种子产量对鼠类扩散栓皮栎坚果的影响

为了解林木种子产量对鼠类种子扩散行为的影响,于2008 年和2009 年,在国有济源市愚公林场调查了栓皮栎的种子雨;在每年的种子雨结束后,选择次生林生境,研究了鼠类对人工释放栓皮栎坚果的扩散,旨在探讨种子产量与种子扩散间的关系。结果表明:1)取食栓皮栎坚果的鼠类主要有大林姬鼠、社鼠和岩松鼠,2008 年的鼠类捕获率(2% )低于2009 年(10%),其差异并未达到显著性水平;2)栓皮栎的种子雨构成和产量存在年际差异,2008 年以完好种子为主且产量高于2009 年,而2009 年以败育种子为主;3)2008 年人工释放栓皮栎坚果的中位存留时间显著高于2009 年;4)2008 年鼠类对栓皮栎坚果的平均扩散距离显著低于2009 年,且两年的搬运距离几乎都集中在9 m 以内;5)2008 年鼠类对栓皮栎坚果的埋藏比例显著高于2009 年。结果提示,在种子高产年份,可能有较多的种子逃脱动物的取食,从而增加种子萌发和幼苗建成的机会,最终促进植物的更新。     

Masting,seed dispersal and seed predation in the cycad Macrozamia communis

Summary Little is known about the adaptive value of mast seeding, a common phenomenon in temperate trees and shrubs. Masting is likely to affect both seed dispersal and seed predation. In systems where similar taxa of animals are involved in these two processes, the consequences of mast seeding are likely to be particularly complicated. This study examined the effects of mast seeding in a cycad, Macrozamia communis, on the dispersal of seeds, the pattern of dispersion of seeds and post-dispersal predation on seeds. Dispersal of seeds by possums was poorer from source plants in a masting population than from source plants in an adjacent, non-masting population. This resulted in fewer seeds per seeding female plant in the masting plot being dispersed to favourable sites. We conclude that this is caused by the feeding behaviour and movements of possums in the masting site. The abundance of seeds in this site did not satiate the post-dispersal predators, native rats. In fact, more seeds in this site were eaten than in the nonmasting site. We suggest that the mast seeding observed in M. communis may not be adaptive, but is more likely a consequence of other factors which synchromize flowering within local populations.     

The seed dimorphism of Spergularia marina in relation to dispersal by wind and water

Summary The seeds of the halophyte Spergularia marina differ both within and between individuals in that they either possess or lack a membranaceous border. This paper presents a morphological study of the length, weight and area of the seed types, and their dispersal characteristics under experimental conditions of wind and water dispersal. The winged seeds are shown to be larger both by length and by weight. Their rate of descent increases with wing loading. If the wing is lacking, however, the rate of descent increases with weight only. The distance of dispersal is equal for both seed types except at low wind speeds, when the winged seeds disperse farther. If the seed wing is removed, the excised seeds have shorter dispersal distances. When dispersed by water, a difference in the distance seeds are dispersed can only be detected in the presence of vegetation. The winged seeds are more frequently trapped in the vegetation as compared to the unwinged seeds. The hypothesis that the seed dimorphism is an adaptation for differential dispersal distances is discussed.     

Dung beetles as secondary seed dispersers in a temperate grassland

The two-phase dispersal event in which dung beetles move seeds after endozoochory is often assumed to be advantageous for plant regeneration. Because seeds are expected to end up in favourable and safe germination sites, it is considered as an example of directed dispersal. However, literature so far is restricted to tropical rain forest ecosystems, while data for temperate regions are lacking. In this study, the effect of dung beetles on seedling establishment of endozoochorically dispersed seeds is evaluated for a temperate grassland ecosystem. We performed a field experiment in which cages excluded dung beetles from horse and cattle dung samples with mixed-in grass seeds. Seed germination from these samples was significantly higher than that from samples which were accessible to dung beetles. This indicates that the effect of dung beetles on short-term seedling establishment was negative, which contrasts with the patterns found for large-seeded species used in tropical studies. This is most likely attributed to the lack of roller species and the larger depth at which tunneling Geotrupes species bury seeds.     

