Intraspecific variation of seed floating ability in Sparganium emersum suggests a bimodal dispersal strategy

Water-mediated spread of seeds (hydrochory) plays an important role in the dispersal of aquatic plants. In this study we investigate intraspecific variation in floating ability and germination capacity of Sparganium emersum seeds in relation to seed mass, within three natural populations along the Rur River (the Netherlands–Germany). Our results suggest that S. emersum produces two types of seeds: (i) short-floating seeds (SFS) that sink within 4 weeks (approximately 71% of all seeds), and (ii) long-floating seeds (LFS) that float at least for 6 months (approximately 28% of all seeds). Our study further shows that short-floating seeds display a significantly higher germination (%) (SFS = 89.9% vs LFS = 32.6%), a faster germination rate (SFS = 8.71 ± 3.3 vs LFS = 9.32 ± 3.1 days to germination) and a higher mean seed mass (SFS = 15.17 ± 4.5 vs LFS = 11.25 ± 3.8 mg), compared to long-floating seeds. It is argued that the production of these two types of seeds by S. emersum plants, each type with a different potential for water-mediated dispersal, represents a bimodal hydrochoric dispersal strategy.     

濒危植物明党参种子散布和种子库动态研究

对伞形科濒危植物明党参（Changium smyrnioides)的种子生产、散布和种子库动态进行了研究。明党参平均每株产生完好种子132粒,86.36%来自顶生花序,不同生境中个体产生的种子千粒重有差异;其平均1.06m高的花葶在种子散布过程中易倒伏,使种子远离母株;种子散布格局为聚集型;种子库密度随时间逐渐下降。明党参种子产量和不同时期种子库密度均低于同科非濒危植物峨参（Anthriscus sylvestris),2个种的种子库均为非持续型;明党参具有低种子数、大粒种子、种子散布远离母株以减少种内竞争的K对策,在受到人类大量采挖和生境干扰后种群不容易迅速恢复;而峨参采取高种子数、小粒种子、种子近母株密集分布和产生无性系的r对策,种群受干扰后易恢复。     

海草床种子扩散过程及种子库形成机制研究进展

海草是唯一一类可以完全生活在海水中的高等被子植物,具有重要的生态服务功能和巨大的经济价值。但受气候变化和人类活动的双重影响,海草床退化趋势日益严峻。海草床生态系统受到外界胁迫后的稳定性和恢复能力很大程度上依赖于有性繁殖即种子繁殖,当胁迫造成海草死亡等不可逆转的伤害时,通过沉积物种子库能够进行种群维持和自我更新,因此研究海草种子扩散过程及种子库形成机制对海草生态系统稳定性的维持具有重要意义。综述了海草生活史类型、种子繁殖特征、种子扩散过程及影响因素、种子库形成机制等。在此基础上总结了目前研究存在的几方面不足和未来展望：1)不同环境胁迫条件对海草有性繁殖努力的影响研究;2)海草种子二次扩散的影响因素和扩散机制研究;3)沉积物沉降和再悬浮对种子扩散和截留的影响研究;4)环境因素变化下种子库的潜在分布和海草适宜生境预测与模拟。本研究以期为海草床生态系统的保护恢复研究提供理论参考。     

Timing and spatial patterning of seed dispersal and redistribution in a South American warm desert

