Seed dynamics of resprouting shrubs in grassy woodlands: Seed rain,predators and seed loss constrain recruitment potential

Abstract Measuring the fate of seeds between seed production and seedling establishment is critical in understanding mechanisms of recruitment limitation of plants. We examined seed fates to better understand the recruitment dynamics of four resprouting shrubs from two families (Fabaceae and Epacridaceae) in temperate grassy woodlands. We tested whether: (i) pre‐dispersal seed predation affected seed rain; (ii) post‐dispersal seed predation limited seed bank accumulation; (iii) the size of the seed bank was related to seed size; and (iv) viable seeds accumulated in the soil after seed rain. There was a distinct difference in seed production per plant between plant families with the legumes producing significantly more seeds per individual than the epacrids. Seed viability ranged from 43% to 81% and all viable had seed or fruit coat dormancy broken by heat or scarification. Pre‐dispersal predation by Lepidopteran larvae removed a large proportion of seed from the legume seed rain but not the epacrids. Four species of ants (Notoncus ectatomoides, Pheidole sp., Rhytidoponera tasmaniensis and Iridomyrmex purpureus) were major post‐dispersal seed removers. Overall, a greater percentage of Hardenbergia (38%) and Pultenaea (59%) seeds were removed than the fleshy fruits of Lissanthe (14%) or Melichrus (0%). Seed bank sizes were small (<15 seeds m^?2) relative to the seed rain and no significant accumulation of seed in the soil was detected. Lack of accumulation was attributed to seed predation as seed decay was considered unlikely and no seed germination was observed in our study sites. Our study suggests that seed predation is a key factor contributing to seed‐limited recruitment in grassy woodland shrubs by reducing the number of seeds stored in the soil.     

Population dynamics of the shrub Acacia suaveolens (Sm.) Willd.: Seed production and predispersal seed predation

Seed production and predispersal seed predation in the shrub Acacia suaveolens were examined over 3 consecutive years in eight populations in south-eastern Australia. Seed-crop sizes varied both between and within populations of different ages. Seed production was maximal in the first one to four flowering seasons after establishment, and then declined with plant age. The size of the annual seed-crop was also influenced by rainfall for that year. Predispersal seed predation varies between populations over fruiting seasons with the initial large seed-crops resulting in predator satiation. Within one fruiting season, no significant variation in the extent of predispersal seed predation was found in any of three populations studied. There were two major forms of predispersal seed loss: toss of whole fruits to Melanterius corosus (Coleoptera: Curculionidae) and external insect seed grazers, and loss of individual seeds within fruits to M. corosus. Exclusion experiments showed that seeds lost to these predispersal seed predators would otherwise have been matured by the parent plant. The effects of predispersal seed predation cannot be directly related to seedling recruitment. Indirectly, such predation may influence the dispersion of seeds in the soil profile and hence, subsequent recruitment.     

Soil seed bank dynamics in the rare obligate seeding shrub,Grevillea barklyana (Proteaceae)

Abstract Grevillea barklyana is a rare, obligate seeding shrub that occurs in fire-prone heath and woodland in southeastern Australia. The size and dynamics of the soil seed bank were investigated in populations 2–29 years after fire in order to identify fire frequencies that would threaten population survival. Plants began flowering 2 years after fire (16%), but maximum inflorescence and seed production did not occur until 15–16 years after fire when plants were full size. Seed production declined in older populations. Fruit:flower ratios were very low (0.1–5.3%). Most flowers (> 95%) failed to initiate fruits. Of the initiated fruits, about 60% aborted, 27% were attacked by parrots and 13% matured. An exclusion experiment showed that fruits attacked by parrots would have otherwise matured. Seeds were consumed on the soil surface by mammals, particularly rodents. Post-dispersal seed losses were consistently high in all months and in all populations (78–100%). Mortality of adult plants increased with time since fire. In the 29-year-old population, 30% of plants were dead and 33% were senescent. Recruitment was negligible in the absence of fire, but following fire the seedling to parent ratio was 20:1. Seedling survival was high (90%) following two years of above-average rainfall. The observed seed bank was small (3–16 seeds irr) and increased during the first 16 years after fire before declining in the 21- and 26-year-old populations. The size and dynamics of the seed bank may limit population numbers, especially if the intervals between fires are less than 10 years or greater than 25 years.     

