The role of pre- and post-dispersal seed predation in determining total seed loss

Most seed predation studies focus on either pre- or post-dispersal predation and may therefore underestimate the role of predation in regulating plant populations. We therefore estimated total seed predation of an invasive tree, mesquite (Leguminoseae: Prosopis spp.), by examining the entire seed pool from tree to seed bank. The spatio-temporal dynamics of total seed predation was examined by sampling across its Australian distribution and through time. The main predator was a host-specialist multivoltine beetle, Algarobius prosopis L. (Bruchidae), previously introduced as a biocontrol agent. Seed predation exceeded 20% in all seed stages (in pods on and off the tree, and seeds within woody endocarps (capsules) and free seeds on and in the ground) but was consistently highest in capsules on the ground (up to 90%). Pre-dispersal predation contributed little. Total seed predation rates were primarily determined by predation rates on the most persistent seed stage, in this case fallen pods if only pods are considered and seeds in capsules for the total seed pool. This pattern was consistent across the surveyed taxa, regions, years and seasonally. Predation rate was relatively unaffected by seed density, potentially because densities were always low (<150 seeds m⁻²). Average total seed predation within a region reached 55%, but we conclude that any population regulation of mesquite by seed predation will principally be through reduced seed bank persistence. Our results highlight the need to consider the entire seed pool, especially the often cryptic and overlooked long-lived stages, when determining seed loss to predation and its likely population consequences.     

Forest edges and fire ants alter the seed shadow of an ant-dispersed plant

Exotic species invade fragmented, edge-rich habitats readily, yet the distinct impacts of habitat edges and invaders on native biota are rarely distinguished. Both appear detrimental to ant-dispersed plants such as bloodroot, Sanguinaria canadensis. Working in northeastern Georgia (USA), an area characterized by a rich ant-dispersed flora, fragmented forests, and invasions by the red imported fire ant, Solenopsis invicta , I monitored the interactions between ants and S. canadensis seeds in uninvaded forest interiors, uninvaded forest edges, invaded forest interiors, and invaded forest edges. I observed 95% of the seed dispersal events that occurred within the 60-min observation intervals. Seed collection rates were similar among all four (habitat × invasion) groups. The presence of invasive ants had a strong effect on seed dispersal distance: S. invicta collected most seeds in invaded sites, but was a poorer disperser than four of five native ant taxa. Habitat type (interior versus edge) had no effect on seed dispersal distance, but it had a strong effect on seed dispersal direction. Dispersal towards the edge was disproportionately rare in uninvaded forest edges, and ants in those habitats moved the average dispersed seed approximately 70 cm away from that edge. Dispersal direction was also skewed away from the edge in uninvaded forest interiors and invaded forest edges, albeit non-significantly. This biased dispersal may help explain the rarity of myrmecochorous plants in younger forests and edges, and their poor ability to disperse between fragments. This is the first demonstration that forest edges and S. invicta invasion influence seed dispersal destination and distance, respectively. These forces act independently.     

Heat increases germination of water-permeable seeds of obligate-seeding <Emphasis Type="Italic">Darwinia</Emphasis> species (Myrtaceae)

We examined the response of seeds to heat in four geographically restricted and one widespread species of shrubby Darwinia from the fire-prone region of southeastern Australia. These shrubs are killed by fire and rely on seed germination after a fire to maintain populations. We replicated the germination trials across several sites and several fruiting seasons for most species. Seeds had a high level of viability and were largely dispersed in a dormant state, except in D. glaucophylla, where seed dormancy varied significantly across fruiting seasons. The indehiscent fruit of all species readily imbibes moisture when wet and seeds are not considered to be ‘hard-seeded’. All species had increased seed germination in response to a limited range of heating temperatures (generally 80–100°C). Higher temperatures killed increasing proportions of seeds. This pattern was broadly consistent across species, population and seasons, although the proportion of seeds whose germination was promoted by heat varied from high (D. diminuta, D. fascicularis, D. glaucophylla) to moderate (D. biflora, D. procera). Our work highlights the importance of heat as a mechanism for influencing germination in species that are not hard-seeded. Consequently, soil temperatures during a fire should strongly influence post-fire germination levels in Darwinia. The roles of other cues that promote germination, i.e. smoke, seasonal temperatures and their interactions with heat, remain to be investigated.     