We measured newly-produced seeds entering the soil (Potential Seed Bank) to assess the timing and spatial patterning of Phase I dispersal in the central Monte desert, Argentina. Rates of forb- (6.13 mg m^-2 d^-1) and shrub-seed input (48.9 mg m^-2 d^-1) were maximum in early summer. The rate of grass-seed input, instead, was similar in early and late summer (7 to 8 mg m^-2 d^-1). About 90% of forb- and shrub-seed mass entered the habitat through protected (i.e., under canopy) areas, whereas 70% of grass-seed mass did so through exposed areas. Adult plant location and the uneven impact of wind on shrub, forb and grass seeds may explain such patterns.We also compared the Potential Seed Bank with the soil seed bank in the following spring (Realized Seed Bank). Seeds that form transient banks in other ecosystems (e.g., shrub seeds of the genus Larrea, or perennial grass seeds like those of Pappophorum and Trichloris) prevailed in the Potential Seed Bank. Some annual forb seeds, instead, appeared to form a more persistent seed bank, and prevailed in the Realized Seed Bank (e.g., Chenopodium). Horizontal redistribution did not affect the spatial patterning of forb and shrub seeds, but produced a more homogeneous distribution of grass seeds in the habitat. The impact of wind could explain the redistribution pattern of grass seeds. Finally, we found almost 80% of total seeds in the top 2 cm of soil. The smallest grass and forb seeds (Sporobolus and Descurainia) as well as some medium-sized and large forb seeds (e.g., Glandularia, Sphaeralcea, Phacelia) were able to reach deeper soil layers in the central Monte desert.     

Seed bank,seed dispersal and vegetation cover: Colonization along a newly‐created river channel

Question : What is the relative importance of the initial seed bank and subsequent seed dispersal for floristic composition of bank vegetation two years after creation of a newly‐cut reach of a river channel? Location : River Cole, West Midlands, United Kingdom. Methods : We took bank and bed sediment samples from a 0.5‐km reach of a new river channel cut into intact flood‐plain. After river diversion, seed samples deposited on artificial turf mats placed on the river banks and flood‐plain edge were taken in summer and winter 2002 and 2003. Seed rain samples from funnel traps were taken during summer 2002 and 2003. We undertook greenhouse germination trials to assess viable seed species within these samples. In summer 2004, we surveyed river bank vegetation. Agglomerative cluster analysis was used to investigate floristic similarity between seed bank, seed rain, seed deposition samples and final bank vegetation cover. DCA was used to explore contrasts between the samples and to assess whether these reflected interpretable environmental gradients. Results : Seed rain samples contained a small subset of species in the summer depositional samples. 38 species were found within the final vegetation, the seed bank, and at least one of the four sets of depositional samples; a further 30 species not present in the seed‐bank samples were present in at least one of the four sets of depositional samples and the final vegetation. Floristic composition of the vegetation was most similar to the depositional samples from winter 2002 and 2003 and summer 2003. DCA axis 1 reflected a time sequence from seed‐bank samples through depositional samples to the final vegetation. Conclusions : Newly cut river banks were colonized rapidly. Seed remobilization and hydrochorous transport from the upstream catchment are important for colonization. Species richness was highest in samples deposited during winter when high river flows can remobilize and transport viable seeds from upstream. This process would also have enhanced the species richness of seed production along the banks during the second summer (2003).     

Aerial seed bank and dispersal traits in Anthemis chrysantha (Asteraceae), a critically endangered species

Mature plant density and fruit production were monitored in the main population of four successive cohorts of the endangered winter annual Anthemis chrysantha (Asteraceae) in southeastern Spain. Experiments were conducted with artificial rainfall and a wind tunnel to determine the temporal and spatial dispersal pattern of the species and the relationship with rain and wind.     

A seed predator drives the evolution of a seed dispersal mutualism

Although antagonists are hypothesized to impede the evolution of mutualisms, they may simultaneously exert selection favouring the evolution of alternative mutualistic interactions. We found that increases in limber pine (Pinus flexilis) seed defences arising from selection exerted by a pre-dispersal seed predator (red squirrel Tamiasciurus hudsonicus) reduced the efficacy of limber pine's primary seed disperser (Clark's nutcracker Nucifraga columbiana) while enhancing seed dispersal by ground-foraging scatter-hoarding rodents (Peromyscus). Thus, there is a shift from relying on primary seed dispersal by birds in areas without red squirrels, to an increasing reliance on secondary seed dispersal by scatter-hoarding rodents in areas with red squirrels. Seed predators can therefore drive the evolution of seed defences, which in turn favour alternative seed dispersal mutualisms that lead to major changes in the mode of seed dispersal. Given that adaptive evolution in response to antagonists frequently impedes one kind of mutualistic interaction, the evolution of alternative mutualistic interactions may be a common by-product.     

