Are tortoises important seed dispersers in Amazonian forests?

According to most studies on seed dispersal in tropical forests, mammals and birds are considered the main dispersal agents and the role played by other animal groups remains poorly explored. We investigate qualitative and quantitative components of the role played by the tortoise Chelonoidis denticulata in seed dispersal in southeastern Amazon, and the influence of seasonal variation in tortoise movement patterns on resulting seed shadows. Seed shadows produced by this tortoise were estimated by combining information on seed passage times through their digestive tract, which varied from 3 to 17 days, with a robust dataset on movements obtained from 18 adult C. denticulata monitored with radio transmitters and spoon-and-line tracking devices. A total of 4,206 seeds were found in 94 collected feces, belonging to 50 seed morphotypes of, at least, 25 plant genera. Very low rates of damage to the external structure of the ingested seeds were observed. Additionally, results of germination trials suggested that passage of seeds through C. denticulata’s digestive tract does not seem to negatively affect seed germination. The estimated seed shadows are likely to contribute significantly to the dispersal of seeds away from parent plants. During the dry season seeds were dispersed, on average, 174.1 m away from the location of fruit ingestion; during the rainy season, this mean dispersal distance increased to 276.7 m. Our results suggest that C. denticulata plays an important role in seed dispersal in Amazonian forests and highlight the influence of seasonal changes in movements on the resulting seed shadows.     

Incorporating patterns of disperser behaviour into models of seed dispersal and its effects on estimated dispersal curves

The processes determining where seeds fall relative to their parent plant influence the spatial structure and dynamics of plant populations and communities. For animal dispersed species the factors influencing seed shadows are poorly understood. In this paper we test the hypothesis that the daily temporal distribution of disperser behaviours, for example, foraging and movement, influences dispersal outcomes, in particular the shape and scale of dispersal curves. To do this, we describe frugivory and the dispersal curves produced by the southern cassowary, Casuarius casuarius, the only large-bodied disperser in Australia’s rainforests. We found C. casuarius consumed fruits of 238 species and of all fleshy-fruit types. In feeding trials, seeds of 11 species were retained on average for 309 min (±256 SD). Sampling radio-telemetry data randomly, that is, assuming foraging occurs at random times during the day, gives an estimated average dispersal distance of 239 m (±207 SD) for seeds consumed by C. casuarius. Approximately 4% of seeds were dispersed further than 1,000 m. However, observation of wild birds indicated that foraging and movement occur more frequently early and late in the day. Seeds consumed early in the day were estimated to receive dispersal distances 1.4 times the ‘random’ average estimate, while afternoon consumed seeds received estimated mean dispersal distances of 0.46 times the ‘random’ estimate. Sampling movement data according to the daily distribution of C. casuarius foraging gives an estimated mean dispersal distance of 337 m (±194 SD). Most animals’ behaviour has a non-random temporal distribution. Consequently such effects should be common and need to be incorporated into seed shadow estimation. Our results point to dispersal curves being an emergent property of the plant–disperser interaction rather than being a property of a plant or species.     

Cheek-pouch dispersal of seeds by Japanese monkeys (Macaca fuscata yakui) on Yakushima Island, Japan

Seed dispersal by Japanese monkeys (Macaca fuscata yakui) via cheek-pouch was studied in a warm temperate evergreen forest on Yakushima Island. Plant list was compiled based on a study during 1986–1995, of which troops of monkeys have been habituated without artificial feeding. We followed the well-habituated monkeys in 1993 and 1994 to observe the feeding behavior and their treatments of fruits and seeds, and collected seeds dispersed by monkeys to record the distance carried from the mother trees. We checked the difference of germination ratio between seeds dispersed via cheek-pouch and seeds taken from mother trees by sowing experiments. Seeds and acorns of 22 species were observed to be dispersed via cheek-pouch of monkeys. Among them, three species with acorns were never dispersed via feces, and 15 species with drupes were seldom dispersed via feces. Plant species of which seeds are dispersed only via cheek-pouch had larger seeds than those of dispersed both via cheek-pouch and via feces, and typically had only one or two seeds in a fruit. As for one of cheek-pouch dispersal species,Persea thunbergii, the mean distance when seeds were carried from the mother trees via cheek-pouch was 19.7 m, and the maximum distance was as long as 105 m although more than 80% of seeds were dispersed within 30 m from mother trees. And 82% of seeds dispersed via cheek-pouch germinated. The easy separation of seeds from other parts of the fruit seems to facilitate cheek-pouch dispersal more than dispersal via feces. Cheek-pouch dispersal by monkeys has possibly enhanced the natural selection for larger seeds which bring forth larger seedlings with high shade-tolerance. In conclusion, cheek-pouch dispersal by monkeys is quite an important mode for trees in the mature stand in a warm temperate evergreen forest on Yakushima Island.     