Fire and flood: Soil‐stored seed bank and germination ecology in the endangered Carrington Falls Grevillea (Grevillea rivularis,Proteaceae)

Abstract Seed germination is dependent on the interaction between the dormancy state of a seed and the presence of favourable environmental conditions. Thus, the spectacular pulse of seedling recruitment in many Australian vegetation communities following disturbances such as fire can be attributed to changes in microsite conditions and/or the dormancy‐breaking effect of the disturbance on accumulated seed banks. Grevillea rivularis is a threatened species endemic to the area immediately above Carrington Falls in the NSW Southern Highlands. Most of the population is confined to the riparian vegetation zone in woodland and heath, and is therefore subject to periodic disturbance from fire and flood. For this species, a pulse of seedling recruitment has been recorded after fire, flood and mechanical soil disturbance. The aims of this study were to examine the density and vertical distribution of the soil‐stored seed bank and to investigate the role of heat and scarification as cues for germination of fresh and soil‐stored seed. There was a large seed bank under the canopies of established individuals (194 ± 73 seeds m^?2) and most seeds were found in the 0–2 cm and leaf‐litter layers of the soil profile. The germination response of soil‐stored and fresh seed was examined using a hierarchical series of laboratory experiments. Seeds of G. rivularis showed marked dormancy polymorphism. Thirty‐six percent of soil‐stored seed germinated without treatment, whereas no untreated fresh seeds germinated. Scarification or heating caused significant germination of dormant soil‐stored seed, but only scarification resulted in germination of dormant fresh seeds. These results highlight important differences in the dormancy state of soil‐stored and fresh seed. Thus, being a riparian species in a fire‐prone environment, the dormancy mechanisms in seeds of G. rivularis suit this species to disturbance by both fire and flood.     

Opposing forces of seed dispersal and seed predation by mammals for an invasive cactus in central Kenya

The invasive erect prickly pear cactus (Opuntia stricta) has reduced rangeland quality and altered plant communities throughout much of the globe. In central Kenya's Laikipia County, olive baboons (Papio anubis) frequently consume O. stricta fruits and subsequently disperse the seeds via defecation. Animal‐mediated seed dispersal can increase germination and subsequent survival of plants. However, consumption of seeds (seed predation) by rodents may offset the potential benefits of seed dispersal for cactus establishment by reducing the number of viable seeds. We investigated foraging preferences of a common and widely distributed small mammal—the fringe‐tailed gerbil (Gerbilliscus robustus), between O. stricta seeds deposited in baboon faeces versus control O. stricta seeds. In addition to providing evidence of seed predation on O. stricta by G. robustus, our data show that seed removal was higher (shorter time to use) for seeds within faeces than for control seeds. G. robustus clearly prefers seeds within faeces compared to control seeds. These results suggest that high abundances of rodents may limit successful establishment of O. stricta seeds, possibly disrupting seed dispersal via endozoochory by baboons.     

Soil seed bank dynamics in the rare obligate seeding shrub,Grevillea barklyana (Proteaceae)

Abstract Grevillea barklyana is a rare, obligate seeding shrub that occurs in fire-prone heath and woodland in southeastern Australia. The size and dynamics of the soil seed bank were investigated in populations 2–29 years after fire in order to identify fire frequencies that would threaten population survival. Plants began flowering 2 years after fire (16%), but maximum inflorescence and seed production did not occur until 15–16 years after fire when plants were full size. Seed production declined in older populations. Fruit:flower ratios were very low (0.1–5.3%). Most flowers (> 95%) failed to initiate fruits. Of the initiated fruits, about 60% aborted, 27% were attacked by parrots and 13% matured. An exclusion experiment showed that fruits attacked by parrots would have otherwise matured. Seeds were consumed on the soil surface by mammals, particularly rodents. Post-dispersal seed losses were consistently high in all months and in all populations (78–100%). Mortality of adult plants increased with time since fire. In the 29-year-old population, 30% of plants were dead and 33% were senescent. Recruitment was negligible in the absence of fire, but following fire the seedling to parent ratio was 20:1. Seedling survival was high (90%) following two years of above-average rainfall. The observed seed bank was small (3–16 seeds irr) and increased during the first 16 years after fire before declining in the 21- and 26-year-old populations. The size and dynamics of the seed bank may limit population numbers, especially if the intervals between fires are less than 10 years or greater than 25 years.     