动物与植物种子更新的关系Ⅱ．动物对种子的捕食、扩散、贮藏及与幼苗建成的关系

植物的繁殖体总是面临来自各类生物（如昆虫、脊椎动物、真菌）的捕食风险。因动物捕食引起的种子死亡率影响植物的适合度、种群动态、群落结构和物种多样性的保持。种子被捕食的时间和强度成为植物生活史中发芽速度、地下种子库等特征的主要选择压力,而种子大小、生境类型等因素也影响动物对植物种子的捕食。捕食者饱和现象被认为是植物和种子捕食者之间的高度协同进化作用的结果,是限制动物破坏种子、提高被扩散种子存活率的一种选择压力。大部分群落中的大多数植物种子被动物扩散。种子扩散影响种子密度、种子被捕食率、病原体攻击率、种子与母树的距离、种子到达的生境类型以及建成的植株将与何种植物竞争,从而影响种子和幼苗的存活,最终影响母树及后代植物的适合度。种子被动物扩散后的分布一般遵循负指数分布曲线,大多数种子并没有扩散到离母树很远的地方。捕食风险、生境类型、植被盖度均影响动物对种子的扩散。植物结实的季节和果实损耗的过程也体现了其对扩散机会的适应。许多动物有贮藏植物种子的行为。动物贮藏植物繁殖体的行为,一方面调节食物的时空分布,提高了贮食动物在食物缺乏期的生存概率;另一方面也为种子萌发提供了适宜条件,促进了植物的扩散。于是,植物与贮食动物形成了一种协同进化关系,这种关系可能是自然界互惠关系（mutualism)的一种。影响幼苗存活和建成的因子包括种子贮蒇点的微生境、湿度、坡向、坡度、林冠盖度等。许多果食性动物吃掉果肉后,再将完好的种子反刍或排泄出来。种子经动物消化道处理后,发芽率常有所提高。     

小果野芭蕉种子散布和不同时空尺度上种子被捕食格局

 以分布在云南西双版纳地区的大型先锋草本植物小果野芭蕉(Musa acuminata)为研究材料,研究其种子初次散布过程和不同时空尺度上种子被 捕食格局。小果野芭蕉的成熟果实有75%在夜间被取食和传播,在白天消失的果实则占25%。蝙蝠是其最主要的种子传播者,鸟类在其种子传播 过程中也起到一定的作用。人工摆放种子试验结果显示小果野芭蕉种子的主要转移者是小型啮齿类(鼠类)和蚁类：在开放处理下3 d后转移率为 86%,排除蚁类(鼠类可进入)处理下种子转移率为69%以及排除鼠类(蚂蚁可进入)处理下种子被转移率为56%。季节、地点和生境均显著影响人工 摆放种子被转移强度：雨季显著高于旱季(p<0.001), 野芭蕉生境显著高于与其相连的自然森林和荒地(p<0.001),在人为干扰较少的补蚌自然 保护区显著低于西双版纳热带植物园和新山,而后两者之间并无显著差异(p>0.05)。同时,地点和生境以及季节、地点和生境都有显著的交互 作用。与相邻的森林和荒地相比,野芭蕉群落中种子被鼠类捕食的强度最大且受蚁类二次转移的比例最少,森林和荒地中种子被鼠类捕食的强 度相对较小且蚁类对种子的二次转移比例较高,从而更好地帮助种子逃避鼠类捕食。因此,依赖于食果动物(主要是蝙蝠, 也包括鸟类)的初次 散布是小果野芭蕉种子逃避捕食的关键。     