Evolution of a unique seed maturity pattern in Croton bonplandianum Baill strengthens ant-plant mutualism for seed dispersal

Summary The female flowers of Croton bonplandianum bear nectar glands which become active during fruit maturation and attain peak activity just prior to the splitting of fruits. This temporal specificity of nectar gland activity is shown to facilitate seed dispersal by ants, which are attracted to the plant only during the fruit maturation period. The nectar glands establish a nectar influence zone with a radius of 60 cm around the plant within which seed dispersal by ants is effective. Seed dispersal by ants is facilitated only if the seeds are placed within this nectar influence zone. This is accomplished by an intriguing evolutionary shift in the maturation pattern of the fruits. Unlike the usual acropetal development, fruit maturation in Croton is temporally asymmetrical, with the fruits nearer the parental axis maturing early. This unique pattern of fruit development together with the polychasial branching system leads to a concentration of seeds within the nectar influence zone and enhances seed dispersal by ants. The proximate factors responsible for this asynchronous fruit maturity were investigated.     

Secondary seed dispersal in Montrichardia arborescens (L.) schott dominated wetlands in Laguna Grande, Venezuela

Laguna Grande, Monagas State, Venezuela, is a shallow, V-shaped lake created by the confluence of two rivers. Montrichardia arborescens (L.) Schott. dominated wetlands cover most of the north and south arms and the littoral zone of the main body of the lake. The vegetation and seed banks of Montrichardia wetland sites were sampled in the north arm, south arm and main body five times from the end of the dry season in 1991 to beginning of the rainy season in 1992. The composition of the vegetation was similar and changed very little at all three sites during the course of the study. These wetlands had 53 species. Besides M. arborescens, other common species were Hamelia patens Jacq., Mikania cordifolia (L.) Wild., Sarcostemma clausum (Jacq.) Roem. & Schult., and Vitis caribaea L. In both the vegetation and seed banks, species richness was highest during the dry season. Altogether, the seed banks contained the seeds of 61 species of which 35 were also found in the vegetation. Seeds of three tree species were found in the seed banks that did not grow anywhere in the lake. In the seed bank, seeds of Cyperus odoratus L., Eleocharis interstincta (Vahl.) R&S, Ludwigia hyssopifolia (G. Don.) Excell, L. lithospermifolia (Mich.) Hara, Polygonum acuminatum H.B.K., and Sacciolepis striata (L.) Nash were the most abundant. Mean total seed density over the entire study was 6,500, 3,800, and 6,000 seeds/m² in the north arm, south arm, and main basin, respectively. Seed production and dispersal occur primarily during the dry season, and the highest seed densities at all sites were found in the dry season when there was no or little standing water. The lowest seed densities at all sites were found during the rainy season during which seed densities declined over 80% at the north and south arm sites. In the main body of the lake, however, seed densities during the rainy season, although lower than during the dry season, actually increased significantly from 3,600 seeds/m² in August 1991 to 6,000 seeds/m² in October 1991. A significant decrease in seed density in either the north or south arms or both and a significant increase in the main body site during the rainy season occurred for 5 of the 8 species whose seeds were the most abundant, for all life-form guilds, except hydrophytes and for the entire seed bank. Secondary dispersal by water currents during the rainy season appears to be transporting seeds from the north and south arms into the seed bank of the main body of the lake.     

Aspects of variation in a neotropical seed dispersal system

This paper explores the causes and consequences of seed and seedling mortality of the tree Virola nobilis (Myristicaceae) Central Panama in order to understand the advantage to local seed dispersal by birds and monkeys. Post-dispersal mortality due to insects (primarily Conotrachelus spp., Curculionidae) accounts for 30–35% of seed and seedling death during the first 12 weeks after seed fall. Because more seeds and seedlings are killed under and near fruiting trees than 15–45 m away, seed dispersal confers a 20–40 fold advantage on seeds carried 45 m from fruiting adults. In contrast, >60% of seed and seedling death during the first year is due to seed predation by mammals, with >90% due to mammals among the <2% seeds that survive until maternal endosperm is exhausted ±12 weeks after seed fall. Mortaliy due to mammals is independent of distance from parent trees, confering no advantage to seed dispersal. Insects account for variation in mortality attributable to distance effects, mammals to between site effects.Early weevil infestations put a premium on seed removal by large birds (Ramphastos swainsonii, R. sulfuratus, Penelope purpurascens), which carry >50% of the seeds that they eat >40 m, as compared with smaller birds (Baryphthengus martii, Tityra semifasciata, Trogon massena) and monkeys (Ateles geoffroyi), which leave most or all of the seeds that they eat under or near the tree crown.     