Ant benefits in a seed dispersal mutualism

Myrmecochorous plant seeds have nutrient rich appendages, elaiosomes, which induce some ant species to carry the seeds back to their nest where the elaiosome is consumed and the seed is discarded unharmed. The benefits to plants of dispersal of their seeds in this way have been well documented, but the benefits to the ants from consuming the elaiosomes have rarely been measured and are less clear. Ant benefits from myrmecochory were investigated in a laboratory experiment using the ant Myrmica ruginodis and seeds of Ulex species. To separate the effects of elaiosome consumption on the development of newly produced larvae versus existing larvae, ten ‘Queenright’ colonies containing a queen were compared to ten ‘Queenless’ colonies. Six measures of colony fitness over a complete annual cycle were taken: sexual production, larval weight and number, pupal weight and number, and worker survival. Queenless colonies fed with elaiosomes produced 100.0±29.3 (mean ± SE) of larvae compared to non-elaiosome fed colonies which produced 49.6±19.0; an increase of 102%. Larval weight increased in both Queenright and Queenless colonies. In colonies fed with elaiosomes, larvae weighed 1.02±0.1 mg, but in non-elaiosome fed colonies larvae weighed 0.69±0.1 mg; an increase of 48%. The food supplement provided by Ulex elaiosomes was trivial in energetic terms, under the conditions of an ample diet, suggesting that these effects might be due to the presence of essential nutrients. Chemical analysis of Ulex elaiosomes showed the presence of four essential fatty acids and four essential sterols for ants.     

Long-term demographic consequences of a seed dispersal disruption

The loss or decline of vertebrate frugivores can limit the regeneration of plants that depend on them. However, empirical evidence is showing that this is still very scarce, as functionally equivalent species may contribute to maintain the mutualistic interaction. Here, we investigated the long-term consequences of the extinction of frugivorous lizards on the population persistence of a Mediterranean relict shrub Cneorum tricoccon (Cneoraceae). We examined the demographic parameters among 26 insular and mainland populations, which encompass the entire plant distributional range, comparing populations with lizards with those in which these are extinct, but in which alien mammals currently act as seed dispersers. Plant recruitment was found to be higher on island populations with lizards than on those with mammals, and the long-term effects of the native disperser's loss were found in all vital phases of plant regeneration. The study thus gives evidence of the cascading effects of human-induced changes in ecosystems, showing how the disruption of native ecological processes can lead to species regression and, in the long term, even to local extinctions.     

Seed bank,seed size and dispersal in moisture gradients of temporary pools in Southern France

Plants reduce risk of extinction due to unpredictable rainfall by soil seed banks, dispersal or large seeds. However, seed size also increases independently in dry habitats, and since seed size is in a trade-off with seed number, size of seed banks is expected to increase in moister habitats. Therefore, we wanted to test if seed abundance in soil increases in wet habitats, if seed size increases in dry habitats, and if spread of seeds along the gradient is higher for plants of intermediate habitats in local moisture gradients.We studied 15 temporary pools in three biogeographically separated wetlands in Southern France. For each pool we studied five different moisture levels, totalling 75 local plant communities. We quantified soil seed bank by the seedling emergence method, seed size and an index of spatial spread of seeds in the soil for every species. We also quantified water levels for each plot.We found increasing abundance of seeds in the soil with increasing water levels but lower seed size and higher spread at intermediate water levels. When we controlled for niche position, we found no trade-off between seed size, spread and abundance in the soil seed bank.Type and importance of risk reduction strategies thus appeared to be strongly driven by the plant species’ moisture niche and the spatial arrangement of water levels.     

