Within- and between-species patterns of allocation to pulp and seed in vertebrate dispersed plants

Non-uniform scaling of pulp and seed mass has been shown to alter dispersal probabilities in vertebrate dispersed species. Since dispersal in tropical forests is strongly linked to establishment success, processes determining allocation to pulp and seed are likely to impact on parental fitness, and therefore should be under the control of natural selection. In this study I examine size-dependent dry mass allocation pattern to pulp and seed mass both among and within 20 fruit-producing plant species of tropical rainforest in northeast Queensland, Australia. Reduced major axis analyses using mean values for each species showed significant isometry, indicating that at the community level, plant species that employ vertebrates as a means of seed dispersal tend to allocate an equivalent mean relative proportion of the overall dry weight investment in fruit to pulp. However, identical analyses conducted for each species separately revealed that relationships within individual species do not reflect the inter-specific relationship. These results imply two influences on dry mass allocation to fruit components; the first (within-species allometry) determines how fruits vary within each species. The second (between-species allometry) operates in a similar manner across species to produce equal ratios of mean pulp and seed mass independent of within-species allometries.     

Effect of phylogeny, life history and habitat correlates on seed germination of 69 arid and semi-arid zone species from northwest China

We tested the effects of phylogeny, life history and habitat conditions for seedling establishment on seed germination of 69 arid/semi-arid zone species from northwest China. Final germination percentages had a bimodal distribution, whereas, days to first germination were skewed toward short periods. Stepwise multi-way ANOVAs showed that the amount of variance in final germination percentage among species for seeds incubated in light was explained by phylogeny (13.6%), dispersal mode (11.9%), seed mass (3.6%), and habitat (0.2%); in darkness by dispersal mode (17.5%), phylogeny (12.7%), seed mass (5.7%), and habitat (0%). A series of ANOVAs revealed that the variance in final germination percentages among species is largely dependent upon phylogeny and dispersal mode but that it is also influenced by seed mass and habitat. The effects of dispersal mode and seed mass on final germination percentage among species were phylogenetic group- and habitat-specific. Wind-dispersed seeds had higher germination percentages than unassisted and vertebrate-dispersed seeds. Compared to xerophytes, mesophytes tended to have smaller seed mass, higher mean germination percentages and a greater effect of seed mass on final germination fractions. Our results suggest that phylogeny and life history should be taken into account in studies on the role of natural selection in regulation of seed germination.     

Seed dispersal by wind,birds, and bats between Philippine montane rainforest and successional vegetation

In the moist Neotropics, vertebrate frugivores have a much greater role in the dispersal of forest and successional woody plants than wind, and bats rather than birds play the dominant role in dispersing early successional species. I investigated whether these patterns also occurred in a Philippine montane rainforest and adjacent successional vegetation. I also asked whether seed mass was related to probability of dispersal between habitats. A greater number of woody species and stems in the forest produced vertebrate-dispersed seeds than wind-dispersed seeds. Although input of forest seeds into the successional area was dominated by vertebrate-dispersed seeds in terms of species richness, wind-dispersed seeds landed in densities 15 times higher. Frugivorous birds dispersed more forest seeds and species into the successional area than bats, and more successional seeds and species into the forest. As expected, seed input declined with distance from source habitat. Low input of forest seeds into the successional area at the farthest distance sampled, 40 m from forest edge, particularly for vertebrate-dispersed seeds, suggests very limited dispersal out of forest even into a habitat in which woody successional vegetation provides perches and fruit resources. For species of vertebrate-dispersed successional seeds, probability of dispersal into forest declined significantly with seed mass.     