Dispersal and recruitment dynamics in the fleshy‐fruited Persoonia lanceolata (Proteaceae)

Question: What is the role of dispersal, persistent soil seed banks and seedling recruitment in population persistence of fleshy‐fruited obligate seeding plant species in fire‐prone habitats? Location: Southeastern Australia. Methods: We used a long‐term study of a shrubby, fleshy‐fruited Persoonia species (Proteaceae) to examine (1) seed removal from beneath the canopy of adult plants; (2) seedling recruitment after fire; (3) the magnitude and location of the residual soil seed bank; and (4) the implications for fire management of obligate seeding species. We used demographic sampling techniques combined with Generalised Linear Modelling and regression to quantify population changes over time. Results: Most of the mature fruits (90%) on the ground below the canopy of plants were removed by Wallabia bicolor (Swamp wallaby) with 88% of seeds extracted from W. bicolor scats viable and dormant. Wallabies play an important role in moving seeds away from parent plants. Their role in occasional long distance dispersal events remains unknown. We detected almost no seed predation in situ under canopies (< 1%). Seedling recruitment was cued to fire, with post‐fire seedling densities 6‐7 times pre‐fire adult densities. After fire, a residual soil seed bank was present, as many seeds (77‐100%) remained dormant and viable at a soil depth where successful future seedling emergence is possible (0‐5 cm). Seedling survival was high (> 80%) with most mortality within 2 years of emergence. Plant growth averaged 17 cm per year. The primary juvenile period of plants was 7–8 years, within the period of likely return fire intervals in the study area. We predicted that the study population increased some five‐fold after the wildfire at the site. Conclusions: Residual soil seed banks are important, especially in species with long primary juvenile periods, to buffer the populations against the impact of a second fire occurring before the seed bank is replenished.     

Aerial seed bank and dispersal traits in Anthemis chrysantha (Asteraceae), a critically endangered species

Mature plant density and fruit production were monitored in the main population of four successive cohorts of the endangered winter annual Anthemis chrysantha (Asteraceae) in southeastern Spain. Experiments were conducted with artificial rainfall and a wind tunnel to determine the temporal and spatial dispersal pattern of the species and the relationship with rain and wind.     

Roles of species-preferential seed dispersal by ants and endozoochory in Melampyrum (Orobanchaceae)

Aims Melampyrum pratense and M. subalpinum are two myrmecochorous species, which possess similar habitat requirements and frequently occur together. Despite this, their population sizes differ markedly. Melampyrum pratense populations are usually very large, whereas M. subalpinum has rather small and isolated populations. We suggest that such an imbalance might be partially influenced by the difference in ant-mediated seed-removal rates. Genus Melampyrum is considered to be exclusively myrmecochorous, though to achieve the recent distribution of some Melampyrum species during the Holocene myrmecochory would be highly insufficient. We suggest that endozoochory takes place in the long-distance migration, whereas myrmecochory is important for the removal of seeds on a local scale.Methods For seed-preference analysis, M. pratense and M. subalpinum mixed seed samples were placed around Formica polyctena anthills. After a period of time, the remaining seeds of both species were counted for each sample. The results were analysed by analysis of variance and generalized linear mixed-effect model. To test myrmecochorous removal distances, M. pratense seeds were covered with fluorescent dactyloscopic powder and placed in the vicinity of a large ant trail. The area around the starting plot was searched in the dark using UV LED torchlight 7h after the beginning. The distance from the starting plot was measured for each seed found. Birds, rodents, leporine and a ruminant were fed with M. pratense seeds and fresh plants to test the possibility of endozoochorous dispersal of the species. Animal droppings were searched for intact seeds.Important findings Our field studies show that from mixed seed samples, containing both species, ants significantly preferred the seeds of M. pratense. This may be one of factors that has positive influence on M. pratense success in seed dispersal on mixed stands and consequently in the colonization of favourable sites. Experiments focusing on ant-mediated dispersal distance revealed that F. polyctena ants are able to move seeds over a distance of 36 m in only 7h. This distance is among the furthest known myrmecochorous removals of forest plant seeds. A new Melampyrum seed disperser Oligolophus tridens (Opiliones) was observed repeatedly. Our pilot study documented that Melampyrum seeds are able to pass through the digestive tract of a cow intact. This suggests that large ruminants such as deer, bison or forest-grazing livestock may function as important long-distance dispersers of Melampyrum species.     

