a direct attack and the influence on food search and mate localization performance.     

Orientation of crustacean burrows in the Bay of Panzano (Gulf of Trieste, Northern Adriatic Sea)

Three burrow systems produced by Jaxea nocturna and one cluster of burrows produced by Upogebia pusilla where investigated in the Bay of Panzano, Northern Adriatic Sea, to determine preferred orientations. The distributions of dip directions differ between both producers. Steep shafts into the consolidated mud followed by large, shallowly inclined tunnels oriented in a manner similar to a spiral are characteristic for Jaxea nocturna burrows. In contrast, the Y-shaped burrows of Upogebia pusilla have entrance shafts that are less steep which are connected by a near-horizontal section, where a blind ending tunnel branches. Orientations of the dominating, shallowly inclined burrow parts are not randomly distributed in either Jaxea or in Upogebia burrows. Three preferred axial orientations with almost equal proportions in each direction are characteristic for the Jaxea burrow from the centre of the bay. This system transforms to a squared structure near the coast, where one direction parallels the shoreline and the other is oriented at right angles to the coast. The latter orientation demonstrates significant constancy in all investigated burrows. Upogebia burrow clusters coincide with the three preferred orientations of the Jaxea burrow from the bay centre, but the proportions of the directions are unequal. Burrow segments connecting the steep entrance shafts in Upogebia are oriented almost parallel to the shoreline, whereas at right angles to the coast the flat blind-ending tunnels incline towards the open sea. The study shows a strong coincidence between empirical and theoretical distributions of the dominating orientations in both species.     

Orientation of crustacean burrows in the Bay of Panzano (Gulf of Trieste, Northern Adriatic Sea)

Three burrow systems produced by Jaxea nocturna and one cluster of burrows produced by Upogebia pusilla where investigated in the Bay of Panzano, Northern Adriatic Sea, to determine preferred orientations. The distributions of dip directions differ between both producers. Steep shafts into the consolidated mud followed by large, shallowly inclined tunnels oriented in a manner similar to a spiral are characteristic for Jaxea nocturna burrows. In contrast, the Y-shaped burrows of Upogebia pusilla have entrance shafts that are less steep which are connected by a near-horizontal section, where a blind ending tunnel branches. Orientations of the dominating, shallowly inclined burrow parts are not randomly distributed in either Jaxea or in Upogebia burrows. Three preferred axial orientations with almost equal proportions in each direction are characteristic for the Jaxea burrow from the centre of the bay. This system transforms to a squared structure near the coast, where one direction parallels the shoreline and the other is oriented at right angles to the coast. The latter orientation demonstrates significant constancy in all investigated burrows. Upogebia burrow clusters coincide with the three preferred orientations of the Jaxea burrow from the bay centre, but the proportions of the directions are unequal. Burrow segments connecting the steep entrance shafts in Upogebia are oriented almost parallel to the shoreline, whereas at right angles to the coast the flat blind-ending tunnels incline towards the open sea. The study shows a strong coincidence between empirical and theoretical distributions of the dominating orientations in both species.     

Sociality and the Rate of rDNA Sequence Evolution in Wasps (Vespidae) and Honeybees (Apis)

Sequence data of mitochondrial 16S ribosomal DNA (mt-rDNA) and nuclear 28S ribosomal DNA (nuc-rDNA) were compared in two honeybee species (Apis mellifera and Apis dorsata) and a selection of 22 wasp species (Vespidae) with different levels of sociality. The averge substitution rates in mt-rDNA and nuc-rDNA were almost-equal in solitary species. In species with larger nests, however, the difference between the nuclear and the mitochondrial substitution rate significantly increased. The average substitution ratio, ψ (nucleotide substitutions in mt-rDNA/nucleotide substitutions in nuc-rDNA) was 1.48 ± 0.12 (SE) among the solitary Eumeninae, 3.70 ± 0.15 among five primitive social Stenogastrinae species, 3.24 ± 0.20 among five Polistinae species, 5.76 ± 0.33 among nine highly eusocial Vespinae, and 12.7 in the two Apis species. The high egg-laying rate and the effective population size skew between the sexes may contribute to the rise of the substitution ratio in the highly eusocial species. Drift and bottleneck effects in the mitochondrial DNA pool during speciation events as well as polyandry may further enhance this phenomenon. Received: 12 January 1998 / Accepted: 28 April 1998     

非嗜食寄主气味蛹期处理对不同寄主来源美洲斑潜蝇雌成虫嗅觉定向的影响

测定用菜豆和丝瓜饲养的美洲斑潜蝇Liriomyza sativae Blanchard蛹暴露于用正己烷稀释的番茄或南瓜叶片挥发性抽提物(TLE或PLE)72h后,羽化雌成虫在Y形嗅觉仪中对TLE或PLE的定向率、平均反应时间以及对正己烷的定向率和平均反应时间,比较幼虫寄主经历和寄主气味对美洲斑潜蝇成虫嗅觉记忆的影响。结果表明,蛹期使用非嗜食寄主植物气味处理,能增加美洲斑潜蝇羽化雌成虫对非嗜食寄主气味的定向;幼虫期的寄主经历对处理后的成虫嗅觉定向也产生影响,饲喂丝瓜的美洲斑潜蝇蛹期处理后,羽化雌成虫显著偏向选择处理气味而非正己烷,饲喂菜豆的美洲斑潜蝇对南瓜的气味也表现出同样的特点,但对番茄的气味则相反,即使处理后对番茄气味的选择比率增加,仍然偏向选择正己烷。     

Experience of natural prey and feeding efficiency in three-spined sticklebacks (Gasterosteus aculeatus L.)