Latitude, seed predation and seed mass

Aim We set out to test the hypothesis that rates of pre‐ and post‐dispersal seed predation would be higher towards the tropics, across a broad range of species from around the world. We also aimed to quantify the slope and predictive power of the relationship between seed mass and latitude both within and across species. Methods Seed mass, pre‐dispersal seed predation and post‐dispersal seed removal data were compiled from the literature. Wherever possible, these data were combined with information regarding the latitude at which the data were collected. Analyses were performed using both cross‐species and phylogenetic regressions. Results Contrary to expectations, we found no significant relationship between seed predation and latitude (log₁₀ proportion of seeds surviving predispersal seed predation vs. latitude, P = 0.63; R² = 0.02; n = 122 species: log₁₀ proportion of seeds remaining after postdispersal seed removal vs. latitude, P = 0.54; R² = 0.02; n = 205 species). These relationships remained non‐significant after variation because of seed mass was accounted for. We also found a very substantial (R² = 0.21) relationship between seed mass and latitude across 2706 species, with seed mass being significantly higher towards the tropics. Within‐species seed mass decline with latitude was significant, but only about two‐sevenths, as rapid as the cross‐species decline with latitude. Results of phylogenetic analyses were very similar to cross‐species analyses. We also demonstrated a positive relationship between seed mass and development time across ten species from dry sclerophyll woodland in Sydney (P < 0.001; R² = 0.77; Standardized Major Axis slope = 0.14). These data lend support to the hypothesis that growing period might affect the maximum attainable seed mass in a given environment. Main conclusions There was no evidence that seed predation is higher towards the tropics. The strong relationship between seed mass and latitude shown here had been observed in previous studies, but had not previously been quantified at a global scale. There was a tenfold reduction in mean seed mass for every c. 23° moved towards the poles, despite a wide range of seed mass within each latitude.     

Variation in seed germination of 86 subalpine forest species from the eastern Tibetan Plateau: phylogeny and life-history correlates

We have examined the seed germination strategy of 86 subalpine woody species from the eastern part of the Tibetan Plateau and assessed correlations between seed germination and phylogeny, life-history attributes, habitat, and altitude using stepwise analyses of variance (ANOVAs) and phylogenetically independent contrasts (PICs). A series of ANOVAs revealed that the total amount of variance in the arcsine-transformed value of germination percentage among species could be accounted for by phylogeny (19.2%) and dispersal mode (14.3%) in seeds incubated in the light and by phylogeny (29.3%) and dispersal mode (11.0%) in those incubated in the dark. In addition, the total amount of variance in T50 (days to half of the final germination) can be accounted for by phylogeny (22.7%) and dispersal mode (17%) in seeds incubated in the light and by phylogeny (32.1%) and dispersal mode (9.9%) in those incubated in the dark. PICs indicated a significant correlation between germination percentage and dispersal mode and a positive correlation between T50 and seed mass, showing that wind-dispersed seeds had higher germination percentages than both unassisted and vertebrate-dispersed seeds and that small seeds germinated faster than large ones. We suggest that phylogeny and dispersal mode may constrain variation in seed germination across species and that the speed of germination and seed mass may co-vary to adapt to a variable environment. Therefore, from an evolutionary perspective, seed germination might be the result of both selective pressures over long-term ecological time and phylogenetic constraints over the long-standing evolutionary history of the phylogenetic group.     

Global patterns in seed size

Aim  To provide the first global quantification of the slope and shape of the latitudinal gradient in seed mass, and to determine whether global patterns in seed mass are best explained by growth form, vegetation type, seed dispersal syndrome, or net primary productivity (NPP).
Location  Global.
Methods  We collected seed mass data for 11,481 species × site combinations from around the world. We used regression to describe the latitudinal gradient in seed mass, then applied general linear models to quantify the relative explanatory power of each of the variables hypothesized to underlie the latitudinal gradient in seed size.
Results  There is a 320-fold decline in geometric mean seed mass between the equator and 60°. This decline is not linear. At the edge of the tropics, there is a sudden 7-fold drop in mean seed mass. The strongest correlates of the latitudinal gradient in seed mass are plant growth form, and vegetation type, followed by dispersal syndrome and NPP. A model including growth form, vegetation type, dispersal syndrome and NPP explains 51% of the variation in seed mass. Latitude explains just 0.2% of the residual variation from this model.
Main conclusions  This is the first demonstration of a major decrease in seed size at the edge of the tropics. This drop in seed mass is most closely correlated with changes in plant growth form and vegetation type. This suggests that the drop in seed mass might be part of a sudden change in plant strategy at the edge of the tropics.     

