DNA fingerprinting validates seed dispersal curves from observational studies in the neotropical legume parkia

Background

Determining the distances over which seeds are dispersed is a crucial component for examining spatial patterns of seed dispersal and their consequences for plant reproductive success and population structure. However, following the fate of individual seeds after removal from the source tree till deposition at a distant place is generally extremely difficult. Here we provide a comparison of observationally and genetically determined seed dispersal distances and dispersal curves in a Neotropical animal-plant system.

Methodology/Principal Findings

In a field study on the dispersal of seeds of three Parkia (Fabaceae) species by two Neotropical primate species, Saguinus fuscicollis and Saguinus mystax, in Peruvian Amazonia, we observationally determined dispersal distances. These dispersal distances were then validated through DNA fingerprinting, by matching DNA from the maternally derived seed coat to DNA from potential source trees. We found that dispersal distances are strongly right-skewed, and that distributions obtained through observational and genetic methods and fitted distributions do not differ significantly from each other.

Conclusions/Significance

Our study showed that seed dispersal distances can be reliably estimated through observational methods when a strict criterion for inclusion of seeds is observed. Furthermore, dispersal distances produced by the two primate species indicated that these primates fulfil one of the criteria for efficient seed dispersers. Finally, our study demonstrated that DNA extraction methods so far employed for temperate plant species can be successfully used for hard-seeded tropical plants.     

Cache site selection by chipmunks (Tamias spp.) and its influence on the effectiveness of seed dispersal in Jeffrey pine (Pinus jeffreyi)

The effectiveness of Jeffrey pine (Pinus jeffreyi) seed dispersal performed by seed-caching yellow pine chipmunks (Tamias amoenus) and lodgepole chipmunks (Tamias speciosus) was compared to that of wind dispersal in the Sierra Nevada of western Nevada. Wind-dispersed seeds typically fall under or near the parent tree. Chipmunks removed 90 and 97% of 1064 radioactive seeds from each of two simulated wind-dispersed seed shadows in less than 24 h. Wind-dispersed seeds were deployed within 12 m of the two source trees, but chipmunk caches were found from 2–69 m from the trees. Chipmunks carried nearly all seeds away from source trees, greatly reducing the density of seeds under and near source trees. Caches contained from 1–35 seeds and most were buried 7–21 mm deep. Chipmunks cached in open bitterbrush shrubland with mineral soils much more than expected and cached in closed-canopy Jeffrey pine and lodgepole pine forests with thick needle litter much less than expected. Many Jeffrey pine seedlings and saplings grow in the bitterbrush habitat and few grow in the pine forests. Ten and 20% of the original caches survived until April, the time of seed germination, at the two sites. The movement of wind-dispersed seeds is random relative to environmental variables important in seedling survival, and the wind in coniferous forests cannot quickly bury seeds. The quality of seed dispersal rendered by chipmunks was superior to that provided by the wind because the chipmunks quickly harvested seeds on the ground, moved them away from source trees, and buried them in the ground in habitats and microhabitats where they were more likely to establish new seedlings. The increased quality of seed dispersal provided by animals relative to the wind may help explain why over twenty species of pines have evolved seeds and cones that are adapted for dispersal by seed-caching animals.     

Splash seed dispersal by raindrops

Splash seed dispersal by raindrops was investigated for plants in southern Japan. Nine families, 10 genera and 19 species were confirmed as raindrop-dispersed plants. The 10 genera were Gentiana, Gratiola, Chrysosplenium, Mazus, Mitella, Ophiorrhiza, Sagina, Sedum, Trigonotis and Veronica. The method of splash rain dispersal in these species was clarified. Raindrop-dispersed species were all small herbaceous plants with a vertical pedicel and an apically opening fresh capsule when the seeds mature. Open capsules were cup-shaped or boat-shaped and can accommodate raindrops easily. The raindrops splashed the seeds from the capsule. In general, the seeds weighed very little, but they were heavier than powder or dust seeds dispersed by wind. A strong negative correlation was found between seed weight and the number of seeds per capsule. In the case of Trigonotis brevipes (Maxim.) Maxim., raindrops were received into the cup-shaped calyx-tube and dispersed the fruitlets. Some species, such as Gentiana thunbergii (G. Don) Griseb., Gentiana zollingeri Fawcett and Ophiorrhiza japonica Blume, had hydroscopic movement capsules that opened widely only when wet. Raindrop-dispersed plants were found in various habitats. For example, some plants grew together on rocks along the mountain torrents where splash water could easily be caught. The results of the laboratory and field experiments indicated that the dispersal distance of seeds by raindrops was 1m or less. For small herbaceous plants, splash dispersal by rain might be an effective and advantageous method of seed dispersal because dispersal is not affected by plant height.     

