昆虫种子捕食与蒙古栎种子产量和种子大小的关系

为了了解昆虫种子捕食者在栎类种群更新中的作用,于2006年秋季,在黑龙江省伊春市带岭区东方红林场研究了昆虫对蒙古栎Quercus mongolica在参园和东山两个种群的种子蛀食情况及其与蒙古栎种子产量和种子大小的关系。结果表明：(1)在参园和东山两个林分内,蒙古栎种子雨动态非常相似,种子雨成分中完好种子的平均密度仅为3.2±4.1个/m²（参园）和1.7±2.8个/m²（东山）,分别仅占种子产量的4.0％和3.2％,而虫蛀种子和败育种子的比例均在38%以上,以虫蛀种子的比例最高,分别为58.2％和57.7％;(2)柞栎象Curculio arakawai是蛀食蒙古栎种子的主要昆虫种类,在虫蛀种子中所占比例高达96.8％（参园）和97.1％（东山）,且象甲蛀食种子中所含虫卵数与种子大小有关,即种子越大,所含象甲的虫卵数就较多。本研究的结果说明2006年蒙古栎成熟种子多遭遇虫蛀,导致完好种子的数量极低,因而可能成为限制蒙古栎种群更新的重要因素。     

Latitude, seed predation and seed mass

Aim We set out to test the hypothesis that rates of pre‐ and post‐dispersal seed predation would be higher towards the tropics, across a broad range of species from around the world. We also aimed to quantify the slope and predictive power of the relationship between seed mass and latitude both within and across species. Methods Seed mass, pre‐dispersal seed predation and post‐dispersal seed removal data were compiled from the literature. Wherever possible, these data were combined with information regarding the latitude at which the data were collected. Analyses were performed using both cross‐species and phylogenetic regressions. Results Contrary to expectations, we found no significant relationship between seed predation and latitude (log₁₀ proportion of seeds surviving predispersal seed predation vs. latitude, P = 0.63; R² = 0.02; n = 122 species: log₁₀ proportion of seeds remaining after postdispersal seed removal vs. latitude, P = 0.54; R² = 0.02; n = 205 species). These relationships remained non‐significant after variation because of seed mass was accounted for. We also found a very substantial (R² = 0.21) relationship between seed mass and latitude across 2706 species, with seed mass being significantly higher towards the tropics. Within‐species seed mass decline with latitude was significant, but only about two‐sevenths, as rapid as the cross‐species decline with latitude. Results of phylogenetic analyses were very similar to cross‐species analyses. We also demonstrated a positive relationship between seed mass and development time across ten species from dry sclerophyll woodland in Sydney (P < 0.001; R² = 0.77; Standardized Major Axis slope = 0.14). These data lend support to the hypothesis that growing period might affect the maximum attainable seed mass in a given environment. Main conclusions There was no evidence that seed predation is higher towards the tropics. The strong relationship between seed mass and latitude shown here had been observed in previous studies, but had not previously been quantified at a global scale. There was a tenfold reduction in mean seed mass for every c. 23° moved towards the poles, despite a wide range of seed mass within each latitude.     

槲栎种子雨进程中昆虫的捕食特征

为了了解昆虫捕食与槲栎Quercus aliena种子大小和产量的关系以及在栎林更新中的作用,于2008和2009年秋季,分别在暖温带伏牛山系天池山国家森林公园内研究了两个年度槲栎的种子雨过程及昆虫捕食特征。结果表明：（1）种子雨过程从8月中下旬到9月底约经历40 d,存在高峰期且时段较明显,高峰期下落种子量分别占全部种子雨量的78.13%（2008年）和75.91%（2009年）;（2）槲栎的种子雨强度年间存在较大差异,2008年明显小于2009年（两年分别为31.75±16.65粒/m²和51.92±29.26粒/m²）,但2009年的橡子明显比2008年小（两年分别为1.94±0.61 cm³和2.46±0.57 cm³）;（3）种子雨构成比例在两个年份间存在差异,完好种子的量分别为59.05%（2008年）和36.12%（2009年）,虫蛀率在大量结实的2009年显著提高;（4）虫蛀橡子（2.29±0.42 cm³）显著大于完整橡子（1.59±0.32 cm³）,且虫蛀种子中所含虫卵数与种子大小显著正相关,昆虫有选择大种子产卵寄生的偏好。本研究的结果说明,昆虫对槲栎大种子有寄生选择偏好;大部分槲栎种子遭遇象甲虫蛀而降低生命活力,这可能是影响槲栎林更新的主要因素之一。     