Neglected seed dispersers: endozoochory by Javan lutungs (Trachypithecus auratus) in Indonesia

Leaf monkeys are known to be leaf eaters, and thus, their potential role as seed dispersers has been neglected. However, they do also feed on fruits. To examine the role of leaf monkeys as endozoochorous seed dispersers, we studied the Javan lutung (Trachypithecus auratus) in Indonesia. We compared multiple aspects of seed dispersal processes (amount and diversity of seeds ingested, dispersal distance, and germination rate) of lutungs with that of the sympatric long‐tailed macaque (Macaca fascicularis). Over the study period, 54 percent of the lutung feces contained intact seeds, which was equivalent to the macaque feces contained seeds (62%). Seeds of at least six plant species were detected in the lutung feces, which was less than those found in the macaque feces (>19 plant species). The main species of seeds defecated by both lutungs and macaques was Ficus spp. (seed size: 0.7 mm). Seed shadow, estimated from travel distance (range: 1–299 m) and gut passage rate (24–96 h), had a unimodal‐distribution with a peak at 51–100 m, and was shorter than that reported in published accounts of macaques and other similar and smaller sized frugivores. Finally, germination rates of Ficus spp. seeds ingested by both lutungs and macaques were lower than that of the control seeds. These results imply that the dispersal effectiveness of lutungs would be lower than that of the sympatric primate frugivores. However, at a population level, lutungs could play a significant role as seed dispersers for the small‐seeded species, and therefore, more research into their frugivorous habits is warranted.     

The role of Orii’s flying-fox (Pteropus dasymallus inopinatus) as a pollinator and a seed disperser on Okinawa-jima Island, the Ryukyu Archipelago, Japan

The role of the Orii’s flying-fox (Pteropus dasymallus inopinatus) as a pollinator and a seed disperser on Okinawa-jima Island was investigated by direct observations and radio-tracking from October 2001 until January 2006. We found that Orii’s flying-fox potentially pollinated seven native plant species. Its feeding behavior and plant morphological traits suggested that this species is an important pollinator of Schima wallichii liukiuensis and Mucuna macrocarpa. The flying-fox also dispersed the seeds of 20 native plant species. The seeds of all plants eaten by the flying-fox were usually dropped beneath the parent tree, although large fruits of four plant species were occasionally brought to the feeding roosts in the mouth, with the maximum dispersal distance—for Terminalia catappa—estimated to be 126 m. Small seeds of 11 species (mostly Ficus species) were dispersed around other trees, during the subsequent feeding session, through the digestive tracts, with the mean dispersal distance for ingested seeds estimated at 150 ± 230.3 m (±SD); the maximum dispersal distance was 1833 m. A comparison of the seed dispersal of available fruits according to the size of flying-foxes and other frugivores suggested that the seed dispersal of eight plant species producing large fruits mostly depended on Orii’s flying-fox. On Okinawa-jima Island, the Orii’s flying-fox plays an important role as a pollinator of two native plants and as a long-distance seed disperser of Ficus species, and it functions as a limited agent of seed dispersal for plants producing large fruits on Okinawa-jima Island.     