The size,distribution and germination requirements of the soil-stored seed-bank of Grevillea barklyana (Proteaceae)

Abstract Fire-triggered release from seed dormancy is a characteristic of many Australian plant species. We investigated aspects of the seed-bank dynamics and dormancy characteristics in seeds of Grevillea barklyana, an understorey shrub of coastal sclerophyll vegetation in the Jervis Bay Region on the south coast of New South Wales. We used two soil core sizes to compare the number and distribution of stored seeds in soil cores taken from underneath and outside the limits of plant canopies at three study sites. Core size did not affect the estimate of seed density. No seeds were found outside the existing canopies. Even in under-canopy samples, seed numbers were small. Mean seed densities were estimated as 10. 9, 14. 1 and 4. 3 seeds per for the three sites. A hierarchical series of laboratory experiments was used to test the germination response of both fresh and soil-stored seeds. Likewise, we attempted to simulate ‘natural’ disturbance conditions in a glasshouse experiment. Both laboratory and glasshouse experiments indicated polymorphism in germination behaviour. A constant proportion of seeds exhibited enforced dormancy, when moist at room temperature, while a smaller number of seeds showed either an induced dormancy or a non-seed-coat linked innate dormancy. The majority (75%) of seeds were innately dormant due to a hard seed-coat. This dormancy was broken when the seed-coat was damaged, for instance, by heat. The level of polymorphic germination behaviour will be dependent on the length of the inter-fire periods. We conclude that the expression of polymorphism within and between species across a range of environments and fire regimes is an important consideration for any further study attempting to assess the role of the seed-bank.     

The role of water chemistry in the distribution of Austropotamobius pallipes (Crustacea Decapoda Astacidae) in Piedmont (Italy)

Over the last decades, the populations of Austropotamobius pallipes have decreased markedly all over Europe. If we evaluate the ecological factors that determine its presence, we will have information that could guide conservation decisions. This study aims to investigate the chemical-physical demands of A. pallipes in NW Italy. To this end, we investigated 98 sites. We performed Principal Component Analysis using chemical-physical parameters, collected in both presence and absence sites. We then used principal components with eigenvalue > 1 to run Discriminant Function Analysis and Logistic Regression. The statistics on the concentration of Ca²⁺, water hardness, pH and BOD₅ were significantly different in the presence and in the absence sites. pH and BOD₅ played the most important role in separating the presence from the absence locations. These findings are further evidence that we should reduce dissolved organic matter and fine particles in order to contribute to species management and conservation.     