昆虫蛀蚀对鼠类介导下的锐齿槲栎种子扩散的影响

森林鼠类的种子贮藏行为对植物的扩散和自然更新有着非常重要的影响。然而,鼠类是否具有鉴别虫蛀种子的能力还存在一定的争议。此外,鼠类的鉴别能力是否受到食物丰富度变化的影响也未见相关报道。采用标签标记法,2011年秋季(9—11月,食物丰富季节)和2012年春季(4—6月,食物匮乏季节)分别在秦岭南坡的佛坪国家级自然保护区内,调查了森林鼠类对完好和虫蛀锐齿槲栎(Quercus aliena)种子的选择差异。结果显示:1)在秋季,尽管2种类型种子的存留动态没有显著差异,但是在后期虫蛀种子的存留时间相对更长;而在春季2种类型种子的存留动态则极为显著,几乎所有的完好种子(99%)在释放后的第3天就被鼠类全部扩散,虫蛀种子的存留时间则相对较长。2)在秋季,鼠类更喜好扩散后取食完好种子;而在春季,鼠类则喜好在原地取食绝大部分的种子,并且优先取食完好种子。3)在秋季,鼠类贮藏了更多的完好种子;而在春季,尽管完好种子在释放后第1天便达到贮藏高峰,然而由于后期的大量被捕食,2种类型种子在贮藏动态上没有显示出显著差异。研究结果表明秦岭地区森林鼠类可以准确区分完好与虫蛀种子,但是食物丰富度会影响鼠类对种子的选择策略。在食物丰富的秋季,鼠类更多地选择贮藏完好种子;而在食物相对匮乏的春季,鼠类更倾向于同时取食2种类型种子。森林鼠类通过对2种类型种子的鉴别和选择,影响不同种子的命运,从而可能对种子的扩散和自然更新产生重要影响。     

Quantity,quality and the effectiveness of seed dispersal by animals

Disperser effectiveness is the contribution a disperser makes to the future reproduction of a plant. Although it is a key notion in studies of seed dispersal by animals, we know little about what determines the effectiveness of a disperser. The role of the present paper is to review the available information and construct a hierarchical framework for viewing the components of disperser effectiveness.Effectiveness has both quantitative and qualitative components. The quantity of seed dispersal depends on (A) the number of visits made to the plant by a disperser and (B) the number of seeds dispersed per visit. The quality of seed dispersal depends on (A) the quality of treatment given a seed in the mouth and in the gut and (B) the quality of seed deposition as determined by the probability that a deposited seed will survive and become an adult. In this paper I review the ways disperser behavior, morphology and physiology can influence these major components of disperser effectiveness, and when data permit present preliminary analyses of relationships among components.     

Seed dispersal by pygmy chimpanzees (Pan paniscus): A preliminary report

The role in seed dispersal played by the pygmy chimpanzees (Pan paniscus) inhabiting Wamba, Republic of Zaïre, was studied. Germination was tested for seeds of 17 plant species recovered from the feces of pygmy chimpanzees at Wamba. The fecal seeds of 13 species germinated, and in six of the species the germination rate for the fecal seeds was higher than that of control seeds. Although five other species showed a higher germination rate in the control seeds than in the fecal seeds, the remaining two species revealed no difference in germination rate between the fecal and control seeds. There was no great difference in germination velocity between the fecal and control seeds of the same species. For comparison, seeds of four plant species collected from the feces of common chimpanzees (Pan troglodytes) and gibbons (Hylobates lar) in captivity in Okinawa were tested for their germinability. In this test, although the seeds had passed through the digestive tract, their germinability demonstrated little change. Based on the behavioral characteristics of the pygmy chimpanzee at Wamba and observations of the captive primates on Okinawa, it seems that pygmy chimpanzees may play an important role in the seed dispersal of fruit plant species at Wamba.     