The effects of kernel mass and nutrition reward on seed dispersal of three tree species by small rodents

In the present study, colored tags were fastened to individual seeds to investigate seed dispersal and seed fates of Pinus koraiensis, Corylus mandshurica, and Corylus heterophylla with different seed traits. Our aim was to determine the role of the proportion of kernel mass, caloric reward per seed, and tag color in affecting seed removal rates and seed fates. We predicted that higher proportion of kernel mass and caloric reward will favor seed removal and caching, while lower ones will facilitate seed consumption either in situ or after removal. Our results showed that the proportion of kernel mass, rather than seed size, played an important role in determining seed removal rates and seed fates. Seeds of C. heterophylla with the lowest proportion of kernel mass, regardless of their largest size and/or mass, had lower removal rates, lower level of caching but higher proportion of consumption in situ compared with P. koraiensis and C. mandshurica seeds. Seeds with higher caloric reward exhibited greater dispersal distances. Seed kernel proportion was closely correlated to seed removal rates and caching rates, while seed size and hull thickness seemed to play less important role in affecting seed dispersal, probably because the higher seed handling ability of small rodents in the experimental areas. Our investigation revealed no significant effect of tag color on seed removal rates and seed fates of the three seed species.     

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.     

Allometric allocation in fruit and seed packaging conditions the conflict among selective pressures on seed size

Selective pressures on seed size could vary among the different stages of plant life cycles, so no simple relation could explain a priori its evolution. Here, we determined the relationships between seed size and two fitness components—seed dispersal and survival from predation—in a bird-dispersed tree, Crataegus monogyna. We interpret these relationships in relation to the patterns of mass allocation to fruit and seed components. Selection patterns were assessed at two levels (1) selection pressures on the parent tree; comparing seed dispersal efficiency among individual plants and (2) selection pressures at the individual seed level; comparing seed size variation (i) before and after dispersal, and (ii) before and after postdispersal seed predation. Dispersal efficiency (percentage of seed crop dispersed) was positively correlated with fruit mass and fruit width. Differences in crop size did not offset this effect, and larger seeds were overrepresented in the seed rain relative to the seed pool before dispersal. However, the advantage of larger seeds during the dispersal stage was cancelled later by an opposite selection pressure exerted by seed predators. As a result, smaller seeds had a higher probability of surviving postdispersal seed predation, establishing an evolutionary conflict imposed by the need for dispersal and the danger of being predated. Birds and rodents preferentially selected highly profitable fruits and seeds in terms of the relative proportion of their components. Larger fruits had a higher pulp to seed proportion than smaller ones, and all seeds had the same proportion of coat relative to the embryo-plus-endosperm fraction. Hence, although predator pressures were stronger than disperser ones, larger seeds invested proportionally less in structural defense than in dispersal.     

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.     

脊椎动物传播植物肉质果中的次生物质及其生态作用

在种子植物-动物的互惠关系中,植物果实成熟后需要吸引种子传播者取食果实,传播其种子至适宜萌发的生境,同时又要防御种子捕食者过度消耗种子。果实内的次生物质(如:配糖生物碱、大黄素、辣椒素)在此过程中起到重要的调控作用。依赖脊椎动物传播的肉质果中往往含有与植物茎、叶内相同的次生物质,其种类繁多,主要分为含氮化合物、酚类化合物和萜类化合物。未成熟果实内富含次生物质(如:单宁、大黄素),主要保护未成熟种子不被潜在的捕食者和食果动物取食,这些次生物质的含量通常随果实成熟而降低;其它次生物质(如:脱辅基类胡萝卜素)的含量随果实成熟而增多,可能起到吸引食果动物的作用。在对脊椎动物捕食的抵御中,果实内不同类型的次生物质促使成熟果实对所有脊椎动物都有毒性(专毒性)或者仅对种子捕食者有毒性(泛毒性)。肉质果内的次生物质对植物-食果动物相互关系的调控作用,还可以通过调节动物取食频次和数量、抑制和促进种子萌发、改变种子在肠道的滞留时间、吸引传播者等生态作用而实现。某种次生物质往往集多种生态作用于一身。目前对肉质果内次生物质与脊椎动物相互关系的探讨还不够深入。未来研究需要综合考虑植物次生物质与果实生理生化、形态学等特征对食果者的综合调控机理;次生物质在种子传播后的调控作用对植物种群或群落结构和分布格局的影响;从动植物协同进化角度探讨植物次生物质的产生、防御和吸引策略与脊椎动物对果实的选择和消费之间的关系等。开展脊椎动物传播肉质果实中次生物质的研究,对完善种子传播机制、植物繁殖和更新格局,丰富动植物相互作用、协同进化理论具有重要的意义。     