亚热带林区啮齿动物对樱桃种子捕食和搬运的作用格局

在都江堰林区,通过在原生林、次生林和灌丛3类生境释放野生和栽培樱桃(Cerasus pseudocerasus)的种子,研究了啮齿动物对樱桃种子的捕食及其对樱桃种群更新的作用。结果表明,啮齿动物对樱桃两类种子的搬运无显著差异,而啮齿动物对野生樱桃种子的收获则明显快于栽培樱桃种子,且在3类生境均有类似的趋势。这说明啮齿动物偏爱于收获具有较高收益(种仁重/种子重)的野生樱桃种子。啮齿动物在小于10 d的时间尺度收获了所有释放的樱桃种子,其中,70%以上为啮齿动物所搬运。春季食物的匮乏可能是导致啮齿动物对樱桃种子有较大捕食压力的主要原因,而生境类型间的差异较小。因此,啮齿动物是都江堰林区樱桃地表种子的主要捕食者,它们对野生樱桃种子的选择性捕食和搬运能影响樱桃种子/果实的进化及其种群更新。     

Insects remove more seeds than mammals in first‐year prairie restorations

Seed sowing is a common early step in restoration, but seed consumers can impede plant establishment and alter community structure. Moreover, seed consumers vary in feeding behaviors and the relative importance of different seed consumer groups during restoration are not well understood. At 12 first‐year prairie restorations in Michigan, we studied seed predation using seed removal trays to ask: What is the relative magnitude of seed removal by insects and mammals? Do seed removal rates change over the growing season? Do habitat edges influence seed removal? At what rates are 10 prairie plant species' seeds removed by mammals and insects? Seed removal depended on consumer type, time of year, and seed species. Insects accounted for the majority of seed removal, contrary to previous research in similar systems. In May, insects removed 1.8 times more seeds than mammals, while in August, they removed 5.1 times more. There was greater seed removal in August. During May 28% of seeds were removed, compared to 54% of seeds removed during August, an increase driven by insects. Edge proximity did not influence seed removal. Certain seed species were removed more than others. For example, Lespedeza capitata (round‐headed bush clover) was always removed at high rates, whereas Coreopsis lanceolata (lance‐leaved coreopsis) and Andropogon gerardii (big bluestem) were always removed at low rates. Mammals and insects showed different preferences for several species. This research suggests a prominent role of seed predation, particularly by insects, for early prairie restoration dynamics, with influences varying temporally and among species.     

土壤种子库的研究进展及若干研究热点

 土壤种子库是指存在于表层土壤（包括凋落物）中的有生命的种子。土壤种子库的研究已是植物生态学研究不可缺少的一部分,现已成为植物种群生态学中比较活跃的领域。土壤种子库时期是植物种群生活史的一个重要阶段,有人称之为潜种群阶段。土壤种子库对一年生植物来说尤其重要。土壤种子库简单地可分为瞬时土壤种子库和长久土壤种子库,即使给予理想的萌发条件如季节、温度、湿度等,土壤种子库中也仍有部分种子保持休眠状态,休眠的种子组成了土壤长久种子库的成分。时空异质性是土壤种子库的基本特性之一,不仅不同植被类型的土壤种子库具有不同的组成、大小和多样性,而且微环境也影响土壤种子库的分布格局。由于萌发、捕食和衰老等原因,土壤种子库具有季节动态,一般在旧种子萌发之后,新种子散布之前达到最低点。在高等植物占据的大多数生境中,以休眠繁殖体形式存在的个体远远超过地上植株的数量;土壤种子库、幼苗库和成年植被相互联系相互影响。由于各种原因如群落类型的差异、群落的演替阶段、取样的时间等,地上植被和土壤种子库之间关系大体上可分为两种情况,即相似性和差异性;研究土壤种子库的方法通常有萌发法和物理分离法。土壤种子库能部分反应群落的历史,对退化生态系统的恢复起着重要的作用。目前土壤种子库的主要研究热点问题可分为以下几个方面：1）土壤种子库的研究方法,2）土壤种子库的分类问题,3）土壤种子库分布的时空格局,4）地上植被和土壤种子库的关系,5）土壤种子库的动态等。     

Spicing up restoration: can chili peppers improve restoration seeding by reducing seed predation?

