How seed traits predict floating times: a biophysical process model for hydrochorous seed transport behaviour in fluvial systems

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Reproductive strategy, clonal structure and genetic diversity in populations of the aquatic macrophyte Sparganium emersum in river systems

Many aquatic and riparian plant species are characterized by the ability to reproduce both sexually and asexually. Yet, little is known about how spatial variation in sexual and asexual reproduction affects the genotypic diversity within populations of aquatic and riparian plants. We used six polymorphic microsatellites to examine the genetic diversity within and differentiation among 17 populations (606 individuals) of Sparganium emersum, in two Dutch-German rivers. Our study revealed a striking difference between rivers in the mode of reproduction (sexual vs. asexual) within S. emersum populations. The mode of reproduction was strongly related to locally reigning hydrodynamic conditions. Sexually reproducing populations exhibited a greater number of multilocus genotypes compared to asexual populations. The regional population structure suggested higher levels of gene flow among sexually reproducing populations compared to clonal populations. Gene flow was mainly mediated via hydrochoric dispersal of generative propagules (seeds), impeding genetic differentiation among populations even over river distances up to 50 km. Although evidence for hydrochoric dispersal of vegetative propagules (clonal plant fragments) was found, this mechanism appeared to be relatively less important. Bayesian-based assignment procedures revealed a number of immigrants, originating from outside our study area, suggesting intercatchment plant dispersal, possibly the result of waterfowl-mediated seed dispersal. This study demonstrates how variation in local environmental conditions in river systems, resulting in shifting balances of sexual vs. asexual reproduction within populations, will affect the genotypic diversity within populations. This study furthermore cautions against generalizations about dispersal of riparian plant species in river systems.     

Influence of an ecosystem engineer,the emergent macrophyte Sparganium erectum,on seed trapping in lowland rivers and consequences for landform colonisation

1. Plant physical ecosystem engineers can influence vegetation population and community dynamics by modifying, maintaining or creating habitats. They may also have the potential to act upon biotic processes, such as seed dispersal. 2. Examples exist of reduction in seed dispersal distances in vegetated compared to unvegetated terrestrial environments, and concentration of seed deposits associated with plant patches. Such effects in aquatic environments have been little studied, but the engineering effect of plant patches on patterns of flow velocity and sediment deposition in streams suggests that they may play a similar role. 3. In this study, we assess the potential of an emergent aquatic species, Sparganium erectum, to play a role in physically modifying river habitats and trapping seeds by examining patterns of seed deposition and substrate type in 47 river reaches across England and southern Scotland, U.K. 4. Areas of the river channel within or adjacent to S. erectum patches harboured more plant seeds and more species than unvegetated areas and had finer, sandier substrates with higher organic matter, total nitrogen and total phosphorus content. Most seed species were competitive, indicating that they were well suited to colonise the competitive environment of an S. erectum patch, and could potentially further stabilise accumulated sediments and contribute to landform development. 5. We demonstrate that S. erectum patches influence both the physical environment and the retention of seeds, in consistent patterns across the channel bed, for a range of lowland rivers that vary in stream power and geology and which can be expected to vary in levels of supply of fine sediment and seeds. 6. Our findings support the hypothesis that the fundamental influence of a riverine ecosystem‐engineering species on slowing fluid flow links the habitat creation process of sediment sorting and retention to seed trapping. We suggest the process is applicable to a wide range of aquatic and riparian vegetation. We also suggest that the mono‐specific and competitive growth, which is typical of these engineering species, will strongly influence the recruitment of trapped seeds.     

Water-borne seed transport and seed deposition during flooding in a small river-valley in Northern Germany

Water-borne seed transport and seed deposition during flooding were studied in the Upper Eider river (N-Germany) by direct sampling of the rivers seed content with aquatic seed traps and by analysing the number of deposited seeds on sedimentation mats which were exposed near the river on the soil surface during a flooding period of approx. three weeks.The number of seeds which were transported at the surface of the river Eider was continuously analysed by four aquatic seed traps for a period of 20 weeks (July–December 2000). To test the capture rate of these traps, a recapture experiment with colour marked seeds of Helianthus annuus L. was carried out. During the investigation period approx. 9000 seeds of 76 species were captured by the four aquatic seed traps. The number of trapped seeds varied both spatially (across the river profile) and temporally. Considering this variation and the capture rate of the traps, the water-borne seed transport was estimated to be 3139 seeds per week and meter of the river profile.The seed deposition during a flood in early spring 2002 was analysed by using 20 sedimentation mats. To distinguish effects of seed dispersal into patches from outside from seed rearrangement within patches, the water-borne seed transport was excluded from one half of the mats by fencing them with a woven fabric which was permeable for water but not for floating seeds. Outside of the exclosures 152 viable seeds of 26 species were deposited on the sedimentation mats while only one single seedling was found on mats from which water-borne seed transport was excluded.The results demonstrate that hydrochorous dispersal processes might play an important role in connecting otherwise fragmented populations in periodically flooded habitats along rivers.     