Importance of fragmentation-tolerant species as seed dispersers in disturbed landscapes

Forest fragmentation can negatively affect plants if animal seed-dispersers become locally extinct in fragments. We conducted a 2-year experiment to evaluate the importance of tree squirrels (Sciurus) as seed dispersers for Quercus, Carya, and Juglans, and to assess dispersal consequences in patches where fragmentation-sensitive eastern gray squirrels (Sciurus carolinensis) are absent. We accounted for fates of ∼15,700 seeds from five tree species in four exclosure treatments at 18 fragments during a high (2003–2004) and low seed (2004–2005) year. Two treatments excluded Sciurus to mimic disperser loss. We sampled nut-tree seedling density at 259 sites across eight watersheds, half of which were too fragmented to support S. carolinensis, but supported fragmentation-tolerant fox squirrels (Sciurus niger). Autumn-to-spring seed survival was low (∼1%) for all species during low seed production. During high seed production, survival was higher for Juglans nigra (20%) and Carya ovata (16%) than for three Quercus species (∼4% for Quercus palustris and Quercus rubra in two exclosure types; ∼1% for Quercus alba in all treatments). Survival of J. nigra, C. ovata, and Q. rubra was ≥2.1–7.7 times higher for seeds in exclosures that Sciurus could access. Seed displacement distance was higher in the low seed than the seed-rich year, but the proportion of seeds surviving to greater distances was higher in seed-rich years for all seed types except Q. rubra. This affirms the importance of masting to seed survival and dispersal, but also suggests an advantage to trees of producing seed in non-mast years. Seedling densities were comparable in watersheds with and without S. carolinensis. These results demonstrate the importance of tree squirrels as dispersers of nut-bearing trees, but suggest that fragmentation may not disrupt dispersal of certain species if losing S. carolinensis from disturbed landscapes is compensated for by fragmentation-tolerant fox squirrels (S. niger).     

Different contributions of birds and mammals to seed dispersal of a fleshy-fruited tree

Birds and mammals are important seed dispersers of fleshy-fruited plants. Although their behaviors are different, they frequently consume the same species. Thus to understand the dispersal of fleshy-fruited plants, the contribution of birds and mammals to seed dispersal has to be evaluated. Besides, within birds or mammals, some species may functionally different from others. In this study, we examined seed dispersal of the fleshy-fruited tree Swida controversa focusing on the difference between two frugivore groups (birds and mammals), and differences between species within groups. Collected seeds and S. controversa trees were identified using simple sequence repeat (SSR) genotyping, thus enabling to determine the distance between mother tree and dispersed seeds. The avian species were identified by DNA barcoding of feces, whereas the mammalian species were identified by the shape and smell of feces. Most seeds that fell near or under the maternal trees were dispersed by birds, resulting in short seed dispersal distances (average, 13 m). DNA barcoding detected five taxa of avian dispersers. No differences were detected in seed dispersal distance by different avian taxa (i.e., the distance between dispersed seeds and their maternal trees within the research plot); however the rate of seed immigration from outside the research plot by some avian taxa varied significantly. The seed dispersal distance by mammals was significantly further (127 m; min > 50 m) than that by birds. Additionally, immigrated seeds accounted for approximately two-thirds of mammal-dispersed seeds, indicating that these seeds were from outside the research plot, and that mammals significantly contributed to the long-distance seed dispersal of S. controversa. No differences in seed dispersal distance were detected between different mammalian taxa. Overall, this study revealed that birds and mammals show clearly different seed dispersal patterns, and thus, they play different roles in the regeneration of S. controversa.     

Novel polymorphic microsatellite loci and patterns of pollen-mediated gene flow in an ex situ population of <Emphasis Type="Italic">Eurycorymbus cavaleriei</Emphasis> (Sapindaceae) as revealed by categorical paternity analysis

Understanding the patterns of contemporary, pollen-mediated gene flow is of great importance for designing appropriate conservation strategies. In this study, ten novel polymorphic microsatellite loci were isolated for the rare dioecious tree, Eurycorymbus cavaleriei, and the patterns of pollen dispersal were investigated in an ex situ conserved population. A combination of microsatellite markers with high-collective exclusion power (0.932) was used to assign paternity to 240 seeds collected from eight maternal trees. The average effective pollen dispersal distance (δ) was 292.6 m and the frequency distribution of pollen movement suggested extensive pollen movement in the population. The effective pollen donors per maternal tree (N _ep) ranged from 5 to 10, and the most isolated maternal trees were observed with the largest number of N _ep = 10. Although a trend of near-neighbor mating was found in seven of eight maternal trees, no significant correlations were detected between the average effective pollen dispersal distance (δ) and the geographic distances (d1 and d2) between maternal and male trees. The increased average effective distance of pollen dispersal and number of N _ep for isolated maternal trees might be a compound consequence of low density and long-distance flight of pollinators of this species. The conservation implications of these results are discussed.     

