Accumulation of buried seeds and establishment of ruderal therophytic communities in disturbed habitat,central Japan

The establishment and maintenance mechanisms of pioneer communities were investigated in ruderal habitats under two disturbance regimes, frequent and infrequent cutting sites. In the infrequent cutting sites, large perennials dominated through the year and inhibited the invasion of annuals, and the perennial community succeeded to forest stage if the cutting was stopped. In the frequent cutting sites, therophytic communities of winter and summer annuals alternated by season. Fresh seeds of both winter and summer annuals are dormant, but they have different germination times and thus can share the same sites in different seasons. Wind-dispersed biennials and large perennials have nondormant seeds and easily invade the sites; however, they are unable to mature to reproductive phase due to recurrent cuttings. The therophytic species, which can complete their life-cycle in a period between cuttings, accumulate seeds in the soils and are maintained by these buried seeds (seed bank annuals) during recurrent disturbances. The seed bank is compensation for the dispersal inefficiency of seed bank annuals. After abandonment of the frequent cutting sites, the buried seeds of seed bank annuals germinate and become the first-year pioneers. Thus, seed bank annuals are not invasive colonizers but are the remnants of the ruderal weed communities before abandonment.     

Restoring a Mediterranean‐climate shrub community with perennial species reduces future invasion

Successful restoration of an invaded landscape to a diverse, invasion‐resistant native plant community requires determining the optimal native species mix to add to the landscape. We manipulated native seed mix (annuals, perennials, or a combination of the two), while controlling the growth of non‐native species to test the hypothesis that altering native species composition can influence native establishment and subsequent non‐native invasion. Initial survival of native annuals and perennials was higher when seeded in separate mixes than when combined, and competition between the native perennials and annuals led to lower perennial cover in year 2 of mixed‐seeded plots. The plots with the highest perennial cover had the highest resistance to invasion by Brassica nigra. To clarify interactions among different functional groups of natives and B. nigra, we measured competitive interactions in pots. We grew one native annual, one native perennial, and B. nigra alone or with different competitors and measured biomass after 12 weeks. Brassica nigra was the strongest competitor, limiting the growth of all native species, and was not impacted by competition with native annuals or perennial seedlings. Results from the potted plant experiment demonstrated the strong negative influence of B. nigra on native seedlings. Older native perennials were the strongest competitors against invasive species in the field, yet perennial seedling survival was limited by competition with native annuals and B. nigra. Management action that maximizes perennial growth in early years may lead to a relatively more successful restoration and the establishment of an invasion‐resistant community.     

How do steppe plants follow their optimal environmental conditions or persist under suboptimal conditions? The differing strategies of annuals and perennials

For a species to be able to respond to environmental change, it must either succeed in following its optimal environmental conditions or in persisting under suboptimal conditions, but we know very little about what controls these capacities. We parameterized species distribution models (SDMs) for 135 plant species from the Algerian steppes. We interpreted low false‐positive rates as reflecting a high capacity to follow optimal environmental conditions and high false‐negative rates as a high capacity to persist under suboptimal environmental conditions. We also measured functional traits in the field and built a unique plant trait database for the North‐African steppe. For both perennial and annual species, we explored how these two capacities can be explained by species traits and whether relevant trait values reflect species strategies or biases in SDMs. We found low false‐positive rates in species with small seeds, flowers attracting specialist pollinators, and specialized distributions (among annuals and perennials), low root:shoot ratios, wide root‐systems, and large leaves (perennials only) (R² = .52–58). We found high false‐negative rates in species with marginal environmental distribution (among annuals and perennials), small seeds, relatively deep roots, and specialized distributions (annuals) or large leaves, wide root‐systems, and monocarpic life cycle (perennials) (R² = .38 for annuals and 0.65 for perennials). Overall, relevant traits are rarely indicative of the possible biases of SDMs, but rather reflect the species' reproductive strategy, dispersal ability, stress tolerance, and pollination strategies. Our results suggest that wide undirected dispersal in annual species and efficient resource acquisition in perennial species favor both capacities, whereas short life spans in perennial species favor persistence in suboptimal environmental conditions and flowers attracting specialist pollinators in perennial and annual species favor following optimal environmental conditions. Species that neither follow nor persist will be at risk under future environmental change.     

Ovule survivorship,brood size,life history,breeding systems,and reproductive success in plants

The percentage of ovules developing into seeds (seed-ovule ratios, S/O ratios) is about 85% in annuals but only approximately 50% in perennials.In both annuals and perennials, these typical S/O ratios occur regardless of the kind of breeding system, although many annuals are normally self-pollinating whereas perennials are virtually all cross-pollinating. The mean number of seeds maturing within individual fruits is defined as brood size, and is correlated with different modes of dispersal and strategies of ovule packging. Annuals also have significantly higher brood sizes (21.7) than perennials (<9.9). Among perennials, woody plants have both lower S/O ratios (32.7%) and brood sizes (3.3) than herbaceous perennials (57.2%, 13.5). S/O ratios appear to be largely determined genetically, whereas resource limitations are perhaps more critical in terms of regulating flower production. Among perennials, increased exposure to predators and pathogens is suggested as the best explanation for theevolution of breeding systems favoring genetic recombination. The maintenance of genetic polymorphisms, however, inevitably increases the frequency of lethal and sub-lethal allelic combinations (and perhaps mutations?), that appear to be responsible for the lower S/O ratios in perennials.     

