Evolution of fruit types and seed dispersal:A phylogenetic and ecological snapshot

Success of flowering plants is greatly dependent on effective seed dispersal. Specific fruit types aid different mechanisms of seed dispersal. However, little is known about what evolutionary forces have driven the diversification of fruit types and whether there were phylogenetic constraints on fruit evolution among angio-sperm lineages. To address these questions, we first surveyed the orders and families of angiosperms for fruit types and found no clear association between fruit types and major angiosperm lineages, suggesting there was little phylogenetic constraint on fruit evolution at this level. We then surveyed fruit types found in two contrasting habitats: an open habitat including the Indian desert and North American plains and prairies, and a closed forest habitat of Australian tropical forest. The majority of genera in the survey of tropical forests in Australia were fleshy fruit trees, whereas the majority of genera in the survey of prairies and plains in central North America were herbs with capsules and achenes. Both capsules and achenes are frequently dispersed by wind in the open, arid habitat, whereas fleshy fruits are generally dispersed by animals. Since desert and plains tend to provide continuous wind to aid dispersal and there are more abundant mammal and bird dispersers in the closed forest, this survey suggests that fruit evolution was driven at least in part by dispersal agents abundant in particular habitats.     

果实类型和种子传播的进化:系统发育和生态学简论

Success of flowering plants is greatly dependent on effective seed dispersal. Specific fruit types aid different mechanisms of seed dispersal. However, little is known about what evolutionary forces have driven the diversification of fruit types and whether there were phylogenetic constraints on fruit evolution among angiosperm lineages. To address these questions, we first surveyed the orders and families of angiosperms for fruit types and found no clear association between fruit types and major angiosperm lineages, suggesting there was little phylogenetic constraint on fruit evolution at this level. We then surveyed fruit types found in two contrasting habitats an open habitat including the Indian desert and North American plains and prairies, and a closed forest habitat of Australian tropical forest. The majority of genera in the survey of tropical forests in Australia were fleshy fruit trees, whereas the majority of genera in the survey of prairies and plains in central North America were herbs with capsules and achenes. Both capsules and achenes are frequently dispersed by wind in the open, arid habitat, whereas fleshy fruits are generally dispersed by animals. Since desert and plains tend to provide continuous wind to aid dispersal and there are more abundant mammal and bird dispersers in the closed forest, this survey suggests that fruit evolution was driven at least in part by dispersal agents abundant in particular habitats.     

Angiosperm growth habit,dispersal and diversification reconsidered

Summary Previous studies have sought to elucidate the relationship between dispersal mode (biotic versus abiotic) and the taxonomic diversification of angiosperm families, but with ambiguous results. In this study, we propose the hypothesis that the combination of (1) the large seed size required of plants germinating in closed, light-poor environments and (2) the necessity to move disseminules away from the maternal plant in order to avoid intraspecific competition, predation and pathogens should favour biotically-dispersed relative to abiotically-dispersed woody arborescent angiosperms, resulting in higher diversification of the former. In this paper, we seek patterns of diversification that support this hypothesis. We examine the association between dispersal mode, growth habit and taxonomic richness of monocotyledon and dicotyledon families using (1) contingency table analyses to detect the effect of dispersal mode on the relative abundances and diversification of woody versus herbaceous taxa and (2) non-parametric analyses of variance to detect the statistical effect of dispersal mode on taxonomic diversification (mean number of species per genus, genera per family and species per family) in monocot and dicot families dominated by biotic or abiotic dispersal. We found a clear statistical effect of dispersal mode on diversification. Among families of woody dicots, dispersal by vertebrates is associated with significantly higher levels of species per genus, genera per family and species per family than is abiotic dispersal. The same pattern is observed among woody monocots, but is not significant at the 0.05 level. Among families of herbaceous monocots and dicots, the situation is reversed, with abiotically-dispersed families exhibiting higher levels of diversification than vertebrate-dispersed families. When woody and herbaceous families are pooled, there is no association between dispersal mode and diversification. These data coincide with evidence from the fossil record to suggest vertebrate dispersal has positively contributed to the diversification of woody angiosperms. We suggest that vertebrate dispersal may have promoted the diversity of extant taxa by reducing the probability of extinction over evolutionary time, rather than by elevating speciation rates. Our results suggest vertebrate dispersal has contributed to, but does not explainin toto, the diversity of living angiosperms.     