华山松种子大小对啮齿动物贮藏行为的影响及其时空变化

华山松(Pinus armandii)是广泛分布在中国中西部海拔1000～3300m的山地、种子较大(约300mg)的松属植物。为探究华山松种子大小对啮齿动物贮藏行为的影响,于2006年和2007年在滇西北3个不同地点进行种子标记和追踪实验。结果表明,在所有年份和地点,啮齿动物都倾向于贮藏更多的大种子和取食更多的小种子;啮齿动物贮藏大种子的数量,以及平均距离和最大距离均显著高于小种子。3个地点具有不同的啮齿动物群落结构,从而对种子命运产生显著影响。种子命运在两个年份间也存在显著差异。     

Spatial patterns of dispersal, seed predation and germination during colonization of abandoned grassland by Quercus petraea and Corylus avellana

Key processes during recruitment of Quercus petraea and Corylus avellana were investigated in abandoned calcareous grasslands and adjacent scrub using the following methods: (1) observation of hoarding animals during the main period of ripening of acorns and hazel nuts, (2) exposition of seeds on the soil surface and in 5–6 cm depth to test differences in predation and germination, and (3) mapping of seedlings in the grasslands. European jays (Garrulus glandarius) and mice were the main disperser of seeds. Jays preferred acorns, whereas the rodents were less selective, but probably more important for dispersal of nuts. The maximum dispersal distance was about 10–20 m for mice and was estimated several hundred metres for jays. Mice collected hoards of several seeds in about 2 cm depth in the soil, whereas jays stored single seeds. Seed predation and probably hoarding by mice were highest under scrub and in unmown grassland, while jays preferred mown sites for hoarding. However, hiding of seeds in the soil reduced losses in all sites. Predation of nuts was slightly less intensive than that of acorns. Seeds of Corylus were more sensitive to desiccation than Quercus, but in both species germination was lower for seeds exposed on the soil surface and in drier sites. Quercus and Corylus were the most abundant woody species in the fallow grasslands, probably due to the effective multi-staged dispersal by jays and mice, whereas wind-dispersed and fleshy-fruited species were less common; the latter restricted to margins of adjacent scrubland. The study provides several examples for discordance in suitability of patches for seeds and seedlings due to different habitat requirements of successive developmental stages. This result emphasizes the need for studies in the multiple stages during recruitment of vertebrate-dispersed plants.     

Monitoring seed dispersal at isolated standing trees in tropical pastures: consequences for local species availability

The tropical rain forest landscape has been transformed to a mosaic composed of patches of crops, secondary vegetation and remnant forest fragments of different shapes and sizes. Isolation of patches and fragments is a critical issue in the maintenance of local species diversity. In this study we focus on the dispersal of propagules by birds to understant the movement of plants between landscape components. Seed deposition and the behavior of frugivorous birds were monitored at four isolated fig trees (Ficus yoponensis and F. aurea) in man-made pastures. Seed deposition was measured by trapping seeds under canopy trees for six months and by direct observation of bird visits to the four trees for one year. Seed deposition densities were 465, 614, 632 and 1097 seeds/m² accumulated over six months under each of the four trees. We recorded 8268 seeds of 107 species under the trees, among them, 6726 seeds (81%) were of 56 species dispersed by vertebrate frugivores. Seeds of tree species accounted for 26% of the total species. Seventy-three species of birds perched in the observed trees, and 3344 visits were made by 47 species of frugivores. Frugivorous birds occurred in two groups: resident species nesting in the pastures and resident species nesting elsewhere. Propagule exchange between landscape components is clearly influenced by the behavior of these two groups. Structure and dynamics of the landscape depend on plant species availability within the mosaic. This availability is high and suggests possibilities for the management of the local species diversity of tropical rain forests.     