Seed dispersal and realized gene flow of two forest orchids in a fragmented landscape

• Species with vast production of dust‐like windborne seeds, such as orchids, should not be limited by seed dispersal. This paradigm, however, does not fit recent studies showing that many sites suitable for orchids are unoccupied and most seeds land close to their maternal plant. To explore this issue, we studied seed dispersal and gene flow of two forest orchid species, Epipactis atrorubens and Cephalanthera rubra, growing in a fragmented landscape of forested limestone hills in southwest Bohemia, Czech Republic.
• We used a combination of seed trapping and plant genotyping methods (microsatellite DNA markers) to quantify short‐ and long‐distance dispersal, respectively. In addition, seed production of both species was estimated.
• We found that most seeds landed very close to maternal plants (95% of captured seeds were within 7.2 m) in both species, and dispersal distance was influenced by forest type in E. atrorubens. In addition, C. rubra showed clonal reproduction (20% of plants were of clonal origin) and very low fruiting success (only 1.6% of plants were fruiting) in comparison with E. atrorubens (25.7%). Gene flow was frequent up to 2 km in C. rubra and up to 125 km in E. atrorubens, and we detected a relatively high dispersal rate among regions in both species.
• Although both species occupy similar habitats and have similar seed dispersal abilities, C. rubra is notably rarer in the study area. Considerably low fruiting success in this species likely limits its gene flow to longer distances and designates it more sensitive to habitat loss and fragmentation.

     

Seed dispersal of the non-native invasive tree <Emphasis Type="Italic">Ailanthus altissima</Emphasis> into contrasting environments

Ailanthus altissima has a long history of invasion in urban areas and is currently spreading into suburban and rural areas in the eastern U.S. The objectives of our study were to (1) determine whether A. altissima seed dispersal distance differed between populations on the edges of open fields and intact deciduous forest, and (2) determine whether dispersal differed for north and south winds. We also assessed the relationship between seed characteristics and distance from source populations in fields and forests, and whether seeds disperse at different rates throughout the dispersal season. Using two fields, two intact forest stands, and one partially harvested stand, we sampled the seed rain at 10 m intervals 100 m into each site from October to April 2002–2003. We compared seed density in field and intact forests using a three-way ANOVA with distance from source, wind direction, and environmental structure as independent variables. To assess the accuracy of common empirical dispersal models, mean seed density data at each site were fitted with alternative regression models. We found that mean seed dispersal distance depended on environmental structure and wind direction, a result driven in large part by dispersal at a single site where seed density did not decline with distance. The two alternative regression models fit each site’s dispersal curve equally well. More seeds were dispersed early than in mid- or late-season. Large, heavy seeds traveled as far as small light seeds. Turbulent winds appear to be necessary for seed release, as indicated by a wind tunnel experiment. A. altissima is able to disperse long distances into fields and into mature forests, and can reach canopy gaps and other suitable habitats at least 100 m from the forest edge. It is an effective disperser and can spread rapidly in fragmented landscapes where edges and other high light environments occur. These conditions are increasingly common throughout the eastern U.S. and in other temperate regions worldwide.     

Spatial genetic structure and restricted gene flow in a functionally dioecious fig, <Emphasis Type="Italic">Ficus pumila</Emphasis> L. var. <Emphasis Type="Italic">pumila</Emphasis> (Moraceae)