Costs and benefits of non‐random seed release for long‐distance dispersal in wind‐dispersed plant species

The dispersal ability of plants is a major factor driving ecological responses to global change. In wind‐dispersed plant species, non‐random seed release in relation to wind speeds has been identified as a major determinant of dispersal distances. However, little information is available about the costs and benefits of non‐random abscission and the consequences of timing for dispersal distances. We asked: 1) to what extent is non‐random abscission able to promote long‐distance dispersal and what is the effect of potentially increased pre‐dispersal risk costs? 2) Which meteorological factors and respective timescales are important for maximizing dispersal? These questions were addressed by combining a mechanistic modelling approach and field data collection for herbaceous wind‐dispersed species. Model optimization with a dynamic dispersal approach using measured hourly wind speed showed that plants can increase long‐distance dispersal by developing a hard wind speed threshold below which no seeds are released. At the same time, increased risk costs limit the possibilities for dispersal distance gain and reduce the optimum level of the wind speed threshold, in our case (under representative Dutch meteorological conditions) to a threshold of 5–6 m s^–1. The frequency and predictability (auto‐correlation in time) of pre‐dispersal seed‐loss had a major impact on optimal non‐random abscission functions and resulting dispersal distances. We observed a similar, but more gradual, bias towards higher wind speeds in six out of seven wind‐dispersed species under natural conditions. This confirmed that non‐random abscission exists in many species and that, under local Dutch meteorological conditions, abscission was biased towards winds exceeding 5–6 m s^–1. We conclude that timing of seed release can vastly enhance dispersal distances in wind‐dispersed species, but increased risk costs may greatly limit the benefits of selecting wind conditions for long‐distance dispersal, leading to moderate seed abscission thresholds, depending on local meteorological conditions and disturbances.     

杂草种子传播研究进展

种子传播将母株生殖周期的末端与它们后代种群的建立连结了起来,广泛认为,其对植被结构具有深刻的影响。种子传播的整个过程称为种子传播循环。研究表明,杂草种子传播的因子多种多样,包括仅依赖自身来完成的主动传播,以及依赖风、水、动物、人类等外界媒介的被动传播。其中,人类传播杂草种子是影响最广泛的一种,对现代植物的分布格局产生了深刻的影响。杂草种子的传播,对杂草种子库的数量和空间动态影响很大。研究种子传播的主要方法有荧光染料标记法、放射性同位素标记法、稳定同位素分析、分子遗传标记等。结合近几年国内外的研究进展,作者就杂草种子传播对种子库数量和空间动态影响的精确直接研究、杂草种子传播的过程及传播后的命运、杂草种子适应传播的机理、生态控草措施研究、外来杂草入侵蔓延与其种子传播的关系等方面提出了展望。     

The structure and phylogenetic significance of the conifer Pseudohirmerella delawarensis nov. comb. from the Upper Triassic of North America

The conifer genus Pseudohirmerella Arndt is based on ovulate cone scales of the type species, Pseudohirmerella platysperma (Mägdefrau) Arndt, from the Upper Triassic of Germany, which bear five distal lobes and two paired structures represented by bulges on the adaxial surface. The paired structures were first described as seeds, but have recently been re-interpreted as arils by Arndt. Similar scales from the Passaic Formation at Milford, NJ, USA were described as Glyptolepis delawarensis Bock, and the new combination Pseudohirmerella delawarensis is proposed. A relatively complete concept of Pseudohirmerella is presented, which includes ovulate cones, possible associated pollen cones, associated shoots with Pagiophyllum-Brachyphyllum morphotype leaves and associated, anatomically preserved wood. Based on the ovulate cone morphology and the presence of organic matter lining the concavities on the scale rather than the corresponding bulges on the counterparts, it is likely that the supposed seeds or arils of Pseudohirmerella are actually casts of empty, seed-bearing depressions. Cheirolepidiaceous affinities are likely based on ovulate cone scale morphology, persistent pollen cones, foliage type and details of wood anatomy. The derived ovulate cone scale morphology of Pseudohirmerella indicates a substantial, but mostly undocumented, Triassic diversification of the Cheirolepidiaceae.     

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.     