Gene flow and genetic structure of the aquatic macrophyte Sparganium emersum in a linear unidirectional river

1. River systems offer special environments for the dispersal of aquatic plants because of the unidirectional (downstream) flow and linear arrangement of suitable habitats.
2. To examine the effect of this flow on microevolutionary processes in the unbranched bur-reed ( Sparganium emersum ) we studied the genetic variation within and among nine (sub)populations along a 103 km stretch of the Niers River (Germany–The Netherlands), using amplified fragment length polymorphisms.
3. Genetic diversity in S. emersum populations increased significantly downstream, suggesting an effect of flow on the pattern of intrapopulation genetic diversity.
4. Gene flow in the Niers River is asymmetrically bidirectional, with gene flow being approximately 3.5 times higher in a downstream direction. The observed asymmetry is probably caused by frequent hydrochoric dispersal towards downstream locations on the one hand, and sporadic zoochoric dispersal in an upstream direction on the other. The spread of vegetative propagules (leaf and stem fragments) is probably not an important mode of dispersal for S. emersum , suggesting that gene flow is mainly via seed dispersal. Realized dispersal distances exceeded 60 km, revealing a potential for long-distance dispersal in S. emersum .
5. There was no correlation between geographical and genetic distances among the nine S. emersum populations (i.e. no isolation by distance), which may be due to the occurrence of long-distance dispersal and/or colonization and extinction dynamics in the Niers River.
6. Overall, the genetic population structure and regional dispersal patterns of S. emersum in the Niers River are best explained by a linear metapopulation model. Our study shows that flow can exert a strong influence on population genetic processes of plants inhabiting stream systems.     

The influence of spatial scale on the genetic structure of a widespread tropical wetland tree, Pterocarpus officinalis (Fabaceae)

Identifying factors that cause genetic differentiation in plant populations and the spatial scale at which genetic structuring can be detected will help to understand plant population dynamics and identify conservation units. In this study, we determined the genetic structure and diversity of Pterocarpus officinalis, a widespread tropical wetland tree, at three spatial scales: (1) drainage basin “watershed” (<10 km), (2) within Puerto Rico (<100 km), and (3) Caribbean-wide (>1000 km) using AFLP. At all three spatial scales, most of the genetic variation occurred within populations, but as the spatial scale increased from the watershed to the Caribbean region, there was an increase in the among population variation (Φ_ST=0.19 to Φ_ST=0.53). At the watershed scale, there was no significant differentiation (P=0.77) among populations in the different watersheds, although there was some evidence that montane and coastal populations differed (P<0.01). At the island scale, there was significant differentiation (P<0.001) among four populations in Puerto Rico. At the regional scale (>1000 km), we found significant differentiation (P<0.001) between island and continental populations in the Caribbean region, which we attributed to factors associated with the colonization history of P. officinalis in the Neotropics. Given that genetic structure can occur from local to regional spatial scales, it is critical that conservation recommendations be based on genetic information collected at the appropriate spatial scale.     

Dispersal of Amazonian Trees: Hydrochory in Swartzia polyphylla1

Hydrochory was investigated in the seeds of the Amazonian floodplain tree, Swartzia polyphylla, in which pods open on the tree to release one large seed. Seeds collected from beach drift along the Rio Negro showed a high percentage of floaters (82%). Yet most seeds sank following collection from: adult trees (89%), unflooded ground under adults (96%), and flooded ground under adults (86%). The specific gravity of the seeds was near that of water, 1.04 ± 0.03 for sinkers and 0.98 ± 0.02 for floaters. The ability to float was correlated directly with the volume of the air pocket between the two cotyledons, which varied from 5.6 to 20.5 percent of the total seed volume. In a long-term floatation test lasting 81 days, 45 percent of the seeds never floated, 33 percent always floated, and 22 percent first sank for one week and then floated for at least one month. Seeds that never floated eventually rotted, but not until days 63-73. Seeds that were floating at day 81, regardless of how long they had been floating, were placed on moistened filter paper for 18 days during which time 36 percent germinated, 45 percent rotted, and 19 percent did neither but remained viable. These results suggest that S. polyphylla achieves dimorphism in flotation of its seeds, some sinking and some floating, by producing seeds of continuous variation in specific gravity around a mean close to 1.00. Seeds that float can be dispersed long distances along river margins, while those that sink may be moved only marginally from the parent tree.