Effects of ants,ground beetles and the seed-fall patterns on myrmecochory of Erythronium japonicum Decne. (Liliaceae)

Erythronium japonicum (Liliaceae) inhabits deciduous mesic forests of Hokkaido, northern Japan. Myrmecochory of this species was investigated, especially the dispersal frequency, the effect of seed predators and the seed fall pattern. In the quadrat census using marked seeds of E. japonicum, the ant Myrmica kotokui frequently transported the seeds. However, the frequency of seed removal was low and most seeds were dispersed as little as 1 m or less. The spatial distribution of E. japonicum individuals was nearly random and most seedlings were established 5–20 cm away from the fertile plants, indicating that even this small scale of seed dispersal contributes to avoiding crowding of seedlings. Some arthropods, e.g. springtails, spiders and ticks, hindered seed dispersal by devouring elaiosomes and seeds. Although ground beetle species also damaged seeds and elaiosomes, a few of them exhibited seed removal behaviour. E. japonicum dropped their seeds not all at once but bit by bit, taking 3–6 days to drop all seeds. This seed-fall pattern was effective in raising the frequency of seed removal by ants and reducing seed predation by some arthropods.     

Integrating population demography, genetics and self-incompatibility in a viability assessment of the Wee Jasper Grevillea (Grevillea iaspicula McGill., Proteaceae)

Grevillea iaspicula is an endangered shrub known from only eight small populations (<250 individuals) in south-eastern Australia. The species is threatened by combined ecological and genetic factors, e.g. land conversion, weed invasion, low recruitment and low gene flow among populations. The populations also show large variance in male fitness and limited mate availability which are thought to arise as a consequence of gametophytic self-incompatibility (GSI). This study has used an individual-based, spatially explicit simulation model to explore the interaction between GSI and mate limitation in this species, as well as its effect on long-term population viability. The model was parameterised with demographic and genetic data obtained from 2 years of population monitoring. Simulation results identified extremely low establishment rates as the most critical factor currently influencing the persistence of G. iaspicula populations and indicated that the extant populations are at serious risk of extinction in the near future unless this is altered by, at very least, an order of magnitude higher. SI was shown to affect the magnitude of variation in establishment but this effect was masked when establishment was critically low. Disassortative mating, owing to low allelic richness at the S-locus, had the negative demographic effect of restricting mating to relatively few compatible plants. Restricted mate availability imposed additional limitations to the viability of populations but, given a 20-fold increase in establishment rate, population fluctuations stabilised. The long-term viability of G. iaspicula is bleak without artificial augmentation of the populations but management planning must also consider genetic processes, including SI, to ensure such strategies optimise the benefits gained.     

Dissection and Preparation of Whole Mounts of Endosperm from the Seeds of Grevillea (Proteaceae)

It is sometimes very desirable to supplement the examination of serially cut microtome sections by dissection and study of whole mounts of objects. In the case of the seeds of Grerillea the endosperm forms a curious worm-like structure, the vermiform appendage, in elucidating the nature of which microtome sections alone are inadequate. The dissection of the endosperm for demonstrating this structure is very necessary; otherwise the structure is altogether overlooked. In dissecting out the endosperm, which is easily done, either fresh seeds or previously preserved seeds may be used; the former are easier for handling and yield more satisfactory results. In the case of fresh material, an immediate killing is necessary and any one of the ordinary killing fluids may be used; Bouin's fluid was tried by the writer and was entirely satisfactory. After killing, the material is stained and dehydrated, the several stages of which are best done on the slide alone on which the material is placed. Mounting is done in any of the mounting media, and Canada balsam answers the purpose quite well.     