Long-distance seed dispersal in plant populations

Long-distance seed dispersal influences many key aspects of the biology of plants, including spread of invasive species, metapopulation dynamics, and diversity and dynamics in plant communities. However, because long-distance seed dispersal is inherently hard to measure, there are few data sets that characterize the tails of seed dispersal curves. This paper is structured around two lines of argument. First, we argue that long-distance seed dispersal is of critical importance and, hence, that we must collect better data from the tails of seed dispersal curves. To make the case for the importance of long-distance seed dispersal, we review existing data and models of long-distance seed dispersal, focusing on situations in which seeds that travel long distances have a critical impact (colonization of islands, Holocene migrations, response to global change, metapopulation biology). Second, we argue that genetic methods provide a broadly applicable way to monitor long-distance seed dispersal; to place this argument in context, we review genetic estimates of plant migration rates. At present, several promising genetic approaches for estimating long-distance seed dispersal are under active development, including assignment methods, likelihood methods, genealogical methods, and genealogical/demographic methods. We close the paper by discussing important but as yet largely unexplored areas for future research.     

Overseas seed dispersal by migratory birds

Long-distance dispersal (LDD) promotes the colonization of isolated and remote habitats, and thus it has been proposed as a mechanism for explaining the distributions of many species. Birds are key LDD vectors for many sessile organisms such as plants, yet LDD beyond local and regional scales has never been directly observed nor quantified. By sampling birds caught while in migratory flight by GPS-tracked wild falcons, we show that migratory birds transport seeds over hundreds of kilometres and mediate dispersal from mainland to oceanic islands. Up to 1.2% of birds that reached a small island of the Canary Archipelago (Alegranza) during their migration from Europe to Sub-Saharan Africa carried seeds in their guts. The billions of birds making seasonal migrations each year may then transport millions of seeds. None of the plant species transported by the birds occurs in Alegranza and most do not occur on nearby Canary Islands, providing a direct example of the importance of environmental filters in hampering successful colonization by immigrant species. The constant propagule pressure generated by these LDD events might, nevertheless, explain the colonization of some islands. Hence, migratory birds can mediate rapid range expansion or shifts of many plant taxa and determine their distribution.     

动物对松属植物种子的传播作用研究进展

松属植物约110种,根据种子传播方式可分为风传播松和动物传播松。风传播松占绝大多数,种子多具有适应风力的翅。动物传播松大约23种,都具有大、可食用、无翅或短翅的种子,无法借助风力传播。动物传播松的分布生境多为贫瘠的山地,而且多位于高海拔地区。目前已知9种松树的动物传播种类,其余14种可推测为动物传播。动物传播者包括鸦科鸟类和啮齿类动物,动物将获得的种子分散贮藏,未被重取的种子可能萌发,完成传播。动物传播是定向传播,微生境多适合种子萌发。啮齿类的传播距离可达数10 m,而鸦科鸟类的传播距离可达数公里。动物传播的松树会出现树丛和多树干现象,一般由同一贮点内贮藏的多粒种子萌发造成的。动物贮藏的种子大部分被重取,称传播后取食。一些具有大种子的风传播松在种子落地后,啮齿类和鸟类会再次埋藏而形成二次传播,可看做是一个单独的传播类型,即风-动物传播松。动物传播者与依赖传播松树之间可看作是互利共生关系。     

Recruitment trade-offs and the evolution of dispersal mechanisms in plants

In this study we place seed size vs. seed number trade-offs in the context of plant dispersal ability. The objective was to suggest explanations for the evolution of different seed dispersal mechanisms, in particular fleshy fruits, wind dispersal and the maintenance of unassisted dispersal. We suggest that selection for improved dispersal may act either by increasing the intercept of a dispersal curve (log seed number vs. distance) or by flattening the slope of the curve. 'Improved dispersal' is defined as a marginal increase in the number of recruits sited at some (arbitrary) distance away from the parent plant. Increasing the intercept of the dispersal curve, i.e. producing more seeds, is associated with a reduction in seed size, which in turn affects the recruitment ability, provided that this ability is related to seed size. If recruitment is related to seed size there will be a recruitment cost of evolving increased seed production. On the other hand, a flattening of the slope by evolving dispersal attributes is likely to be associated with a fecundity cost. An exception is wind dispersal where smaller (and hence more numerous) seeds may lead to more efficient dispersal. We derive two main predictions: If recruitment is strongly related to seed size, selection for improved dispersal acts on the slope of the dispersal curve, i.e. by favouring evolution of dispersal attributes on seeds or fruits. If, on the other hand, recruitment is only weakly related to seed size (or not related, or negatively related), selection for improved dispersal favours increased seed production. Despite its simplicity, the model suggests explanations for (i) why so many plant species lack special seed dispersal attributes, (ii) differences in dispersal spectra among plant communities, and (iii) adaptive radiation in seed size and dispersal attributes during angiosperm evolution. This revised version was published online in July 2006 with corrections to the Cover Date.     