Aerial seed bank in a cold desert annual-ephemeral species: Role of anatomical structure of stem and delayed fruit dehiscence in timing of seed dispersal

Why the fruits are retained on dead upright herbaceous plants and how this relates to seed dispersal and timing of germination remain unclear. Stems of the annual Euclidium syriacum (Brassicaceae) with infructescences bearing indehiscent silicles remain upright after plants die in the spring. We investigated the effect of anatomical structures of stem and pedicle and delayed silicle dehiscence on seed dispersal phenology of this species. For comparison, sections were made of the stem of the annual Goldbachia laevigata (Brassicaceae), which has stems that fall over when plants die. Compared to G. laevigata, the stem of E. syriacum has vascular bundles that are closer together, a thicker xylem and phloem, more fibers, a thicker perimedullary zone and a smaller pith diameter:stem diameter ratio. The thickened pedicle did not form an abcission layer. By late October, 5–20% of seeds were dispersed, depending on the position of infructescences on the plant. Snow covered the plants in late autumn and when it melted in mid-April many of the plants had fallen over, with a high number of seeds germinating in attached silicles; seedlings became rooted in soil. After snowmelt, 14–15% of the silicles on the remaining upright plants contained seeds; all seeds were dispersed by early July. The anatomical structures of the stem and pedicle plus the delayed dehiscence of silicles explain the presence of an aerial seed bank in E. syriacum and delay of germination of many of seeds until spring. Further, pieces of upright plants are broken off and dispersed by wind, which helps to explain the wide distribution of E. syriacum in the cold desert.     

Mechanisms of long-distance seed dispersal

Growing recognition of the importance of long-distance dispersal (LDD) of plant seeds for various ecological and evolutionary processes has led to an upsurge of research into the mechanisms underlying LDD. We summarize these findings by formulating six generalizations stating that LDD is generally more common in open terrestrial landscapes, and is typically driven by large and migratory animals, extreme meteorological phenomena, ocean currents and human transportation, each transporting a variety of seed morphologies. LDD is often associated with unusual behavior of the standard vector inferred from plant dispersal morphology, or mediated by nonstandard vectors. To advance our understanding of LDD, we advocate a vector-based research approach that identifies the significant LDD vectors and quantifies how environmental conditions modify their actions.     

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.     

Mitochondrial haplotype distribution,seed dispersal and patterns of postglacial expansion of ponderosa pine

Maternally inherited mtDNA in a secondary contact zone of ponderosa pine revealed a cline less than 10 km wide - much narrower than previously described. A survey of 76 populations gave no evidence either of intermixing or of a mosaic contact zone. Such sharp contact zones are consistent with diffusive range expansion, rather than long distance colonization. However, evidence for long distance seed dispersal events was found in two populations where haplotypes were observed far from their main area of occurrence. The results suggest a small number of long distance colonists with diffusive dispersal from these centres.     

The conservation physiology of seed dispersal

At a time when plant species are experiencing increasing challenges from climate change, land-use change, harvesting and invasive species, dispersal has become a very important aspect of plant conservation. Seed dispersal by animals is particularly important because some animals disperse seeds to suitable sites in a directed fashion. Our review has two aims: (i) to highlight the various ways plant dispersal by animals can be affected by current anthropogenic change and (ii) to show the important role of plant and (particularly) animal physiology in shaping seed-dispersal interactions. We argue that large-bodied seed dispersers may be particularly important for plant conservation because seed dispersal of large-seeded plants is often more specialized and because large-bodied animals are targeted by human exploitation and have smaller population sizes. We further argue that more specialized seed-dispersal systems on island ecosystems might be particularly at risk from climate change both owing to small population sizes involved but also owing to the likely thermal specialization, particularly on tropical islands. More generally, the inherent vulnerability of seed-dispersal mutualisms to disruption driven by environmental change (as well as their ubiquity) demands that we continue to improve our understanding of their conservation physiology.     