Seed predation by rodents presents a significant barrier to native plant recruitment and can impede restoration seeding efforts. In nature, some plants contain secondary defense compounds that deter seed predators. If these natural defense compounds can be applied to unprotected seeds to inhibit rodent granivores, this approach could improve restoration seeding. Capsaicin is the active ingredient in chili pepper (Capsicum spp.) seeds that creates the burning sensation associated with human consumption of hot peppers. This compound has a similar effect on other mammals and is believed to have evolved as a deterrent to rodent seed predators. We used seed‐coating techniques to attach powder ground from Bhut Jolokia (Capsicum chinense) peppers to native plant seeds and evaluated the efficacy of these seed coatings for deterring rodent seed predation and enhancing native plant recruitment using laboratory and field experiments. Laboratory feeding trials demonstrated that native deer mice (Peromyscus maniculatus) consumed far fewer pepper‐coated seeds compared to untreated control seeds. Field seed‐addition experiments consistently demonstrated that rodent seed predation reduced native plant recruitment over the 4‐year study. Coating techniques used in the first 3 years were not persistent enough to reduce rodent seed predation effects on plant recruitment. However, a more persistent coating applied in conjunction with late‐winter sowing negated rodent seed predation effects on recruitment in year 4. Our results demonstrate that coating seeds with natural plant defense compounds may provide an effective, economical way to improve the efficacy of plant restoration by deterring seed predation by ubiquitous rodent granivores.     

Reproductive potential and seedling establishment of the invasive alien tree Schinus molle (Anacardiaceae) in South Africa

Schinus molle (Peruvian pepper tree) was introduced to South Africa more than 150 years ago and was widely planted, mainly along roads. Only in the last two decades has the species become naturalized and invasive in some parts of its new range, notably in semi‐arid savannas. Research is being undertaken to predict its potential for further invasion in South Africa. We studied production, dispersal and predation of seeds, seed banks, and seedling establishment in relation to land uses at three sites, namely ungrazed savanna once used as a military training ground; a savanna grazed by native game; and an ungrazed mine dump. We found that seed production and seed rain density of S. molle varied greatly between study sites, but was high at all sites (384 864–1 233 690 seeds per tree per year; 3877–9477 seeds per square metre per year). We found seeds dispersed to distances of up to 320 m from female trees, and most seeds were deposited within 50 m of putative source trees. Annual seed rain density below canopies of Acacia tortillis, the dominant native tree at all sites, was significantly lower in grazed savanna. The quality of seed rain was much reduced by endophagous predators. Seed survival in the soil was low, with no survival recorded beyond 1 year. Propagule pressure to drive the rate of recruitment: densities of seedlings and sapling densities were higher in ungrazed savanna and the ungrazed mine dump than in grazed savanna, as reflected by large numbers of young individuals, but adult : seedling ratios did not differ between savanna sites. Frequent and abundant seed production, together with effective dispersal of viable S. molle seed by birds to suitable establishment sites below trees of other species to overcome predation effects, facilitates invasion. Disturbance enhances invasion, probably by reducing competition from native plants.     

LIFE HISTORY AND SEED DISPERSAL OF THE SHORT-LIVED CHAPARRAL SHRUB DENDROMECON RIGIDA (PAPAVERACEAE)

The natural history of the short-lived, fire-following shrub Dendromecon rigida Benth. was studied with emphasis on reproductive output and the consequences of seed dispersal, by intensive study of one population and comparisons with several others.