Directed seed dispersal towards areas with low conspecific tree density by a scatter‐hoarding rodent

Scatter‐hoarding animals spread out cached seeds to reduce density‐dependent theft of their food reserves. This behaviour could lead to directed dispersal into areas with lower densities of conspecific trees, where seed and seedling survival are higher, and could profoundly affect the spatial structure of plant communities. We tested this hypothesis with Central American agoutis and Astrocaryum standleyanum palm seeds on Barro Colorado Island, Panama. We radio‐tracked seeds as they were cached and re‐cached by agoutis, calculated the density of adult Astrocaryum trees surrounding each cache, and tested whether the observed number of trees around seed caches declined more than expected under random dispersal. Seedling establishment success was negatively dependent on seed density, and agoutis carried seeds towards locations with lower conspecific tree densities, thus facilitating the escape of seeds from natural enemies. This behaviour may be a widespread mechanism leading to highly effective seed dispersal by scatter‐hoarding animals.     

Inter‐group variability in seed dispersal by white‐handed gibbons in mosaic forest

Seed dispersers, like white‐handed gibbons (Hylobates lar), can display wide inter‐group variability in response to distribution and abundance of resources in their habitat. In different home ranges, they can modify their movement patterns along with the shape and scale of seed shadow produced. However, the effect of inter‐group variability on the destination of dispersed seeds is still poorly explained. In this study, we evaluate how seed dispersal patterns of this arboreal territorial frugivore varies between two neighboring groups, one inhabiting high quality evergreen forest and one inhabiting low quality mosaic forest. We predicted a difference in seed dispersal distance between the two groups (longer in the poor quality forest). We hypothesized that this difference would be explained by differences in home range size, daily path length, and ranging tortuosity. After 6 months of data collection, the evergreen group had a smaller home range (12.4 ha) than the mosaic group (20.9 ha), significantly longer daily path lengths (1507 m vs. 1114 m respectively) and greater tortuosity (39.1 vs. 16.1 respectively). Using gut passage times and displacement rates, we estimated the median seed dispersal distance as 163 m for the evergreen group (high quality forest) and of 116 m for the mosaic group (low quality forest). This contradiction with our initial prediction can be explained in term of social context, resource distribution, and habitat quality. Our results indicate that gibbons are dispersers of seeds between habitats and that dispersal distances provided by gibbons are influenced by a range of factors, including habitat and social context.     

Broad-scale reciprocity in an avian seed dispersal mutualism

Aim Coevolved relationships between individual species of birds and plants rarely occur in seed dispersal mutualisms. This study evaluates whether reciprocal relationships may occur between assemblages of bird and plant species. Location Vancouver Island, British Columbia, Canada (48°50′‐N, 125°22′‐W). Methods The distribution and fruiting phenologies of seven shrub species were compared to seasonal changes in habitat selection and seed dispersal by six fruit‐eating bird species. Results Shrub species inhabiting forest understorey habitat had earlier fruiting phenologies than shrub species inhabiting forest edge habitat along lake and bog margins. Birds showed a parallel pattern in habitat selection, being more abundant in the forest understorey early in the fruiting season, and more abundant in the forest edge later in the season. Rates of seed deposition covaried with avian habitat selection, in such a way that birds directed seed dispersal into habitats preferred by shrubs. Conclusions These results depict a broad‐scale pattern in the abundance of birds and fruits indicative of reciprocal interactions. Seasonal changes in seed dispersal to each habitat appear to reinforce the relationship between shrub habitat affinities and fruiting phenologies. Phenological differences between habitats may also reinforce seasonal changes in avian habitat selection. Therefore, although reciprocal interactions between pairs of bird and plant species are rare, broad‐scale reciprocal relationships may occur between assemblages of bird and plant species.