Ant seed removal in a non-myrmecochorous Neotropical shrub: Implications for seed dispersal

This study investigated ant seed removal of Piper sancti-felicis, an early successional Neotropical shrub. Neotropical Piper are a classic example of bat-dispersed plants, but we suggest that ants are underappreciated dispersal agents. We identified eleven ant species from the genera Aphaenogaster, Ectatomma, Paratrechina, Pheidole, Trachymyrmex, and Wasmannia recruiting to and harvesting P. sancti-felicis seeds in forest edge and secondary forest sites at La Selva, Costa Rica. We also tested for differences in ant recruitment to five states in which ants can commonly encounter seeds: unripe fruit, ripe fruit, overripe fruit, bat feces, and cleaned seeds. Overall, ants harvested more seeds from ripe and overripe fruits than other states, but this varied among species. To better understand the mechanisms behind ant preferences for ripe/overripe fruit, we also studied how alkenylphenols, secondary metabolites found in high concentrations in P. sancti-felicis fruits, affected foraging behavior in one genus of potential ant dispersers, Ectatomma. We found no effects of alkenylphenols on recruitment of Ectatomma to fruits, and thus, these compounds are unlikely to explain differences in ant recruitment among fruits of different maturity. Considering that P. sancti-felicis seeds have no apparent adaptations for ant dispersal, and few ants removed seeds that were cleaned of pulp, we hypothesize that most ants are harvesting its seeds for the nutritional rewards in the attached pulp. This study emphasizes the importance of ants as important additional dispersers of P. sancti-felicis and suggests that other non-myrmecochorous, vertebrate-dispersed plants may similarly benefit from the recruitment to fruit by ants.     

Endozoochory by beetles: a novel seed dispersal mechanism

Background and Aims

Due in part to biophysical sized-related constraints, insects unlike vertebrates are seldom expected to act as primary seed dispersers via ingestion of fruits and seeds (endozoochory). The Mediterranean parasitic plant Cytinus hypocistis, however, possesses some characteristics that may facilitate endozoochory by beetles. By combining a long-term field study with experimental manipulation, we tested whether C. hypocistis seeds are endozoochorously dispersed by beetles.

Methods

Field studies were carried out over 4 years on six populations in southern Spain. We recorded the rate of natural fruit consumption by beetles, the extent of beetle movement, beetle behaviour and the relative importance of C. hypocistis fruits in beetle diet.

Key Results

The tenebrionid beetle Pimelia costata was an important disperser of C. hypocistis seeds, consuming up to 17·5 % of fruits per population. Forty-six per cent of beetles captured in the field consumed C. hypocistis fruits, with up to 31 seeds found in individual beetle frass. An assessment of seeds following passage through the gut of beetles indicated that seeds remained intact and viable and that the proportion of viable seeds from beetle frass was not significantly different from that of seeds collected directly from fruits.

Conclusions

A novel plant–animal interaction is revealed; endozoochory by beetles may facilitate the dispersal of viable seeds after passage through the gut away from the parent plant to potentially favourable underground sites offering a high probability of germination and establishment success. Such an ecological role has until now been attributed only to vertebrates. Future studies should consider more widely the putative role of fruit and seed ingestion by invertebrates as a dispersal mechanism, particularly for those plant species that possess small seeds.     

淡水湿地种子库的小尺度空间格局

以长江中下游淡水湿地湖里沼泽为对象,研究了种子库在小尺度空间范围的水平分布格局。在沼泽中选择地表植被分布比较均一的16m×4m大小的方形区域,按1m间隔获取64个内径7·8cm、深5cm的圆柱状土样,通过幼苗萌发法鉴定每个土样的种子库组成。采用2个格局指数(离散系数和Lloyd平均拥挤指数)以及Moran空间自相关系数分析了种子库中优势种的空间分布格局。结果显示种子库由17个物种组成,多年生的锐棱荸荠(Eleocharisacutangula)和龙师草(E·tetraqueter)是种子库中密度最大的物种。两个格局指数显示7个分布频率大于10%的物种的种子全部为聚集分布。MoransI统计分析显示其中只有3个物种为显著的正的空间自相关,表明这3个物种的种子斑块大小超过了目前的两个取样单位之间的距离(1m),而其它4个物种的种子斑块则小于这个尺度。