1. In 16 populations throughout California, mean seed weight ranged from 10 to 16 mg, and was not correlated with rainfall, elevation or latitude. Seed number per fruit ranged from 3 to 11. Adult populations ranged in size up to several million, but each was probably even-aged, established after recent disturbance. Seed-dispersing ants were found at all localities.
2. Reproduction began one year after establishment. Precocious reproduction did not increase mortality; rather, longer-lived plants were more fecund at all ages.
3. The percentage of the population reproducing and fruit number in the most fecund plants were directly related to rainfall in the preceding six months, regardless of plant age. Reproduction also differed according to topographic position.
4. Mean life expectancy of one-year-old plants was about six years, and only 5% survived to age 10 years. Intraspecific density effects were not apparent. Seed longevity probably exceeds adult longevity by a factor of five or ten.
5. The fruits are explosive and secondary dispersal is by gravity and ants. The ants are attracted to a caruncle which represents 14% of the propagule's caloric content. In the intensive study area ants removed more seeds from more sites than did vertebrate seed predators. Pogonomyrmex subnitidus developed large nests on open sites on ridges, and thus provided uphill (even interdrainage) dispersal; it discarded seeds on the soil surface, free to roll but open to predation. Camponotus species had fewer foragers but many small nests in well-covered sites, and discarded the seeds in underground refuse galleries. Dispersal thus affects exposure to predation, the microsite of germination, conspecific aggregation, and position in the mosaic patterns of topography and fire.

     

Seed predation of Copaifera langsdorffii (Fabaceae): a tropical tree with supra‐annual fruiting

Seed predation is an important ecological and evolutionary force that directly affects the distribution of plant species. Copaifera langsdorffii is a tropical tree species with supra‐annual fruiting, which has its seeds predated by a specialist endogenous insect (Rynochenus brevicollis: Curculionidae) in the Brazilian savanna. Three hypotheses were addressed: (i) the predator satiation hypothesis, (ii) the resource concentration hypothesis and (iii) the larger seed predation hypothesis. A total of 112 individual C. langsdorffii were monitored monthly from January to August during four consecutive years (from 2008 to 2011) to determine the presence of fruits on each plant. All trees produced fruits in the year 2008, whereas none of them produced flowers or fruits in 2009 or 2010. Moreover, only 65 individuals (58%) marked in 2008 produced fruits in 2011. The number of fruits per plant was approximately 21% greater in 2008 than in 2011, while the percentage of seed predation was 76% greater in 2011, thereby supporting the predator satiation hypothesis. The percentage of seeds predated was not affected by the number of fruits per plant. Therefore, our data did not support the resource concentration hypothesis. Plants producing large seeds experienced more seed predation by R. brevicollis, supporting the larger seed predation hypothesis. In addition, we also observed a positive relationship between seed volume and adult R. brevicollis weight. This study demonstrates the importance of supra‐annual fruiting for increasing survivorship of C. langsdorffii seeds both at the individual and the population level, and suggests that seed predators select plants producing large seeds as a way of increasing the number of offspring.     

Effects of Dispersal Pattern and Mammalian Herbivores on Seedling Recruitment for Virola michelii (Myristicaceae) in French Guiana1

The effects of dispersal pattern (seeds in small clumps vs. seeds scattered in pairs) and distance to the nearest Carapa procera (Meliaceae; a tree that produces seeds preferred by terrestrial vertebrates) on survival of seeds and seedlings were examined for the animal–dispersed tree species Virola michelii (Myristicaceae) in a mature forest‘at Paracou, French Guiana, in 1992 and 1993. We assessed the putative role of ground–dwelling mammalian herbivores, rodents, and ungulates that filter the seed shadow, acting either as dispersers or predators and thus modifying the original pattern of seed dispersal made by frugivores. We measured the effects of simulated seed burial by rodents using marked seeds and quantified the effect of protecting seeds and seedlings from ground–dwelling vertebrates on seedling germination and survival with fence exclosures in 1992. Dispersal pattern had short–term but no long–term effects on the proportion of V. michelii seeds that survived one year later as seedlings. In the short term, within six weeks, clumped seeds survived better than scattered seeds in both years. Marked seeds that were removed from their site of dispersal were eaten; rodents only rarely buried seeds of V. michelii, and seed burial reduced seed and seedling survivorship. The combined effect of the factors year and Carapa proximity significantly affected seed survival within six weeks. Although six–week seed survival was greater in 1993 than in 1992, seedling establishment was lower in 1993 than in 1992 following a lower rainfall regime during the key period of seed germination (February). One–year seed and seedling survivorship was similar between treatments and years. Seed survival and seedling establishment in V. michelii was dependent on vertebrates in the short term and on climate in the long term. Overall, seed and seedling survivorship depended on a combination of these factors.     