Large-scale geographical trends in fruit traits of vertebrate-dispersed temperate plants

Aim  To assess large-scale geographical trends in the character of fleshy, vertebrate-dispersed fruits.
Location  Europe between central Sweden and southern Spain.
Methods  Analyses of fruit of sixty-three plant species from twenty-nine families were compiled from four regional data sets. Four structural and five chemical fruit traits were analysed intraspecifically to control rigorously for phylogenetic lineage effects. Trends were examined in relation to various biological features of the considered species.
Results  Contents of soluble carbohydrate and lipids decreased markedly northwards. Fruit diameter and fresh mass peaked at the wettest site, while the pulp water content remained more constant throughout the gradient than any other fruit trait. Ash content, seed number and seed mass did not change, while the nitrogen content showed conflicting trends. No relation was detected between observed variation in fruit traits and fruit type, fruit colour, ripening season, plant growth form, leaf longevity, or geographical distribution of the considered plant species.
Main conclusions  Considerable intraspecific variability exists in vertebrate-dispersed fruits on large geographical scales. Climate presumably affects particularly those traits related to carbon and water gain and storage. Most research on fruit–frugivore interactions has been carried out on small spatial scales and failed to find matchings between frugivore communities and the character of fleshy fruits. I suggest that explicitly addressed large-scale surveys on the geographical variability of fruits and their disperser assemblages are needed to elucidate their spatial patterns and to determine the extent to which fleshy fruit traits are shaped by animals and/or abiotic factors.     

Ecological correlates of seed mass variation in Phoradendron juniperinum,a xylem-tapping mistletoe

Summary We investigated several ecological correlates of seed mass variation in the hemiparasitic, xylemtapping mistletoe, Phoradendron juniperinum. Mean seed mass varied two-fold among plants between the ages of 4 and 14 years old and was positively correlated with parental plant age. Both the standard deviation and the coefficient of variation in mean seed mass decreased with increasing plant age demonstrating that, on average, younger plants produced seed with more variable mass. Nitrogen concentrations (mg nitrogen per gram of seed) of both the seed and fruit (pericarp) were not correlated with mass or the age of the parent plant from which the seed was taken. However, the nitrogen content per seed (mg nitrogen per seed) was positively correlated with the mean seed dry mass and the age of the seed parent, suggesting that the carbon to nitrogen ratio of individual seeds remained relatively constant as seed mass increased and plants grew older. Seed germination ranged between 20% and 86% and was positively correlated with mass and parental plant age. Heavier seeds (seeds from older plants) also had the highest root radicle growth rates. Furthermore, the final root radicle length after 76 d of growth was positively correlated with seed dry mass. When grown on a medium containing an extract prepared from the host plant foliage, all seeds showed lower germination, grew more slowly and had shorter overall root radicles, but had significantly greater development of the haustorial disks (the holdfast which forms the host-parasite junction in Phoradendron) than seeds grown on a control medium. Our results suggest that, on average, seeds of greater mass produced by older plants have a greater total resource pool per propagule (fruit + seed). This resource pool may be important in conferring a greater potential for dispersal (fruit), survival, colonization, and establishment.     