Aggregated seed dispersal by wreathed hornbills at a roost site in a moist evergreen forest of Thailand

Hornbills (Bucerotidae) are widely regarded as important seed dispersers in tropical forests in Africa and Asia. We investigated how the roosting behavior of wreathed hornbills (Aceros undulatus) influences seed deposition and seedling survival at a roost site in a moist evergreen forest of Khao Yai National Park, Thailand. Fallen fruits and seeds were collected in traps that were placed around a roosting site for 14 months, and seedlings were monitored in adjacent quadrats for 3 years. Seedfall and seedlings of species represented in the hornbill diet occurred at significantly higher densities in the traps and quadrats located beneath the crown of the roosting tree than in those located beyond the crown. With the exception of Cinnamomum subavenium, the seeds and seedlings of most diet species rarely survived beyond the first year. The quality of hornbill dispersal to this roosting site may be poor due to the highly concentrated seedfall, which results in high seed and seedling mortality. However, the number of seeds deposited by each hornbill each day at roosting sites is relatively low. Wreathed hornbills are primarily scatter dispersers during the day and probably serve as agents of seed dispersal in the moist evergreen forest of Khao Yai.     

The effects of seed abundance on seed predation and dispersal by rodents in <Emphasis Type="Italic">Castanopsis fargesii</Emphasis> (Fagaceae)

Based on the animal dispersal hypothesis and the predator satiation hypothesis, we examined the effects of seed abundance at both population (i.e., mast seeding) and community levels on seed predation and dispersal of Castanopsis fargesii (Fagaceae), a rodent-dispersed mast species in Eastern Asia. In a subtropical evergreen broadleaved forest in the Dujiangyan region of Sichuan Province, China, individual seeds with coded tin tags were tracked in two contrasting stands (seed-poor and seed-rich) over two years (2000, a low-seed year; 2001, a high-seed year). Our results showed that: (1) small rodents did not harvest the tagged seeds of C. fargesii more rapid in the high-seed year than in the low-seed year in either stand. But, seed harvest was significantly faster in the seed-rich stand than in the seed-poor stand. (2) The removal proportion was significantly lower in the high-seed year than in the low-seed year for either stand, but the removal proportion was slightly higher in the seed-poor stand than in the seed-poor stand. This indicates that high seed abundance decreases seed removal (predator satiation hypothesis). (3) There were only small differences about seed caching, seed survival and seedling establishment of C. fargesii between years and stands. During the survey, no cached seeds survived to geminate in the spring for both stands and years. (4) Mean dispersal distances of the cached seeds are much shorter in the high-seed year (3.1 m) than in the low-seed year (8.1 m) in the seed-rich stand, though similar trend is not examined in the seed-poor stand. Our results indicate that seed predation and dispersal of C. fargesii are influenced by both mast seeding and community-level seed abundance, which is not completely consistent with either the animal dispersal hypothesis or the predator satiation hypothesis, but seems more related to the predator satiation hypothesis.     

Extremely long-distance seed dispersal by an overfished Amazonian frugivore

Throughout Amazonia, overfishing has decimated populations of fruit-eating fishes, especially the large-bodied characid, Colossoma macropomum. During lengthy annual floods, frugivorous fishes enter vast Amazonian floodplains, consume massive quantities of fallen fruits and egest viable seeds. Many tree and liana species are clearly specialized for icthyochory, and seed dispersal by fish may be crucial for the maintenance of Amazonian wetland forests. Unlike frugivorous mammals and birds, little is known about seed dispersal effectiveness of fishes. Extensive mobility of frugivorous fish could result in extremely effective, multi-directional, long-distance seed dispersal. Over three annual flood seasons, we tracked fine-scale movement patterns and habitat use of wild Colossoma, and seed retention in the digestive tracts of captive individuals. Our mechanistic model predicts that Colossoma disperses seeds extremely long distances to favourable habitats. Modelled mean dispersal distances of 337-552 m and maximum of 5495 m are among the longest ever reported. At least 5 per cent of seeds are predicted to disperse 1700-2110 m, farther than dispersal by almost all other frugivores reported in the literature. Additionally, seed dispersal distances increased with fish size, but overfishing has biased Colossoma populations to smaller individuals. Thus, overexploitation probably disrupts an ancient coevolutionary relationship between Colossoma and Amazonian plants.     