The mutualism between fig plants and fig wasps has been recognized as one of the most specialized systems of symbiosis. Figs are pollinated by their highly specific pollinating fig wasps, and the pollinating fig wasps are raised within the syconia of figs. Recent studies indicated a difference between monoecious and dioecious figs in the dispersal range of pollinating wasps, which has potential consequences for gene flow. In this study, we detected the gene-flow pattern of the dioecious climbing fig, Ficus pumila L. var. pumila, at both local and regional scales. At the local scale, spatial autocorrelation analysis indicated strong genetic structure at short distances, a pattern of limited gene flow. This result was also supported by a high inbreeding coefficient (F _IS = 0.287) and significant substructuring (F _ST = 0.060; P < 0.001). Further analysis indicated that the effective gene dispersal range was 1,211 m, and the relative contribution of seed dispersal was smaller than that of pollen dispersal. The inferred effective range of pollen dispersal ranged from 989 to 1,712 m, while the effective seed dispersal range was less than 989 m. Lack of long-distance dispersal agents may explain the limited seed dispersal. The high density of receptive fig trees was the most likely explanation for limited pollen dispersal, and the position of syconia and relatively low wind speed beneath the canopy may contribute to this phenomenon. At the regional scale, significant negative correlations (kinship coefficient F _ij ranging from −0.038 to −0.071) existed in all comparisons between the studied population and other populations, and the assignment test grouped almost all individuals of the studied population into a distinct cluster. Asynchronous flowering on the regional scale, which provides a barrier for the pollinating wasps to fly from the studied population to the other populations, is probably responsible for the limited gene flow on the regional scale.     

云南拉沙山黑白仰鼻猴对种子传播潜力的评估

灵长类是森林生态系统中植物种子的主要传播者,有助于森林植被的更新,然而受研究方法的限制,灵长类种子传播潜力常被低估。为全面评估温带灵长类动物的种子传播潜力,采用直接观察法和粪便分析法评估珍稀濒危灵长类动物黑白仰鼻猴的种子传播潜力。于2018年11月—2019年10月采用直接观察法(瞬时扫描取样法)收集云岭省级自然保护区拉沙山黑白仰鼻猴的活动时间分配数据,获取每月取食果实的比例;同时每月收集黑白仰鼻猴的粪便,采用粪便分析法分拣猴粪中残留的植物种子,统计有完整种子残留的月份和粪便比例,应用这两种方法评估黑白仰鼻猴种子传播潜力及其差异。结果表明：直接观察法收集到黑白仰鼻猴取食果实的月份数为6个月(7—12月),月均取食果实的比例为(15.31±20.15)%,共取食13种果实;而粪便分析法发现黑白仰鼻猴粪粒内全年都有完整种子残留,粪便中月均完整种子残留比例(35.19±35.43)%,其中9月至第二年1月粪便中种子残留比例都大于50%,共取食18种果实;综合两种方法发现云南拉沙山黑白仰鼻猴共取食20种植物果实,具有较高的种子传播潜力。直接观察法可确定黑白仰鼻猴取食果实的物种数,而粪便分析法能...     

Survivorship and growth of seedlings of four dipterocarp species in a tropical rainforest of Sarawak,East Malaysia

Survivorship and growth of seedlings of four dipterocarp species (Dipterocarpus actangulus, D. globosus, Dryobalanops aromatica, Dryobalanops lanceolata) were studied for 2.5 years in a mixed dipterocarp forest in Sarawak, East Malaysia. Predispersal seed predation rates were larger forD. globosus (75%) thanD. lanceolata (27–34%) andD. aromatica (18–26%). Less than 20% of the twoDryobalanops seeds were damaged by vertebrates after seed dispersal. During the period from seed dispersal to the time when the seedlings had shed cotyledons, more dispersed seeds died in the twoDipterocarpus (ca 90%) than the twoDryobalanops (ca 60–70%). The major mortality factors during this period were uprooted and seed/seedling predation by insects or vertebrates. After the seedlings shed cotyledons, all species showed constant mortality rates of 34, 15–16, 17 and 6%/year forD. actangulus, D. lanceolata, D. aromatica andD. globosus, respectively, in the forest understorey. Mortality was lower in less shaded conditions than in more shaded ones forD. aromatica andD. actangulus, but not significantly different forD. lanceolata andD. globosus. A majority of dead seedlings were killed by fallen branches or were found standing with wilted leaves, probably due to water stress. No significant correlation was found between seed/seedling mortality and distance from mother trees or the initial density of seeds/seedings for all species. The mean leaf production was positively correlated with the estimated diffuse light factor of their habitats for each species.     