The Role of Currents and Waves in the Dispersal of Submersed Angiosperm Seeds and Seedlings

We tested the hypothesis that currents, waves, and sediment grain size affect the dispersal of seeds and seedlings of the submersed angiosperms Ruppia maritima, Potamogeton perfoliatus and Stuckenia pectinata. Seed settling velocities and initiation of motion of seeds and seedlings and distance transported were quantified on four sediment types under a range of currents and waves in a flume. The rapid settling velocities of R. maritima and S. pectinata seeds and the increased settling velocity of P. perfoliatus in currents above 8 cm/second suggest that primary dispersal of these species is localized to the general area colonized by their parents. Once settled within a bed, seeds are exposed to weak currents and waves, and are likely to be subject to sediment deposition which may further limit dispersal. In contrast, in restoration projects, the absence of vegetation is likely to make seeds more vulnerable to grazing and transport, and may contribute to the lack of plant establishment. If seeds germinate without being buried, they are susceptible to secondary dispersal at relatively low current velocities and small wave heights due to the drag exerted on the cotyledon. Sand grains tend to stick to the seed coat and rootlet of P. perfoliatus seedlings, perhaps a mechanism to reduce the chances of being displaced following germination. These data reveal the close links between sediment, water flow, and submersed angiosperm seedling establishment; these parameters should be considered when using seeds for restoration of submersed angiosperms.     

Fate of plants from buried seeds on Volcano Usu,Japan, after the 1977–1978 eruptions

The eruptions of 1977–1978 on Mount Usu in northern Japan resulted in the almost complete destruction of vegetation by a 1–3-m-thick accumulation of volcanic deposits. Erosion created gullies that removed these deposits and frequently exposed the old original soil. I previously confirmed that seedlings of 14 species (eight annual and six perennial herbs) emerged from seeds buried in the original topsoil. To clarify the role of the seed bank on volcanic succession, I monitored seedlings of seed bank species from 1983 to 1992. Nearly all the annuals, such as Polygonum longisetum and Rorippa islandica, originated from the seed bank. However, seed supply from the seed bank declined with time. Because seedling mortality was extremely high and reproductive success was low, due mostly to the instability of ground surface, these annuals emerged for several years but disappeared after 1989. The nitrogen-fixing and stoloniferous perennials, Trifolium repens and Lotus corniculatus var. japonicus, were derived only from the seed bank. They have gradually increased in cover and have become large enough to flower. The other perennials derived from the seed bank, most of which produce short rhizomes, did not increase in cover annually and thus contributed less to revegetation than nitrogen-fixing stoloniferous species. The fate of perennials seems to be dependent on the types of rhizome and stolon systems. Although recruitment from the seed bank was restricted to gullies, the seed bank was the major source of annuals and of nitrogen-fixing perennials in the gullies. Succession in the gullies was substantially different from that which occurred outside the gullies where plants mostly originated from seed immigration and vegetative reproduction from surviving plants, and annual and nitrogen-fixing plants were absent.     

Seed dispersal of alien and native plants by vertebrate herbivores

Seed dispersal is crucial for the success and spread of alien plants. Herbivores often establish a dual relationship with plants: antagonist, through herbivory, and mutualist, through seed dispersal. By consuming plants, herbivores may disperse large amounts of seeds, and can facilitate the spread of alien plants. However, seed dispersal of alien plants by herbivores has been largely uninvestigated. I studied factors associated with dispersal of alien and native seeds by the three most important vertebrate herbivores in SW Australia: emus (Dromaius novaehollandia), western grey kangaroos (Macropus fuliginosus) and European rabbits (Oryctolagus cuniculus). Overall frequencies of alien and native seeds dispersed by these herbivores were determined by differences among them in (1) the plant groups they predominantly disperse, that differed in frequencies of aliens versus natives, and (2) the predominant dispersal of aliens or natives within those plant groups. Emus and kangaroos (natives) tended to disperse predominantly alien seeds within plant groups (defined by life forms, dispersal syndromes, and diaspore size), whereas rabbits (alien) tended to disperse predominantly natives. This agrees with the hypothesis that herbivores will use predominantly plants that have evolved in different areas, because of less effective defences against new enemies. Overall frequencies were consistent with this pattern in kangaroos and rabbits, but not in emus. Kangaroos dispersed mostly plant groups that were mainly aliens (herbaceous species and small and medium sized dispersal units and seeds), which together with their predominant use of aliens over natives within groups resulted in the highest overall frequency of alien seeds (73%). Rabbits were similar to kangaroos in the type of plants dispersed, but their predominant use of natives over aliens within groups contributed to an overall predominance of native seeds in their pellets (88%). Emus dispersed mostly plant groups that were mainly natives (e.g. woody species with big diaspores), resulting in low overall frequency of alien seeds (11%), despite their predominant use of aliens over natives within plant groups. Thus, the within-groups trend pointed to a facilitative role of native herbivores of plant invasions through seed dispersal, but was obscured by the different use by herbivores of plant groups with different frequency of aliens.     