Seed size predicts global effects of small mammal seed predation on plant recruitment

Recent studies demonstrate that by focusing on traits linked to fundamental plant life‐history trade‐offs, ecologists can begin to predict plant community structure at global scales. Yet, consumers can strongly affect plant communities, and means for linking consumer effects to key plant traits and community assembly processes are lacking. We conducted a global literature review and meta‐analysis to evaluate whether seed size, a trait representing fundamental life‐history trade‐offs in plant offspring investment, could predict post‐dispersal seed predator effects on seed removal and plant recruitment. Seed size predicted small mammal seed removal rates and their impacts on plant recruitment consistent with optimal foraging theory, with intermediate seed sizes most strongly impacted globally – for both native and exotic plants. However, differences in seed size distributions among ecosystems conditioned seed predation patterns, with relatively large‐seeded species most strongly affected in grasslands (smallest seeds), and relatively small‐seeded species most strongly affected in tropical forests (largest seeds). Such size‐dependent seed predation has profound implications for coexistence among plants because it may enhance or weaken opposing life‐history trade‐offs in an ecosystem‐specific manner. Our results suggest that seed size may serve as a key life‐history trait that can integrate consumer effects to improve understandings of plant coexistence.     

The role of pre- and post-dispersal seed predation in determining total seed loss

Most seed predation studies focus on either pre- or post-dispersal predation and may therefore underestimate the role of predation in regulating plant populations. We therefore estimated total seed predation of an invasive tree, mesquite (Leguminoseae: Prosopis spp.), by examining the entire seed pool from tree to seed bank. The spatio-temporal dynamics of total seed predation was examined by sampling across its Australian distribution and through time. The main predator was a host-specialist multivoltine beetle, Algarobius prosopis L. (Bruchidae), previously introduced as a biocontrol agent. Seed predation exceeded 20% in all seed stages (in pods on and off the tree, and seeds within woody endocarps (capsules) and free seeds on and in the ground) but was consistently highest in capsules on the ground (up to 90%). Pre-dispersal predation contributed little. Total seed predation rates were primarily determined by predation rates on the most persistent seed stage, in this case fallen pods if only pods are considered and seeds in capsules for the total seed pool. This pattern was consistent across the surveyed taxa, regions, years and seasonally. Predation rate was relatively unaffected by seed density, potentially because densities were always low (<150 seeds m⁻²). Average total seed predation within a region reached 55%, but we conclude that any population regulation of mesquite by seed predation will principally be through reduced seed bank persistence. Our results highlight the need to consider the entire seed pool, especially the often cryptic and overlooked long-lived stages, when determining seed loss to predation and its likely population consequences.     

植物种子大小与幼苗生长策略研究进展

种子大小和幼苗更新对策是植物生活史策略的重要组成部分.本文从不同侧面综述了当前国内外关于种子大小和幼苗之间关系的生态学研究,包括:种子大小对萌发出土的影响,种子大小与幼苗形态生长特性之间的关系,种子大小与幼苗存活、竞争能力之间关系等,并对今后的相关研究进行了展望.在不同的微环境和植被类型下,种子大小与幼苗生长之间的关系可能有所差异;种子大小对植物幼苗生长的影响导致种子大小不同的植物对植被幼苗更新补充的贡献不同;种子大小与幼苗生活史策略关系在大尺度的群落空间水平上和小尺度的物种间与物种内的研究对于天然植被的更新恢复研究具有重要意义.     

Offspring allocation in externally ovipositing fig wasps with varying clutch size and sex ratio

The simultaneous optimization of clutch size and sex ratio isa tricky problem. Unless parameters such as host size or fecundityexist to pin down the optimal clutch size, this problem remainselusive to analytical analysis. This is because the fitnesslandscape with respect to clutch size and sex ratio does nothave one single evolutionarily stable peak toward which thepopulation can evolve. To solve this problem, I used a computeremulation to optimize both clutch size and sex ratio using externallyovipositing fig wasps as a model taxon. The simulation approachallows the use of integer numbers of eggs rather than assumingthat females can produce any sex ratio between 0 and 1. Whenfemales have no information about the patches on which theyoviposit, they produce either large clutches with a strong femalebias or clutches of a single male egg. When females have completeknowledge of their oviposition site, a set of conditional substrategiesis evolutionarily stable. Again, these substrategies are eitherlarge clutches with a female bias or dutches consisting of asingle male egg. This dichotomous oviposition pattern resultsin unrelated males sharing a fig, a condition conducive to theevolution of fatal fighting. Selection on female ovipositionstrategies may therefore be an important driving force behindhigh levels of fighting observed between male fig wasps.     