Growth form distribution and genetic relationships in tree clusters of Pinus flexilis,a bird-dispersed pine

Seed dispersal by the Clark's nutcracker (Nucifraga columbiana Wilson) may markedly influence the growth form and genetic population structure of limber pine (Pinus flexilis James). The nutcracker buries clusters of seeds in subterranean caches; germination of clustered seeds often results in a growth form characterized by two or more genetically distinct trees with fused or contiguous trunks (tree clusters). The occurrence of a morphologically similar form, the multi-trunk tree (a single genet branched near the base), as well as the typical single-trunked tree, complicates the study of limber pine populations. We examined growth form distribution and genetic relationships in tree clusters in limber pine populations at four elevations (from 2585 m to 3460 m) in the Colorado Front Range. At three study areas, relative occurrence of limber pine growth forms, as well as that of associated pines, was examined by a point-centered quarter survey. From the four study areas, we collected foliage from each trunk from a total of 74 clumps (combined tree clusters and multi-trunk trees) in order to differentiate the two growth forms using starch gel protein electrophoresis. Tree clumps were significantly more common in limber pine than in ponderosa or lodgepole pine (P<0.010). Although single-trunk limber pine was the most common growth form, except at the highest elevation, both multi-trunk trees and tree clusters were present in each stand. Tree clusters were estimated to comprise about 20% of the tree sites in each limber pine stand; the estimated proportion of multi-trunk trees varied by site from 5% to 77%. Trees in clusters were related, on average, as half to full siblings (mean r=0.43), but were unrelated to trees in other clusters (mean r=0.01). Electrophoretic analysis suggests possible genetic differentiation in limber pine that may be the result of different selection pressures on the growth forms.     

The Effects of Habitat on Dispersal Patterns of an Invasive Thistle, Cynara Cardunculus

Dispersal of offspring is a critical step in the spread of invasive species, yet dispersal patterns are seldom well studied, inhibiting effective management and ecological understanding of invasions. Dispersal patterns can be affected by characteristics of the parent plant and by climatic or site characteristics, including community vegetation structure, but these factors have not been studied in the context of plant invasion processes. Cynara cardunculus (L.) is a polycarpic perennial thistle invasive in coastal grasslands in California that produces large numbers of wind-dispersed seeds. This study quantified and compared C. cardunculus dispersal patterns in an exotic grassland (vegetated site) and an agricultural field (non-vegetated site). Seed size variation in C. cardunculus was also quantified within inflorescences, between inflorescences, and between years of production, and compared across dispersal distances. Results indicate that dispersal distance increased dramatically from less than 20 m in the vegetated site to more than 40 m in the non-vegetated site. Plants producing fewer seeds produced heavier seeds and dispersal distance decreased over time, but seed size was not related to dispersal distance, which may serve to spread the risk to seeds or seedlings across the environment. C. cardunculus has great potential for dispersal in open areas such as agricultural fields or disturbed sites, but may be limited in highly structured natural communities. Management of C. cardunculus and other wind-dispersed plant invasions may be improved by prioritizing populations with open or disturbed areas downwind and minimizing the removal of vegetation during dispersal.     

Primate seed dispersal leaves spatial genetic imprint throughout subsequent life stages of the Neotropical tree Parkia panurensis

Key message

The Neotropical tree Parkia panurensis shows a spatial genetic structure from the seed to the adult stage that is most likely the outcome of the seed dispersal provided by primates.

Abstract

Seed dispersal and pollination determine the gene flow within plant populations. In addition, seed dispersal creates the template for subsequent stages of plant recruitment. Therefore, the question arises whether and how seed dispersal affects the spatial genetic structure (SGS) of plant populations. In this study, we used microsatellites to analyse the SGS of the Neotropical tree Parkia panurensis (Fabaceae). This plant species is a major food resource for primates and its seeds are mainly dispersed by primates. Seeds were collected during behavioural observations of a tamarin mixed-species troop in north-eastern Peru. Additionally, leaf samples of juveniles and of adults trees of this species were collected throughout the home range of the tamarin troop. A significant SGS for embryos (located within the dispersed seeds) and for non-reproductive plants are found up to a distance of 300 m. This matches the distance within which most seeds are dispersed. In the adult stage, the scale of a significant SGS is reduced to 100 m. While we cannot explain this scale reduction, our study provides the first evidence that primate seed dispersal does influence the SGS of a tropical tree species.     