Seed weight increases with altitude in the Swiss Alps between related species but not among populations of individual species

Seed weight is a crucial plant life history trait, determining establishment success and dispersal ability. Especially in stressful environments, larger seeds may be selected at the expense of seed number, because larger seeds have a better chance of giving rise to an established offspring. We tested the hypotheses that between related species-pairs and among populations of single species a similar trend for increasing seed weight with increasing altitude should be present. Firstly, we measured seed weights from 29 species-pairs, with one species occurring in lowland areas and a congeneric species from high altitudes. Seeds of the alpine species were 28±8% larger than seeds from lowland species (P<0.01). Compared to the related lowland species, 55% of the alpine species had heavier seeds, 3% (one species) had lighter, and 41% had seeds of approximately equal weight. Secondly, we compared seed weights among populations of four species from different habitats and with different life histories. Seeds from between 11 and 34 populations per species were sampled along altitudinal gradients of 800–1,500 m (ca. 800 m in Scabiosa lucida, ca. 1,000 m in Saxifraga oppositifolia, ca. 1,000 m in Epilobium fleischeri, and ca. 1,500 m in Carex flacca). In all the four species, we found no indication for heavier seeds at higher altitudes. Our results indicate a selection pressure for species with heavier seeds at higher altitude, but the trend does not seem to operate across all cases. Phylogenetic constraints may limit the correlation among altitude and seed weight, operating particularly against selection for larger seed size, the closer populations and species are related to each other.     

Fruits foraging patterns and seed dispersal effect of frugivorous birds on Hippophae rhamnoides sinensis

Behaviors of 18 species of birds eating fruits of Hippophae rhamnoides spp. sinensis were observed from September 2003 to March 2004. Their foraging patterns were found to be very different and can be divided into five classes: (1) direct swallowing the fruits on crown of the shrubs and sometimes regurgitating seeds soon after; (2) carrying the fruits to their perching sites and swallowing; (3) pecking the fruits from the shrubs to the ground, eating pulp and seeds but leaving pericarp; (4) pecking through the pericarp, eating pulp and leaving pericarp and seeds; (5) pecking through the pericarp on the top of fruits, and only eating seeds. These foraging patterns have different effects on seed dispersal of H. rhamnoides spp. sinensis. The germination experiment of three groups of seeds (seeds from feces, dry fruits and extracted seeds from dry fruits) was carried out. Although ingestion processes of birds had some adverse effects on the seed germination of H. rhamnoides spp. sinensis, the seeds from feces still have a relatively higher germination ratio. H. rhamnoides spp. sinensis provides food to a variety of frugivorous birds, and the birds disperse its seeds. Thus, a mutually beneficial relationship between the bird and the seed is formed. __________ Translated from Chinese Journal of Ecology, 2005, 24(6): 635–638 [译自: 生态学杂志, 2005, 24(6): 635–638]     

Effect of Resting Patterns of Tamarins (<Emphasis Type="BoldItalic">Saguinus fuscicollis</Emphasis> and <Emphasis Type="BoldItalic">Saguinus mystax</Emphasis>) on the Spatial Distribution of Seeds and Seedling Recruitment

The spatial distributions of dispersed seeds have important evolutionary consequences for plants. Repeated defecations in sites frequently used by seed dispersers can result in high seed concentrations. We observed the resting behavior of a mixed-species group of tamarins in Peru and recorded the occurrence of seed dispersal (over 8 mo) and seed fate (over 11–22 mo) to determine whether the location and use of resting sites influenced the spatial distribution of dispersed seeds and seedlings. The tamarins rested mostly on trees (Saguinus fuscicollis: 60.6%, S. mystax: 89.2%) and dead trunks (S. fuscicollis: 24.4%) and used 61% of their resting sites repeatedly. During both the dry and wet seasons, tamarins dispersed significantly more seeds within resting areas (0.00662 and 0.00424 seeds/m², respectively) than outside them (0.00141 and 0.00181 seeds/m²). Seed survival and seedling recruitment did not differ significantly between resting and other areas, resulting in a higher seedling concentration around the resting sites. Seed density did not increase with the duration or the frequency of use of the resting sites but did increase when we pooled the seasonal resting sites together in 50 m × 50 m quadrats, ultimately causing a clumped distribution of dispersed seeds. The use of resting sites in secondary forest, particularly during the dry season, allows the creation of seedling recruitment centers for species coming from the primary forest. Our findings show that tamarin resting behavior affects the spatial distribution of dispersed seeds and seedlings, and their resting sites play an important role in plant diversity maintenance and facilitate forest regeneration in degraded areas.     