Effects of seed predation by ants on Mediterranean grassland related to seed size

Questions: 1. Do harvester ants (Messor barbarus) promote seed mortality in Mediterranean grassland?; 2. Is this effect greater in large‐seeded species? Location: Central Spain. Methods: We established an ant‐exclusion experiment of five circular (1.5 m diameter) plots from where ants were excluded during one year, along with ten control plots. We recorded the seed bank of all species in the plots both before and after the treatment. The effect of seed length and weight was analysed after transforming data into phylogenetically independent contrasts, and alternatively by dividing the species data set into morphological groups. Results: Longer and heavier seeded species significantly increased in the seed banks under the exclusion treatment, although ants did not significantly modify overall seed densities. Conclusions Although the ants do not collect large numbers of seeds, they differentially affect the composition of the seed banks by selecting the longest or heaviest seeds, or both. The persistence of this short‐term effect in the seed bank may result, over a number of years, in the system evolving towards a predominance of small‐seeded annuals, congruent with the species composition actually observed in Mediterranean grasslands.     

Parent-offspring conflict over clutch size

Summary We present a model for sexually-reproducing diploids in which a female can produce a variable (generally large) clutch size, where the sibs then compete over some fixed resource, and where certain offspring use siblicide to reduce the primary clutch/brood size created by the mother. Where siblicide involves neither direct energy loss (e.g. fighting cost) nor gain (e.g. cannibalism) to an offspring, the optimal clutch size for an offspring can differ from the optimum for the mother, i.e. there can be parent-offspring conflict over clutch size. The magnitude of this evolutionary conflict (measured in terms of difference between clutch size optima) increases with multipaternity of the brood and with the steepness of the initial decline in offspring survivorship (through sib-competition as further offspring are added to the brood). However, the disparity in clutch size optima may not be great. Where the integer clutch size optima are the same, there will clearly be no conflict. Where this differs, resolution of the evolutionary conflict could involve much apparent behavioral conflict, commonly manifest as siblicidal aggression.The ESS (evolutionarily stable strategy) for such a game will depend upon the direct costs and benefits of siblicide, as well as on the indirect costs to sibs via relatedness. If the only costs of siblicide arise through relatedness, then offspring will win in the sense that the eventual clutch size will match the offspring optimum. Whether or not the mother will produce this clutch size depends on the mechanism controlling siblicide. A siblicidal ESS will occur when offspring are programmed to kill a fixed number/proportion of a brood (victim-based siblicide), but not if programmed to reduce the sibship to the offspring optimum (survivor-based siblicide). With survivor-based siblicide, the mother can do no better than to lay the offsprings' optimal clutch size.     

Empty seeds reduce seed predation by birds in Juniperus osteosperma

Utah juniper (Juniperus osteosperma) is one of many plant species that produce large numbers of fruits containing parthenocarpic or otherwise empty or inviable seeds. We tested the hypothesis that production of empty fruits in this species results in reduced levels of predation on fertile seeds. In a population in west-central Utah, we estimated the proportion of fruits with filled seeds in trees suffering high levels of fruit destruction by the seed-eating bird Parus inornatus and in neighbouring trees similar in crown and fruit-crop size but suffering negligible predation. We found that the heavily attacked trees had higher proportions of filled seeds. Thus, juniper may benefit from producing fruits that contain no offspring. This is the first study to demonstrate that empty seeds may reduce predation by vertebrate seed eaters and the first to demonstrate discrimination based on seed filling at the level of whole plants.     