Agouti reintroduction recovers seed dispersal of a large-seeded tropical tree

The aim of animal reintroductions has mainly been species recovery; only few reintroduction initiatives focus on ecosystem restoration. Therefore, reintroduction consequences on ecological interactions are seldom assessed. We used the interaction between a reintroduced population of agoutis (Dasyprocta leporina) and a vulnerable tropical endemic tree (Joannesia princeps) to examine reintroduction effects on seed dispersal and seedling establishment. To test the outcomes of this interaction, we tracked seeds of J. princeps in two adjacent forest areas with and without reintroduced agoutis. We also assessed if dispersal distances affected seedling survival. To determine seed fate and dispersal distance, we used spool-and-line tracking, together with camera traps to identify dispersers. Agoutis were the only species removing J. princeps seeds, thus dispersal only occurred where agoutis had been reintroduced; in the area without agoutis, all seeds remained intact on the soil, even one year after the experiment's beginning. At the reintroduction area, most seeds were preyed upon by agoutis but 7% remained dispersed and 2% germinated after ten months. Only seeds buried by agoutis were able to germinate. Most dispersed seeds were dispersed 15 m or farther and longer dispersal distances benefited J. princeps, since seedlings farther from a conspecific adult tree had greater survival probability. Agoutis were also seen burying seeds of two other plant species; these mammals have the potential to benefit dozens of large-seeded species in our study system. Agouti reintroduction thus exemplifies the value of trophic rewilding programs to re-establish ecological interactions and restore ecosystem functioning.     

Biological and environmental effects on fine-scale seed dispersal of an invasive tree in a secondary subtropical forest

Dispersal abilities of invading species emerge from the interaction between the species and some features of the target community. Ligustrum lucidum is a tree species invading different ecosystems. Major spatial patterns of Ligustrum invasions and their ecological consequences have been analyzed, but no study addressed the dispersal process at a fine scale, assessing the effects of different biological and environmental factors. Ligustrum lucidum is an ornithochoric species. The structure of the environment determines bird movements and thus affects seed dispersal. We used inverse modeling to analyze bird-mediated dispersal of L. lucidum seeds in a secondary Yungas forest and surrounding crop-fields. We assessed the effects of egestion mode (regurgitation and defecation) and tree density (as an environment character) on seed dispersal. Seed dispersal presented different spatial patterns depending on the egestion mode. Tree density was positively associated with the number of regurgitated dispersed seeds and negatively associated with the number of defecated dispersed seeds. In both cases, dispersal distance increased in open areas, but absence of perches inhibited seed arrival. Thus, spread of L. lucidum is facilitated in open areas with some trees; inside the native forest, short distance dispersal facilitates the gradual invasion by this exotic species. Our results suggest that processes like crop abandonment and forest succession, which are active in subtropical montane systems, may facilitate L. lucidum invasion. Our seed dispersal models should be combined with actual distribution maps of L. lucidum to identify areas vulnerable to new invasions.     

Larger seeds in tropical floras: consistent patterns independent of growth form and dispersal mode

Many species of tropical moist forests have large seeds compared to those found in temperate floras. This could be attributed to a prevalence of woody growth forms, or adaptation to dispersal by vertebrates, or the dense shade of a closed canopy, rather than to an intrinsic tropical v. temperate difference. We compared tropical v. temperate seed mass data at two geographic scales; firstly within Australia between tropical, subtropical and temperate open woodlands, then between six tropical and temperate datasets from five continents. Within Australia we found that seed mass increased with decreasing latitude in ten out of eleven growth form-dispersal mode combinations: only wind-dispersed graminoids showed no significant effect. While the pattern of generally larger seeds in the tropical flora was associated with tropical families tending to have larger seeds than temperate families, we also found that tropical species had larger seeds than temperate cofamilials. As a preliminary test of the generality of these findings we included a further four tropical and temperate data sets in a second analysis. Average tropical seed masses were larger than temperate seed masses in all testable growth form-dispersal mode combinations, and statistically larger in five out of seven cases: unassisted, vertebrate- and wind-dispersed woody species, and unassisted and vertebrate-dispersed forbs. No difference was found for wind-dispersed forbs or for graminoids lacking dispersal structures. That an effect was found in these combinations rules out explanations based solely on characteristics of tropical v. temperate dispersers, although this may be a contributing factor. Instead we suggest that some aspect of tropical systems, for example higher metabolic costs of seedling production, has selected for a general increase in seed mass among tropical taxa.     