Correlated evolution of genome size and seed mass

Previous investigators have identified strong positive relationships between genome size and seed mass within species, and across species from the same genus and family. Here, we make the first broad-scale quantification of this relationship, using data for 1222 species, from 139 families and 48 orders. We analyzed the relationship between genome size and seed mass using a statistical framework that included four different tests. A quadratic relationship between genome size and seed mass appeared to be driven by the large genome/seed mass gymnosperms and the many small genome size/large seed mass angiosperms. Very small seeds were never associated with very large genomes, possibly indicating a developmental constraint. Independent contrast results showed that divergences in genome size were positively correlated with divergences in seed mass. Divergences in seed mass have been more closely correlated with divergences in genome size than with divergences in other morphological and ecological variables. Plant growth form is the only variable examined thus far that explains a greater proportion of variation in seed mass than does genome size.     

Long-term variation in seed mass and seed production in populations of Paris quadrifolia

Seed production of the perennial herb Paris quadrifolia L. (Liliaceae) was investigated in five populations in northern Poland. The long-term seed production per square metre differed significantly among populations and years. Moreover, throughout the 7-year study period, 30% of both whole ripe fruits and seeds alone were predated. Variation in seed mass per fruit in both space and time was significant. Throughout the 7-year study, nearly all the marked individuals produced fruits once every 2 years. The most frequent break between fruiting was 2 years and the longest was 5 years. Only the mean seed mass in fruits of the same individual varied significantly over subsequent years. In the five populations, the number of ovules, number of seeds in the fruit and seed mass varied significantly between populations. However seed/ovule ratio did not differ in fruits in the five populations. The seed mass/number trade-off in fruits was strongly partially correlated when the effect of total seed mass was considered. Breeding experiments suggest that P. quadrifolia has a substantial capacity for both inbreeding and outbreeding. There were no significant differences in the seed/ovule ratio, seed number or seed mass in fruits produced from bagged or control flowers. However, both seed/ovule ratio and number of seeds were significantly lower in fruits from emasculated flowers.     

Reserve mass and dispersal investment in relation to geographic range of plant species: phylogenetically independent contrasts

Recent studies have reported conflicting evidence about correlations between seed size and plant species geographic range sizes. Using phylogenetically independent contrasts (PICs) within genera, we found no consistent differences in reserve mass between species with similar dispersal morphology and «wide>> versus «narrow>> geographic ranges. There was also no tendency within genera for broad ranged species to be those that allocate a larger percentage of the resources invested in each diaspora to dispersal structures. PICs were also constructed between species having a tenfold difference in seed size. In these PICs, the larger seeded species often occupied a greater number of regions than species with smaller seed sizes. This result was generated primarily through the comparison of species from different genera, families or higher level taxa which differed not only in seed mass but also in dispersal modes and growth forms. Finally, comparing species within Acacia and Eucalyptus having similar seed size but different dispersal modes, we found that bird dispersal (in Acacia ) and possession of a wing for wind dispersal (in Eucalyptus ) was associated with wider geographic range compared to lower-investment dispersal modes. Taken together, these comparisons indicate that seed size is not itself important as a factor influencing breadth of geographic range. Dispersal mode and growth form may have an influence, however, and seed size differences may be associated with contrasts in dispersal mode or growth form.     

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.     

Post-dispersal seed predation and the establishment of vertebrate dispersed plants in Mediterranean scrublands