Two-phase seed dispersal: linking the effects of frugivorous birds and seed-caching rodents

Frugivorous birds disperse the seeds of many fruit-bearing plants, but the fate of seeds after defecation or regurgitation is often unknown. Some rodents gather and scatter hoard seeds, and some of these may be overlooked, germinate, and establish plants. We show that these two disparate modes of seed dispersal are linked in some plants. Rodents removed large (>25 mg) seeds from simulated bird feces (pseudofeces) at rates of 8–50%/day and scatter hoarded them in soil. Ants (Formica sibylla) also harvested some seeds and carried them to their nests. Rodents carried seeds 2.5±3.2 m to cache sites (maximum 12 m) and buried seeds at 8±7 mm depth. Enclosure studies suggest that yellow pine chipmunks (Tamias amoenus) and deer mice (Peromyscus maniculatus) made the caches. In spring, some seeds germinated from rodent caches and established seedlings, but no seedlings established directly from pseudofeces. This form of two-phase seed dispersal is important because each phase offers different benefits to plants. Frugivory by birds permits relatively long-range dispersal and potential colonization of new sites, whereas rodent caching moves seeds from exposed, low-quality sites (bird feces on the ground surface) to a soil environment that may help maintain seed viability and promote successful seedling establishment.     

The effect of bat (Rousettus aegyptiacus) dispersal on seed germination in eastern Mediterranean habitats

The fruit-bat Rousettus aegyptiacus (Pteropodidae) in Israel consumes a variety of cultivated and wild fruits. The aim of this study was to explore some of its qualities as a dispersal agent for six fruit-bearing plant species. The feeding roosts of the fruit-bat are located an average of 30 m from its feeding trees and thus the bats disperse the seeds away from the shade of the parent canopy. The bat spits out large seeds but may pass some (2%) of the small seeds (<4 mg) through its digestive tract. However, neither the deposited seeds nor the ejected seeds (except in one case) had a significantly higher percentage germinating than intact seeds. Although the fruit-bat did not increase the percentage germinating, seeds of three plant species subject to different feeding behaviors (deposited in feces or spat out as ejecta) had a different temporal pattern of germination from the intact seeds. The combined seed germination distribution generated by these different treatments is more even over time than for each treatment alone. It is sugested that this increases asynchronous germination and therefore enhances plant fitness by spreading the risks encountered during germination, especially in eastern Mediterranean habitats where the pattern of rainfall is unpredictable.     

Frugivory and the effects of ingestion by bats on the seed germination of three pioneering plants

The dispersion and seedling establishment of pioneering plants can be favoured by the presence of frugivorous bats because the bats usually improve seed germination after ingestion. Although seed germinability is known to vary greatly after ingestion by different bats, the relative contribution of each bat species to seed germination within plant communities is poorly understood. In this study, we first determined the fauna of frugivorous bats in a semideciduous seasonal forest remnant in southern Brazil and subsequently identified the plant species of the seeds passed through their guts. Second, the germination performance (i.e., germination percentage and speed) of the seeds of three pioneering plants (Piper aduncum, Piper hispidinervum and Solanum granuloso-leprosum) ingested by the most abundant bats was compared with that of the non-ingested seeds (seeds collected from fruits). Additionally, the effects on seed germination of different bat species were compared. During one year, five species of frugivorous bats were caught, and the seeds of eleven identifiable plant species (not counting those of undetermined species) were found in their faeces. We found that the germination performance of the seeds of Piper species was significantly enhanced after ingestion by bats, whereas S. granuloso-leprosum seeds had neutral or reduced germinability when seeds in faeces were compared with pulp-removed seeds. Our results revealed that the bat species that were captured exerted different effects upon seed germination; such a disparity is expected to result in different rates of early establishment of these pioneer plants in tropical forests, most likely affecting forest composition and structure, particularly during the initial stages of succession.     

Genetic structure in a population of a tropical tree Ocotea tenera (Lauraceae): influence of avian seed dispersal

We studied the influence of avian seed dispersal on the structuring of genetic diversity in a population of a tropical tree, Ocotea tenera (Lauraceae). The seeds of O. tenera are principally dispersed by four, relatively specialized, fruit-eating bird species (emerald toucanets, keel-billed toucans, resplendent quetzals, and three-wattled bellbirds). We found high genetic diversity within the overall population and significant, nonrandom structuring of that diversity among subpopulations. Subpopulations contained members of several sibling groups, and most saplings within subpopulations were shown not to be the progeny of adult trees within the same subpopulation. Our data indicate that O. tenera subpopulations are founded with several seeds from few maternal families, and that this mode of establishment is an important determinant of population genetic architecture.