Spatial Segregation of the Sexes and Nutrients affect Reproductive Success in a Dioecious Wind-pollinated Grass

In many dioecious plant species in which spatial distributions of males and females have been examined, the sexes are spatially segregated – usually along an environmental gradient. Unless pollen is uniformly distributed in a population, spatial segregation of the sexes should reduce the average mating success of individuals. In three Californian populations of Distichlis spicata – a wind-pollinated grass species that exhibits spatial segregation of the sexes – I examined patterns of pollen movement and the effects of pollen load and nutrient availability on seed set to determine whether spatial segregation of the sexes actually reduces mating success for both males and females. In two of the populations, pollen dispersal was restricted, and pollen augmentation consistently, significantly increased seed set. However, in the third population – which had the lowest seed set – I found that although there were some indications of pollen limitation, pollen dispersal was not restricted, and seed production was limited primarily by nutrient availability. These results imply that in some populations of D. spicata nutrient limitation on the production of seeds by females may be sufficiently strong that spatial segregation of the sexes causes a fairly low cost to reproductive success compared with a more random distribution of the sexes. However, in other populations, pollen does limit mating success, and the spatial segregation of males and females in these populations is reducing the fecundity of both males and females.     

A keystone ant species promotes seed dispersal in a “diffuse” mutualism

In order to understand the dynamics of co-evolution it is important to consider spatial variation in interaction dynamics. We examined the relative importance of ant activity, diversity and species identity in an ant seed dispersal mutualism at local, regional and continental scales. We also studied the determinants of seed dispersal rates and dispersal distances at eight sites in the Eneabba sandplain (29.63 S, 115.22 E), western Australia to understand local variation in seed dispersal rate and distance. To test the generality of the conclusions derived from the eight local sites, we established 16 sites along a 1650-km transect in western Australia, covering 11° of latitude and a six-fold increase in rainfall, at which we sampled the ant assemblage, estimated ant species richness and ant activity and observed the removal rate of myrmecochorous seeds. We also assessed the importance of ant species identity at a continental scale via a review of studies carried out throughout Australia which examined ant seed dispersal. Among the eight sandplain shrubland sites, ant species identity, in particular the presence of one genus, Rhytidoponera, was associated with the most dispersal and above average dispersal distances. At the landscape scale, Rhytidoponera presence was the most important determinant of seed removal rate, while seed removal rate was negatively correlated with ant species richness and latitude. Most ant seed removal studies carried out throughout Australia reinforce our observations that Rhytidoponera species were particularly important seed dispersers. It is suggested that superficially diffuse mutualisms may depend greatly on the identity of particular partners. Even at large biogeographic scales, temporal and spatial variation in what are considered to be diffuse mutualisms may often be linked to variation in the abundance of particular partners, and be only weakly – or negatively – associated with the diversity of partners.     

Spatial genetic structure in populations of the terrestrial orchid Orchis cyclochila (Orchidaceae)

Orchid seeds are minute, dust-like, wind-borne and, thus, would seem to have the potential for long-distance dispersal. Based on this perception, one may predict near-random spatial genetic structure within orchid populations. In reality we do not know much about seed dispersal in orchids and the few empirical studies of fine-scale genetic structure have revealed significant genetic structure at short distances (< 5m), suggesting that most seeds of orchids fall close to the maternal plant. To obtain more empirical data on dispersal, Ripley’s L(d)-statistics, spatial autocorrelation analyses (coancestry, f_ij analyses) and Wright’s F statistics were used to examine the distribution of individuals and the genetic structure within two populations of the terrestrial orchid Orchis cyclochila in southern Korea. High levels of genetic diversity (H_e = 0.210) and low between-population variation were found (F_ST = 0.030). Ripley’s L(d)-statistics indicated significant aggregation of individuals, and patterns varied depending on populations. Spatial autocorrelation analysis revealed significant positive genetic correlations among individuals located <1 m, with mean f_ij values expected for half sibs. This genetic structure suggests that many seeds fall in the immediate vicinity of the maternal plant. The finding of significant fine-scale genetic structure, however, does not have to preclude the potential for the long distance dispersal of seeds. Both the existence of fine-scale genetic structure and low F_ST are consistent with a leptokurtic distribution of seed dispersal distances with a very flat tail.