The post-dispersal fate of seeds and fruit (diaspores) of three vertebrate-dispersed trees, Crataegus monogyna, Prunus mahaleb and Taxus baccata, was studied in the Andalusian highlands, south-eastern Spain. Exclosures were used to quantify separately the impact of vertebrates and invertebrates on seed removal in relation to diaspore density and microhabitat. The three plant species showed marked differences in the percentage of diaspores removed, ranging from only 5% for C. monogyna to 87% for T. baccata. Although chaffinches (Fringilla coelebs) fed on diaspores, rodents (Apodemus sylvaticus) were the main vertebrate removers of seed and fruit. Two species of ant (Cataglyphis velox and Aphaenogaster iberica) were the only invertebrates observed to remove diaspores. However, the impact of ants was strongly seasonal and they only removed P. mahaleb fruit to any significant extent. While removal of seed by rodents was equivalent to predation, ants were responsible for secondary dispersal. However, their role was limited to infrequent, small-scale redistribution of fruit in the vicinity of parent trees. Rodents and ants differed in their use of different microhabitats. Rodents foraged mostly beneath trees and low shrubs and avoided open areas while the reverse was true of ants. Thus, patterns of post-dispersal seed removal will be contigent on the relative abundance and distribution of ants and rodents. Studies which neglect to quantify separately the impacts of these two guilds of seed removers may fail to elucidate the mechanisms underlying patterns of post-dispersal seed removal. The coincidence of both increased seed deposition by the main avian dispersers (Turdus spp.) and increased seed predation with increasing vegetation height suggested that selection pressures other than post-dispersal seed predation shape the spatial pattern of seed dispersal. Rather than providing a means of escaping post-dispersal seed predators, dispersal appears to direct seeds to microhabitats most suitable for seedling survival. Nevertheless, the reliance of most vertebrate-dispersed trees on regeneration by seed and the absence of persistent soil seed banks imply that post-dispersal seed predators may exert a strong influence on the demography of the plants whose seeds they consume. Even where microsites are limited, the coincidence of the most suitable microhabitats for seedling establishment with those where seed predation is highest provide a means by which selective seed predators can influence community composition. Received: 19 August 1996 / Accepted: 25 January 1997     

Standard yet unusual mechanisms of long-distance dispersal: seed dispersal of Corymbia torelliana by bees

Mellitochory, seed dispersal by bees, has been implicated in long-distance dispersal of the tropical rain forest tree, Corymbia torelliana (Myrtaceae). We examined natural and introduced populations of C. torelliana for 4 years to determine the species of bees that disperse seeds, and the extent and distance of seed dispersal. The mechanism of seed dispersal by bees was also investigated, including fruit traits that promote dispersal, foraging behaviour of bees at fruits, and the fate of seeds. The fruit structure of C. torelliana , with seed presented in a resin reward, is a unique trait that promotes seed dispersal by bees and often results in long-distance dispersal. We discovered that a guild of four species of stingless bees, Trigona carbonaria, T. clypearis, T. sapiens , and T. hockingsi, dispersed seeds of C. torelliana in its natural range. More than half of the nests found within 250 m of fruiting trees had evidence of seed transport. Seeds were transported minimum distances of 20–220 m by bees. Approximately 88% of seeds were dispersed by gravity but almost all fruits retained one or two seeds embedded in resin for bee dispersal. Bee foraging for resin peaked immediately after fruit opening and corresponded to a peak of seed dispersal at the hive. There were strong correlations between numbers of seeds brought in and taken out of each hive by bees ( r =  0.753–0.992, P  < 0.05), and germination rates were 95 ± 5%. These results showed that bee-transported seeds were effectively dispersed outside of the hive soon after release from fruits. Seed dispersal by bees is a non-standard dispersal mechanism for C. torelliana, as most seeds are dispersed by gravity before bees can enter fruits. However, many C. torelliana seeds are dispersed by bees, since seeds are retained in almost all fruits, and all of these are dispersed by bees.     

Parental effects on seed mass: seed coat but not embryo/endosperm effects

Many biologists studying environmentally induced parental effects have indirectly suggested that the parental environment alters seed mass by altering the amount of endosperm or embryo tissue in the seed. We tested this hypothesis by measuring the effects of parental temperature on total seed mass, seed coat mass, and embryo/endosperm mass in offspring of Plantago lanceolata. Parental temperature significantly affected total seed and coat mass but not endosperm/embryo mass. Thus, larger seeds do not contain more resources in the embryo or endosperm than do small seeds. Rather they have more coat mass, which probably strongly influences germination. These results suggest caution when making assumptions about the pathways by which environmentally induced parental effects are transmitted in plant species. We also observed that controlled crosses differed significantly in their response to parental temperature, which provides evidence for genetic variation in environmentally induced parental effects, i.e., intergenerational phenotypic plasticity, in natural populations of P. lanceolata.