The possibility that efficiency of feeding on different prey in three-spined sticklebacks ( Gasterosteus aculeatus L.) increases by experience of such items was investigated experimentally by comparing sticklebacks reared with only non-moving prey (frozen brine shrimps) with sticklebacks caught in the wild from localities where a wide range of prey species was available. Fish without experience of natural food scored less well than those from the wild on various components of foraging and so had reduced feeding efficiency. However, feeding efficiency in naive fish improved after just a few days of experience with natural prey. Such experience-induced changes in foraging efficiency will alter the profitability of prey items for individual fish.     

美洲斑潜蝇蛹期化学环境对成虫嗅觉定向的影响

将美洲斑潜蝇Liriomyzasativae Blanchard蛹分别暴露于3种植物挥发物-芳 樟醇、β-子丁香烯和3- 己烯-1-醇,羽化后用Y形嗅觉仪测定雌成虫对相应挥发物的定向反 应。蛹期暴露于3-己烯-1-醇后羽化的雌成虫趋向该化合物的比率（37.7%）和平均反应时 间（21.5 s）,与对照组（30.4%,35.0 s）差异不显著;β-子丁香烯处理组雌成虫选择 该化合物的比率（46.2%）与对照组（42.0%）差异不显著,但平均反应时间（21.0 s）却显著短于对照组（41.5 s）。蛹期经芳樟醇处理后雌成虫选择该化合物的比率（52.9%）显著 高于对照组（28.4%）,平均反应时间（19.5 s）也显著短于对照组（34.5 s）。以芳樟醇为处理化合物的进一步研究表明,蛹发育早期是诱导成虫产生定向反应的敏感时期。将1、3、5 、7 日龄蛹分别暴露于芳樟醇48 h后,只有1日龄组蛹羽化的雌成虫趋向芳樟醇的比率（54%） 显著高于对照组（26%）;但1、3、5 日龄组蛹羽化的雌成虫对芳樟醇的平均反应时间均显著 短于对照组。此外,蛹期暴露于芳樟醇的持续时间也影响雌成虫对该化合物的定向反应,2日龄 蛹分别暴露于芳樟醇24、48、72、96和120 h后,只有处理时间大于72 h的雌成虫选择该化合物 的比率才显著高于对照组;但所有处理组羽化雌成虫的平均反应时间均显著低于对照组。由此 推断,美洲斑潜蝇蛹期经历的化学环境会影响成虫的嗅觉定向反应。     

Influence of Social Context on the Use of Blended and Graded Facial Displays in Chimpanzees

Our understanding of social communication and emotional behavior in nonhuman primates has advanced considerably through research over the past half century. Chimpanzee facial displays have typically been described as highly graded communicative signals, but we propose an additional distinction: blended displays. They appear to be morphologically and acoustically similar to the expressions in 2 prototypical/parent categories. We describe the facial and vocal communicative repertoire of chimpanzees and examine how they use graded and blended signals in different social contexts. Data from behavioral observations revealed that they used facial displays differently depending on the social context. Specifically, the variability can be explained by 7 factors representing nervousness and distress, agonism, contact reassurance, excitement, greetings, play, and vigilance. Additionally, the use of blended displays was not simply divided between the contexts that elicited the parent types, nor were they used in totally unique contexts. Instead, the data showed that the contextual use of blended displays is primarily correlated with the social contexts that elicited only one of the parent expressions. Thus, the blended displays appeared to reflect conflicting internal motivational states in the sender, instead of expressing features of the external environment. We proffer several possible explanations for how the blended signals may be interpreted by receivers and why they would be contextually associated with only one parent group.     

Influence of previous mating experience on future mating success in maleLucilia cuprina (Diptera: Calliphoridae)

Receptive and virginL. cuprina females were placed with a virgin male and an experienced male that had mated between one and five times previously. The experienced males secured significantly more matings than virgin males. Males with previous matings also gained experience in competing with males. Males directed mating attempts at each other, seemingly in the context of intrasexual competition. Experienced males directed more mating attempts at virgin males than vice versa. As their number of previous matings increased, experienced males made the first mating attempt at females more often and directed more mating attempts at females compared with virgin males. Females did not actively discriminate against experienced males, even though the proportion of matings secured on the first attempt by experienced males declined with increasing mating experience. Alternative behavioral explanations are discussed.     

Orientation of flying male Anobium punctatum (Coleoptera: Anobiidae) to sex pheromone: separating effects of visual stimuli and physical barriers to wind

Abstract. Flying male woodworm, Anobium punctatum , are able to locate a source of female pheromone offered as a point source but land more quickly if the same odour source is presented on a model 'tree' (cylinder). We tested the response to a two-dimensional black 'tree' marked as a silhouette on the upwind screen of the wind tunnel, with or without a hidden upwind barrier to the wind. The beetles responded equally to pheromone presented on a visual silhouette with wind barrier and to the hidden upwind wind-barrier alone, suggesting that the wind-barrier itself has a significant effect on insect orientation and landing: visual stimuli do not add to the landing response if the barrier is present. However, without the barrier upwind, landing was faster when the visual stimulus was present than without. This is the third forest-living species to be shown to respond in this way, which suggests this phenomenon may be more widespread. The possibility that this phenomenon is present in other, non-forest, phytophagous insects, their predators and parasitoids is discussed.