The role of spines in anthropogenic seed dispersal on the Galápagos Islands

1. Dispersal has important ecological and evolutionary consequences for populations, but understanding the role of specific traits in dispersal can be difficult and requires careful experimentation. Moreover, understanding how humans alter dispersal is an important question, especially on oceanic islands where anthropogenic disturbance through species introductions can dramatically alter native ecosystems.
2. In this study, we investigated the functional role of spines in seed dispersal of the plant caltrop (Tribulus cistoides L., Zygophyllaceae) by anthropogenic dispersal agents. We also tested whether humans or wildlife are more important seed dispersers of T. cistoides on the Galápagos.
3. Tribulus cistoides is found on tropical mainland and oceanic island habitats. The dispersal structure of T. cistoides is called a mericarp, and they are typically protected by one pair of upper spines and a second pair of lower spines, but the presence and size of spines varies within and between populations. On the Galápagos, the upper and lower spines protect mericarps from seed predation by Darwin's finches. We tested whether spines play a dual role in dispersal by factorially manipulating the presence/absence of the upper and lower spines to simulate natural variation in mericarp morphology.
4. The upper spines greatly facilitated seed dispersal, whereas the lower spines had no discernible effect on dispersal. The presence of upper spines increased dispersal rate on shoes by pedestrians 23‐fold, on fabrics (e.g., towels) and cars by nearly twofold, and the presence of upper spines increased dispersal distance by cars sixfold. When comparing dispersal rates in habitats with high (roads and foot paths) versus low (arid forest) anthropogenic activity, dispersal rates were demonstrably higher in the habitats with more human activity.
5. These results have important implications for understanding the ecology and evolution of plant dispersal in the Anthropocene. Spines on the fruits of T. cistoides play important functional roles in anthropogenic dispersal, whereas native and introduced wildlife plays a minor role in dispersal on inhabited islands of the Galápagos. Our results imply that seed predators and humans are jointly shaping the ecology and evolution of contemporary populations of T. cistoides on the Galápagos.

     

The phylogeny and biogeography of Hakea (Proteaceae) reveals the role of biome shifts in a continental plant radiation

The frequency of evolutionary biome shifts during diversification has important implications for our ability to explain geographic patterns of plant diversity. Recent studies present several examples of biome shifts, but whether frequencies of biome shifts closely reflect geographic proximity or environmental similarity of biomes remains poorly known. We explore this question by using phylogenomic methods to estimate the phylogeny of Hakea, a diverse Australian genus occupying a wide range of biomes. Model‐based estimation of ancestral regions indicates that Hakea began diversifying in the Mediterranean biome of southern Australia in the Middle Eocene–Early Oligocene, and dispersed repeatedly into other biomes across the continent. We infer around 47 shifts between biomes. Frequencies of shifts between pairs of biomes are usually similar to those expected from their geographic connectedness or climatic similarity, but in some cases are substantially higher or lower than expected, perhaps reflecting how readily key physiological traits can be modified to adapt lineages to new environments. The history of frequent biome‐shifting is reflected in the structure of present‐day assemblages, which tend to be more phylogenetically diverse than null‐model expectations. The case of Hakea demonstrates that the radiation of large plant clades across wide geographic areas need not be constrained by dispersal limitation or conserved adaptations to particular environments.     

Limitations to reproductive output and genetic rescue in populations of the rare shrub Grevillea repens (Proteaceae)

Background and Aims

When conserving rare plant species, managers are often faced with small and/or isolated populations displaying low levels of sexual reproduction and genetic variation. One option for reinvigorating these populations is the introduction of genetic material from other sites, but in some cases fitness may be reduced as a result of outbreeding depression. Here the pollination biology of the rare shrub Grevillea repens is studied across its natural range and reproductive responses following cross-pollination among populations are examined to determine factors that may be limiting sexual reproduction and the potential for genetic rescue.

Methods

Pollen manipulation treatments (self-, autogamous self-, cross- and open pollination) were applied to flowers to examine the breeding system and fruit and seed production in five populations of G. repens. Pollen production, presentation and viability were investigated and interpopulation crosses of increasing genetic distance performed among the populations.

Key Results

The study species is self-incompatible and displayed very low natural seed set over two seasons, due partly to low pollen viability in one of the populations. Within-population crossing increased fruit and seed production at some sites, indicating pollinator limitation. Interpopulation crosses further increased reproductive output in one population, suggesting mate limitation, and for this site there was a positive relationship between genetic distance among populations and the size of genetic rescue benefits. However, in other populations there was a decrease in fruit and seed set with increasing genetic distance.

Conclusions

The results highlight that management strategies involving interpopulation crosses can improve reproductive output in small, isolated populations of rare plants, but guidelines need to be developed on a population by population basis.Key words: Grevillea repens, Proteaceae, genetic rescue, pollination ecology, self-incompatibility, breeding system, interpopulation cross, outbreeding depression, pollinator limitation, mate limitation, resource limitation