Dispersal and recruitment dynamics in the fleshy‐fruited Persoonia lanceolata (Proteaceae)

Question: What is the role of dispersal, persistent soil seed banks and seedling recruitment in population persistence of fleshy‐fruited obligate seeding plant species in fire‐prone habitats? Location: Southeastern Australia. Methods: We used a long‐term study of a shrubby, fleshy‐fruited Persoonia species (Proteaceae) to examine (1) seed removal from beneath the canopy of adult plants; (2) seedling recruitment after fire; (3) the magnitude and location of the residual soil seed bank; and (4) the implications for fire management of obligate seeding species. We used demographic sampling techniques combined with Generalised Linear Modelling and regression to quantify population changes over time. Results: Most of the mature fruits (90%) on the ground below the canopy of plants were removed by Wallabia bicolor (Swamp wallaby) with 88% of seeds extracted from W. bicolor scats viable and dormant. Wallabies play an important role in moving seeds away from parent plants. Their role in occasional long distance dispersal events remains unknown. We detected almost no seed predation in situ under canopies (< 1%). Seedling recruitment was cued to fire, with post‐fire seedling densities 6‐7 times pre‐fire adult densities. After fire, a residual soil seed bank was present, as many seeds (77‐100%) remained dormant and viable at a soil depth where successful future seedling emergence is possible (0‐5 cm). Seedling survival was high (> 80%) with most mortality within 2 years of emergence. Plant growth averaged 17 cm per year. The primary juvenile period of plants was 7–8 years, within the period of likely return fire intervals in the study area. We predicted that the study population increased some five‐fold after the wildfire at the site. Conclusions: Residual soil seed banks are important, especially in species with long primary juvenile periods, to buffer the populations against the impact of a second fire occurring before the seed bank is replenished.     

动物对花楸树种实的取食与传播

花楸树是我国东北林区重要的非木质资源树种,其种实既有自然散布方式,也有动物散布方式.本文通过对花楸树种实散布过程中动物活动特点的研究,探索动物取食和传播花楸树种实的规律及其对花楸树天然更新的影响.在2008和2009年花楸树果实成熟期,通过定期观察取食花楸树果实的鸟类及其取食方式,确定对花楸树果实有拜访行为的鸟类有8种,其中食果肉鸟类斑鸫、灰喜鹊和白背啄木鸟对花楸树种实有传播作用,它们对花楸树果实的拜访频率分别为54%、12%和7%,取食后第一落点集中于距离母树5～10 m之间（占68.2%）,其次为距离母树5 m以内（占27.3%）,距离母树10 m以外的比例很小（占4.5%）.果实在鸟类消化道内的滞留时间可达20 min,表明潜在传播距离会很长.人工摆放果实和种子试验表明,在不同生境地面摆放的果实6～7 d内全部消失,果实的取食者主要是啮齿类和地面取食的鸟类,取食率均较高（50%～70%）;种子的取食者为啮齿类、地面取食的鸟类和蚁类,取食率均较低（1%～5%）.花楸树为多种动物提供食物,而动物为花楸树传播种子,动物的取食对花楸树的天然更新有重要影响.     

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.     

Co-evolution of seed size and seed predation

Using the evolutionarily stable strategy (ESS) approach in a model for the co-evolution of seed size and seed predation, I show that seed size variation within individual plants is favoured if there is a trade-off in the predator's attack rate for different seed sizes. A single seed size is not evolutionarily stable because a predator that is optimally adapted to one particular seed size cannot prevent invasion by plants with a different seed size. The model generates the following predictions. The ESS consists of a continuous range of seed sizes. Small seeds tend to be attacked more frequently than big seeds. Plants with many resources and plants with low (frequency-independent) juvenile mortality have more variable seeds than plants with few resources and a high juvenile mortality. Seed size variation is higher in fluctuating populations regulated by seed predation alone than in stable populations (partially) regulated by seedling competition. Predator searching behaviour does not directly affect the ESS seed size range, but may have an indirect effect by affecting population stability or the significance of seedling competition as a population regulating mechanism. Moreover, seed size distributions are found to be more skewed in favour of small seeds if predation is spatially non-uniform than if predation is more even. Application of the model to systems of several co-evolving plant and predator species is discussed.     