Early stages of secondary succession on abandoned cropland in north-east Borneo Island

A study on community structure and floristic change in the early stages of the tropical old-field succession in an intensive shifting cultivation area, Sabah, north-east Borneo Island, was conducted in March and September 1997. Dense therophytic communities were established soon after abandonment, and rapidly changed: namely, from Galinsoga parviflora (in approximately 1–2 months) to Crassocephalum crepidioides (in approximately 3–5 months) and further to Conyza sumatrensis (in approximately 1 year). These therophytic communities were replaced by perennial grass and shrub species, such as Imperata cylindrica and Eupatorium odoratum, within 3 years after abandonment. At this stage, a few pioneer tree species such as Trema orientalis made a sparse canopy layer approximately 3–4 m high. The three pioneer annuals have wind-dispersed small seeds. The plant densities of the three annuals did not appear to change between areas fallow for 1 month and 4 months. Almost all of the seedlings of the three annuals might simultaneously have invaded the fallowed field soon after abandonment. The three pioneer annuals take a different amount of time to reach their respective mature sizes. This resulted in the rapid successional changes in the tropical old-field succession. The seed-bank annuals, whose seeds germinate after the winter pre-chilling, occurred in the temperate old-fields as the first-year pioneers under seasonal condition. Intermittent shifting cultivation and tropical conditions without seasonality favour the occurrence of wind-dispersed annuals with a spatially fugitive strategy.     

Destination‐based seed dispersal homogenizes genetic structure of a tropical palm

As the dominant seed dispersal agents in many ecosystems, frugivorous animals profoundly impact gene movement and fine‐scale genetic structure of plants. Most frugivores engage in some form of destination‐based dispersal, in that they move seeds towards specific destinations, resulting in clumped distributions of seeds away from the source tree. Molecular analyses of dispersed seeds and seedlings suggest that destination‐based dispersal may often yield clusters of maternal genotypes and lead to pronounced local genetic structure. The long‐wattled umbrellabird Cephalopterus penduliger is a frugivorous bird whose lek mating system creates a species‐specific pattern of seed dispersal that can potentially be distinguished from background dispersal processes. We used this system to test how destination‐based dispersal by umbrellabirds into the lek affects gene movement and genetic structure of one of their preferred food sources Oenocarpus bataua, a canopy palm tree. Relative to background dispersal processes, umbrellabird mating behaviour yielded more diverse seed pools in leks that included on average five times more seed sources and a higher incidence of long‐distance dispersal events. This resulted in markedly lower fine‐scale spatial genetic structure among established seedlings in leks than background areas. These species‐specific impacts of destination‐based dispersal illustrate how detailed knowledge of disperser behaviour can elucidate the mechanistic link driving observed patterns of seed movement and genetic structure.     

Estimation of the retention times and distances of seed dispersed by two monkey species, Alouatta seniculus and Lagothrix lagotricha, in a Colombian forest

Isolated-bout method to estimate the retention times and dispersal distances was applied to the seed dispersal by red howler monkeys (Alouatta seniculus) and Humboldts woolly monkeys (Lagothrix lagotricha) in a lowland tropical forest at La Macarena, on the border of the Macarena and Tinigua National Parks, the Department of Meta, Colombia. Continuous observations were made on the feeding and ranging behavior of well-habituated troops of howler monkeys and woolly monkeys as well as continuous collection of monkeys feces. We selected out the isolated-bout as a feeding bout on the specific species that was only once recorded within 48 h before the seeds of that species appeared in the feces of monkeys. In that case, the seeds were strongly suggested to come from that isolated bout. Then retention times, route seed dispersal distances and direct seed dispersal distances were estimated. Howler monkeys, which are regarded as generalist herbivores, showed longer retention times and dispersal distances along monkeys route than did woolly monkeys, which are specialist frugivores. However, the direct distances that seeds were carried from the mother tree were not significantly greater for howler monkeys than for woolly monkeys. This shows that both retention time and movement patterns by the monkeys, especially the total ranging area, influence the direct distance that seeds are carried from the mother tree.     