种子特征和结实量对啮齿动物取食和扩散种子行为的影响

为了深入了解啮齿动物在不同种子丰富度条件下对不同大小和单宁含量种子的觅食行为策略及其与植物种群更新的关系,在宁夏六盘山区的华北落叶松人工林,研究了不同大小和单宁含量[0%Tannin(T)、2%T、8%T和15%T]的人工种子在模拟结实小年和结实大年对啮齿动物取食和扩散行为的影响.结果表明: 啮齿动物消耗种子速度在结实小年更快,结实大年的种子消耗速度相对缓慢. 种子就地取食率(ISPR)在不同结实年份间无显著差异,扩散后取食率(PRAD)在结实小年显著高于结实大年,但前者的扩散后贮藏率(HRAD)显著低于后者;种子扩散后的取食距离(PDAD)和贮藏距离(HDAD)在结实小年均显著大于结实大年.在结实小年,大种子的PDAD和HDAD均大于小种子,前者在不同大小种子间均差异显著,而后者仅在2%T和15%T的不同大小种子间差异显著;在结实大年,除0%T外的其他单宁含量种子的PDAD和HDAD在不同大小种子间均差异显著.ISPR在中等单宁含量种子最大,高单宁含量种子最小;PRAD分别在结实小年的高单宁含量种子和结实大年的无单宁种子最大;不论在结实大年还是结实小年,HRAD均在高单宁含量种子最大,中等单宁含量种子最小.这说明结实大年可延缓啮齿动物对种子的消耗速率,提高种子的HRAD,但种子扩散距离减小;啮齿动物在结实大年和小年均表现出对大种子的扩散偏好,且大种子被扩散的距离更远;啮齿动物在不同结实年份均偏好于就地取食中等单宁含量种子,而扩散高单宁含量种子.     

食种子动物对三种锦鸡儿属植物繁殖更新的影响

锦鸡儿属(Caragana)植物是黄土高原植被的重要组分,具有固氮、耐旱的特性,对黄土丘陵沟壑区的植被恢复具有重要意义。作者于2002–2003年对该属本地种白毛锦鸡儿(C.licentiana)和甘蒙锦鸡儿(C.opulens)及外来种中间锦鸡儿(C.intermedia)种子被动物取食情况进行了比较研究。2003年白毛锦鸡儿种子散落前的虫害率为88.2%,甘蒙锦鸡儿为29.7%±1.7%,中间锦鸡儿为43.2%±4.8%。种子落地后,白毛锦鸡儿和甘蒙锦鸡儿的种子被小啮齿动物取食的比例在各种微生境下均为100%,且取食速度很快,在种子放置后4d内即全部被取食。中间锦鸡儿种子被取食率则因微生境的不同而不同:在灌丛下被取食率为100%,在半阳坡的开阔草地上为85.0%±10.0%,在种植植物未成功的水平阶地上为31.0%±8.7%,且取食速度相对较慢。研究结果表明,食种子动物的取食大大减少了白毛锦鸡儿和甘蒙锦鸡儿的种子量以及种子传播萌发的机会。但食种子动物可能充当了白毛锦鸡儿4.0%硬实种子的传播者,从而有助于白毛锦鸡儿实现种子途径的更新;中间锦鸡儿种子无论在散落前还是散落后受食种子动物的影响均很小。     

壳斗科三种植物种子大小对昆虫寄生及种子存活率的影响

种子内的寄生昆虫可以严重影响种子的发育、损害种子活力。种子足余策略理论认为大种子有利于抵御和适应昆虫寄生取食,但动物最优觅食理论推测,大种子更易遭受昆虫寄生。为对这两种对立观点进行验证,本实验以青冈、苦槠和麻栎各2个种群的种子为材料,对昆虫寄生与完好种子间的体积和萌发率进行比较,并对寄生种子萌发率与种子体积的关系进行了分析。结果显示:(1)在6个种群的种子中,只有松阳麻栎和青冈种群的寄生种子体积大于完好种子,其余4个种群的寄生种子体积小于完好种子,但这种差异不显著;(2)所有寄生种子的整体萌发率(18%)显著低于完好种子(45.66%)(P<0.001),在不同种群内,寄生种子的萌发率也分别显著低于完好种子。(3)比较同种植物体积差异显著的寄生种子的萌发率发现,大种子总比小种子具有更高的萌发率,但差异不显著;在不同植物的寄生种子间比较时,体积最大的麻栎种子萌发率显著高于体积较小的青冈和苦槠种子。研究结果表明,象虫在种子上产卵时对大种子没有选择偏好,在昆虫寄生取食严重损害种子活力的压力下,大种子比小种子具有更强的耐受力。     

Post‐dispersal seed predation and its relations with seed traits: a thirty‐species‐comparative study

Post‐dispersal seed predation is a key process determining the variability in seed survival in forests, where most seeds are handled by rodents. Seed predation is thought to affect seedling regeneration, colonization ability and spatial distribution of plants. Basic seed traits are the essential factors affecting rodent foraging preferences and thus seed survival and seedling recruitment. Many studies have discussed several seed traits and their effects upon seed predation by rodents. However, the results of those previous studies are usually equivocal, likely because few seed traits and/or plant species tend to be incorporated into these studies. In order to elucidate the relationships between seed predation and seed traits, we surveyed the predation of 48 600 seeds in a natural pine forest, belonging to 30 species, for three consecutive years. The results demonstrated that: (i) seed size and seed coat hardness did not significantly affect seed predation; (ii) total phenolics had a negative effect upon seed predation; (iii) positive effects of nitrogen content upon seed predation were found. From our study, it seems that the better strategy to prevent heavy predation is for plants to produce seeds with higher total phenolics content rather than physical defenses (i.e. hard seed coat) or larger seeds. Additionally, rodent foraging preference may depend more on Nitrogen content than other nutrient content of seeds.     