Fruit Production and Seed Predation in Two Miombo Woodland Trees in Zambia

During a six year period (1990–1995) fruiting frequency among marked trees and seed production and predispersal predation by beetle larvae of lsoberlinia angolensis and Julbernardia globiflora of the Leguminosae family and Caesalpinoideae subfamily were studied at two dry miombo woodland sites in central Zambia. Fruit production varied from year to year and was not related to annual rainfall. On average I. angolensis trees fruited once every two years while J. globiflora trees fruited once every two and one-half years. Number of fruits per tree was correlated with tree size for both species. In a peak year fruit biomass represented two to four percent of total above ground biomass but for J. globiflora about 5.5 and 4.5 percent of nitrogen and potassium, respectively, were in fruits compared to 0.6 and 1.8 percent for I. angolensis. The lower fruiting frequency in J. globifora was probably related to the depletion of N reserves during a fruiting year which required a longer replenishment period than in I. angolensis. Predispersal seed predation was low for J. globiflora (10%) and very high in I. angolensis (65–85%) and for the latter species predation rate increased from August to November 1991 and significant variations were observed among years and pod types. Seed predation rate was lowest in a peak fruiting year and this observation provides support for the satiation hypothesis. The results of the study also suggest that I. angolensis and J. globiflora may have different regeneration strategies although reproductive allocation in both species was within the range reported for other iteroparous plants.     

Temporal Variation in Seed Predation by Insects in a Population of Syagrus romanzoffiana (Arecaceae) in Santa Catarina Island, SC, Brazil

Insect seed predation may vary depending on seed production. The present study considers the hypothesis that the rates of seed predation tend to be smaller in years of higher fruit production. Thus, we monitored the production of fruits and predation of seeds of the palm Syagrus romanzoffiana over 2?years in the Atlantic Forest (Parque Municipal da Lagoa do Peri, Florianópolis, SC, Brazil), between July 2006 and June 2008. Plots of 0.25?m² were fitted under 20 mother plants and fruits were monthly collected for assessment of abundance and seed predation. There was variation in fruit production between the 2?years and among reproductive plants. Predation rates were high and occurred in the predispersal phase by the Curculionidae Revena rubiginosa Boheman, Anchylorhynchus aegrotus Fahraeus, and Anchylorhynchus variabilis Gyllenhal. Seed predation by these species of Anchylorhynchus is first registered in the present study. In average, about 60% of the seeds monthly produced in the population tend to escape insect predation in year of high or low production, becoming available for recruitment. The predation rate was not related to the amount of fruits produced per reproductive plant. Also, different than expected, there was a positive relation between the rates of seed predation and the total of fruits produced monthly on the plots. Thus, no evidence for the satiation of insect seed predators was found in this study with S. romanzoffiana.     

The role of invertebrates in seedling establishment in a heterocarpic plant,Atriplex sagittata

Seed heteromorphism is a marked character of many Chenopodioideae (Amaranthaceae). Seed morphs differ in dormancy, germination and seedling biology, but differences in their predation have not yet been studied. Atriplex sagittata produces small black dormant and large brown non‐dormant seeds. In this study, the timing of seed release and seedling establishment were ascertained, and their consumption by invertebrates (carabids, isopods and slugs) was studied. Seeds dispersed in the autumn passed the winter on the ground surface, protected from invertebrate predation by low temperatures. In the following vegetative season, ungerminated black seeds exposed to predation on ground surface were preferred by a large carabid species, Pseudoophonus rufipes. Some black and all brown seeds escaped predation by germinating in early spring. The seedlings were little endangered by carabids and isopods but were preferred by an invasive slug, Arion vulgaris, the feeding of which can exterminate seedlings at places in which slugs are abundant. Invertebrate predation is important factor of seed and seedling mortality of A. sagittata and seed heteromorphism modifies its intensity and timing.