Seed dispersal anachronisms: rethinking the fruits extinct megafauna ate

Background

Some neotropical, fleshy-fruited plants have fruits structurally similar to paleotropical fruits dispersed by megafauna (mammals >10³ kg), yet these dispersers were extinct in South America 10–15 Kyr BP. Anachronic dispersal systems are best explained by interactions with extinct animals and show impaired dispersal resulting in altered seed dispersal dynamics.

Methodology/Principal Findings

We introduce an operational definition of megafaunal fruits and perform a comparative analysis of 103 Neotropical fruit species fitting this dispersal mode. We define two megafaunal fruit types based on previous analyses of elephant fruits: fruits 4–10 cm in diameter with up to five large seeds, and fruits >10 cm diameter with numerous small seeds. Megafaunal fruits are well represented in unrelated families such as Sapotaceae, Fabaceae, Solanaceae, Apocynaceae, Malvaceae, Caryocaraceae, and Arecaceae and combine an overbuilt design (large fruit mass and size) with either a single or few (<3 seeds) extremely large seeds or many small seeds (usually >100 seeds). Within-family and within-genus contrasts between megafaunal and non-megafaunal groups of species indicate a marked difference in fruit diameter and fruit mass but less so for individual seed mass, with a significant trend for megafaunal fruits to have larger seeds and seediness.

Conclusions/Significance

Megafaunal fruits allow plants to circumvent the trade-off between seed size and dispersal by relying on frugivores able to disperse enormous seed loads over long-distances. Present-day seed dispersal by scatter-hoarding rodents, introduced livestock, runoff, flooding, gravity, and human-mediated dispersal allowed survival of megafauna-dependent fruit species after extinction of the major seed dispersers. Megafauna extinction had several potential consequences, such as a scale shift reducing the seed dispersal distances, increasingly clumped spatial patterns, reduced geographic ranges and limited genetic variation and increased among-population structuring. These effects could be extended to other plant species dispersed by large vertebrates in present-day, defaunated communities.     

Seed Rain and Seed Dispersal Agents in Pure and Mixed Plantations of Native Trees and Abandoned Pastures at La Selva Biological Station, Costa Rica

Lack of seed dispersal can be an important obstacle to natural regeneration (NR) of degraded pastures in the humid tropics. Tree plantations can facilitate secondary forest succession by attracting seed dispersal agents from nearby forests. We studied seed rain and seed dispersal agents in 12–13 years old pure and mixed native tree plantations at La Selva Biological Station, Costa Rica from July to December 2004. Plantations of Balizia elegans (5,522), Dipteryx panamensis (2,263), and Jacaranda copaia (2,091) had the greatest total seed abundance; treatments with the least total seed abundance were Calophyllum brasiliense (56), nonplanted abandoned pasture control 2 (353), Mixed Species 2 (389), and control 1 (836). Plantations of J. copaia and Hyeronima alchorneoides had the greatest seed species richness density, whereas the lowest seed species richness was found in the control treatments. The NR plots had more seeds dispersed by wind, whereas in the plantations, the most important dispersal agents were birds and mammals. The most abundant seeds were those of Miconia spp. (14,492), Psychotria bracheata (2,252), and the Poaceae family (1,346), all species from early successional stages. Plantations of native species are effective in attracting seed dispersal agents and thus facilitating regeneration of degraded pasturelands in the region.     

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.     