Shell strength and primate seed predation of nontoxic species in eastern and southern Africa

The large African primates that eat fruit destroy the seeds of a number of fruiting species. This paper addresses several questions about seed-eating: What is the nature of the dietary niche provided by the nonpoisonous seeds of eastern and southern Africa? How well are these seeds mechanically protected? What other means of reducing seed predation are employed by the plants? and is the niche ecologically stable? Measurements of seed shell strength on 37 species from 17 families reveal a range of values, from <100-kg (numerous species) to over 2000-kg (palm nuts) breaking load. Primates crack open with their teeth seed shells from species exhibiting test strengths less than 600 kg. Variation in shell strength appears to increase dramatically for average species strengths above 100 kg. Plant species are not characterized by specific shell strengths but instead, display envelopes of shell strength overlapping broadly with other species. Taking this into account, adult male baboons (Papio spp.) appear to be dentally capable of preying upon most of the seed species of eastern and southern Africa. The possibility for predation of nonpoisonous seeds exists primarily because the plants periodically produce large crops in synchrony and the hard-shelled seeds are effectively dispersed, sometimes explosively but more often by means of edible fruits. The concomitant primate seed predation is a facultative specialization, of little apparent threat to the community of plants that support it.     

Pre- and postdispersal seed predation by rodents: balance of food and safety

Seed presentation and availability for seed predators changeduring every plant reproductive cycle. We know very little abouthow those changes impinge on both the ability of seed predatorsto impact plant populations and the foraging costs associatedwith seed consumption. Therefore, we conducted several fieldexperiments to evaluate whether wood mice Apodemus sylvaticusbalance food and safety while foraging on Helleborus foetidusseeds during both the pre- and early postdispersal phases ofthe plant reproductive cycle. Both food and safety were keydeterminants of mouse foraging on H. foetidus seeds, thoughtheir roles were not consistent along the plant reproductivecycle. Thus, augmenting ambient food reduced fruit removal bymice during the predispersal phase. During the postdispersalphase, seeds in sheltered microsites experienced higher removalrates than those located in nonsheltered microsites; however,no effect of food augmentation was detected. This apparent reversedrole of food and safety on decision making by mice seemed closelylinked to both the dramatic changes in accessibility and presentationof H. foetidus seeds and the coupled changing foraging costsfaced by mice at different phases of the plant reproductivecycle. For instance, because the cost of foraging for predispersalseeds was higher than for postdispersal seeds, the effect offood augmentation on foraging by wood mice was greater duringthe predispersal phase. Thus, our study illustrates the needof considering differences between pre- and postdispersal seedpredation in the study of granivore rodents and their impacton plant populations.     

Why do some nectar foragers perch and others hover while probing flowers?

Diurnal hawkmoths, Hemaris fuciformis, and bumblebees, Bombus pasquorum, were observed foraging for nectar in flowers of Viscaria vulgaris. The hawkmoths hovered in front of the flowers, while the bees perched on them. The hawkmoths had a faster probing rate than the bees, and consequently also had higher gross and net rates of energy gain. A model is presented that shows that hovering only yields a higher net rate of energy gain (NREG) than perching when nectar volumes are high due to low competition for the resource. The difference in NREG of perchers and hoverers decreases with an increase of competition, and eventually perching yields the highest NREG. This is an effect of the higher cost of hovering. The results suggest that hovering can only evolve as a pure evolutionarily stable strategy (ESS) if competition is reduced, for example by co-evolutionary specializations with plants. The possibility that it has evolved as a mixed ESS (i.e. individuals can both hover and perch depending on the resource level) is discussed. The evolution of optimal foraging strategies is discussed, and it is pointed out that the rate of gain of an animal is independent of the strategy used when all competing foragers use the same strategy, but competitively superior strategies will nevertheless evolve because they are ESSs. Competition between strategies with different energy costs are special, because resource availability determines which strategy is competitively superior. A high-cost strategy can only evolve as a pure ESS at high resource levels, or as a mixed ESS at intermediate levels.