AN ANALYSIS OF VARIABILITY IN SEED SETTLING VELOCITIES OF SEVERAL WIND-DISPERSED ASTERACEAE

Dispersal is an important life history component. Seed settling velocity may be a useful surrogate for the measurement of dispersal ability in wind-dispersed plants, particularly those whose seeds have plumose dispersal structures. I measured settling velocities on seeds of eight species of Asteraceae, including annuals, biennials, and perennials, and including both native and introduced species. The species are Aster exilis, Picris echioides, Chrysopsis villosa, Heterotheca grandiflora, Conyza bonariensis, Sonchus oleraceous, Senecio vulgaris, and Taraxacum officinale. From these data I estimated components of total variation in seed settling velocities due to differences among species, among plants within species, and among inflorescences and seeds within plants. Significant amounts of variability were found at all levels. Contrasts among mean settling velocities showed that the five introduced species have lower settling velocities than the three native species; this result continues to be true when annuals are considered separately from biennials and perennials. Also, over all eight species, annuals have lower settling velocities than biennials and perennials. Variability among species apparently reflects different dispersal “strategies” employed by the species; these different strategies may be correlated with other life-history traits and with ecological characteristics. Variability within species also may have ecological consequences in that such variability may represent an example of risk-spreading.     

Fine-scale spatial genetic structure in the frankincense tree <Emphasis Type="Italic">Boswellia papyrifera</Emphasis> (Del.) Hochst. and implications for conservation

The fine-scale genetic structure and how it varies between generations depends on the spatial scale of gene dispersal and other fundamental aspects of species’ biology, such as the mating system. Such knowledge is crucial for the design of genetic conservation strategies. This is particularly relevant for species that are increasingly fragmented such as Boswellia papyrifera. This species occurs in dry tropical forests from Ethiopia, Eritrea and Sudan and is an important source of frankincense, a highly valued aromatic resin obtained from the bark of the tree. This study assessed the genetic diversity and fine-scale spatial genetic structure (FSGS) of two cohorts (adults and seedlings) from two populations (Guba-Arenja and Kurmuk) in Western Ethiopia and inferred intra-population gene dispersal in the species, using microsatellite markers. The expected heterozygosity (H _E) was 0.664–0.724. The spatial analyses based on kinship coefficient (F _ij) revealed a significant positive genetic correlation up to a distance of 130 m. Spatial genetic structure was relatively weak (Sp = 0.002–0.014) indicating that gene dispersal is extensive within the populations. Based on the FSGS patterns found, we estimate indirectly gene dispersal distances of 103 and 124 m for the two populations studied. The high heterozygosity, the low fixation index and the low Sp values found in this study are consistent with outcrossing as the (predominant) mating system in B. papyrifera. We suggest that seed collection for ex situ conservation and reforestation programmes of B. papyrifera should use trees separated by distances of at least 100 m but preferably 150 m to limit genetic relatedness among seeds from different trees.     

Liana diversity and reproductive attributes in two tropical forests in Mexico

We describe liana diversity and variation in morphology of flowers and diaspores in two tropical forests in Mexico: a seasonally deciduous forest at Chamela, Jalisco and a lowland rain forest at Chajul, Chiapas. Flowers were classified as inconspicuous (1 cm in length and white or pale green flowers) or conspicuous (> 1 cm and brightly colored flowers). Dispersal syndrome was classified as anemochory, barochory, and zoochory. We recorded a higher number of lianas species in Chajul (128 spp.) than in Chamela (71 spp.). In both sites, families with the highest number of species were Bignoniaceae, Leguminosae, Sapindaceae, and Malphigiaceae, and almost half of the liana species had inconspicuous flowers. Most of the species had an abiotic syndrome of dispersal with 40 (56.3%) and 71 (55.5%) wind-dispersed species in Chamela and Chajul, respectively. Zoochory was significantly associated with inconspicuous flowers and anemochory with conspicuous flowers. Our results suggest that (1) flower morphology and dispersal type are not related with the amount of rainfall and (2) lianas are more prone to be wind-dispersed.     

Dispersal mode,seed shadows,and colonization patterns

This review assesses the state of our knowledge about comparative seed shadows. Using data presently available in the literature, I compare the slopes (on a log-linear scale) of seed shadows for plants with different morphologically characterized modes of dispersal. The seeds of many species have no evident morphological adaptation for dispersal and seem to achieve only short-distance dispersal. Seed shadows for herbaceous species with devices for wind have flatter slopes and more distant modes and maxima than those of ballists, which in turn exceed those with no special devices. Seed shadows for wind-dispersed trees and shrubs had similar or steeper slopes than those for vertebrate-dispersed species in this sample. Species with poor mechanisms for dispersal in space only sometimes had the capacity for better dispersal in time (dormancy). Although some species exhibited seed shadows sufficiently steep to be predicted to colonize new-habitat in a front or phalanx pattern, actual colonization patterns must reflect many other factors.     

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.