FACILITATION OF SUCCESSION BY THE NONMYCOTROPHIC COLONIZER SALSOLA KALI (CHENOPODIACEAE) ON A HARSH SITE: EFFECTS OF MYCORRHIZAL FUNGI

The effects of inoculation with vesicular-arbuscular mycorrhizal fungi on nonmycotrophic (plants which do not form a mycorrhizal association) colonizing annuals and on mycotrophic perennials were investigated in southwestern Wyoming. A subsoil containing no initial mycorrhizal inoculum was used, and seeds of the later successional perennials were planted. The annuals were removed from one-half of the sampling quadrats to test for the interaction of competition and mycorrhizae on the perennials. During the third and fourth years of succession, the density and percent cover of Salsola kali, the most abundant nonmycotrophic annual, decreased by one-half to one-third with inoculation. Mycorrhizal hyphae and spores were found in the rhizosphere of S. kali, with only occasional (1–2% of the root length) penetration of hyphae into the cortex. There were no differences in tissue phosphorus and nitrogen concentrations or water relations of inoculated vs. uninoculated S. kali. The planted grasses, all Agropyron species, had no significant increase in density or percent cover with inoculation. The percent root infection of A. smithii was 5–30%. Grass density and percent cover was greatest where S. kali was present, suggesting facilitation of grass establishment by annuals on this harsh, windy site. However, grass density was lower where 5. kali density was lower following inoculation. Mycorrhizal fungi were hypothesized to hasten the rate of succession on other sites. On this site where facilitation is an important process, inoculation of early seral plants and their subsequent decline may slow primary succession in the early years.     

Flowering requirements of Polymnia canadensis (Asteraceae) and their influence on its life history variation

The purpose of this study is to determine the flowering requirements of Polymnia canadensis and how they correspond to the occurrence of winter annuals, biennials, and short-lived monocarpic perennials in this species. Polymnia canadensis has a vernalization requirement for flowering, and even very small plants (i.e., those with one pair of leaves) can be vernalized. Vernalized plants can flower under both long- and short days. However, to flower plants must attain a minimum postvernalization size. Plants of this primarily monocarpic species that do not die after they flower once require another period of vernalization to flower a second time (i.e., to be dicarpic). Vernalized plants exposed to high temperatures can be devernalized; these must be re-vernalized in order to flower.     

GERMINATION ECOPHYSIOLOGY OF HERBACEOUS PLANT SPECIES IN A TEMPERATE REGION

Germination phenology data have been collected from 75 winter annuals, 49 summer annuals, 28 monocarpic perennials, and 122 polycarpic perennials, and experimental investigations of dormancy breaking and germination requirements have been conducted on 56 winter annuals, 32 summer annuals, 18 monocarpic perennials, and 73 polycarpic perennials. The purpose of these studies was to determine if there are correlations between the dormancy breaking and germination requirements of seeds and the germination phenology, life cycle type, habitat requirements, range of geographical distribution, and phylogenetic relationships of the species. Germination phenology is highly correlated with the responses of seeds to the yearly temperature cycle. Species with winter and summer annual life cycles have predictable germination characteristics, but monocarpic and polycarpic perennials do not. Several dormancy types may be found in a given habitat, and narrowly endemic and widely-distributed species in the same genus may have similar germination characteristics. Within some families there is a tendency for a particular type of seed-temperature response to be very important, but frequently this is related to the predominance of a given life cycle type in the family.     

Variation in the establishment of a non-native annual grass influences competitive interactions with Mojave Desert perennials

Competition between native and non-native species can change the composition and structure of plant communities, but in deserts, the highly variable timing of resource availability also influences non-native plant establishment, thus modulating their impacts on native species. In a field experiment, we varied densities of the non-native annual grass Bromus madritensis ssp. rubens around individuals of three native Mojave Desert perennials—Larrea tridentata, Achnatherum hymenoides, and Pleuraphis rigida—in either winter or spring. For comparison, additional plots were prepared for the same perennial species and seasons, but with a mixture of native annual species as neighbors. Growth of perennials declined when Bromus was established in winter because Bromus stands had 2–3 months of growth and high water use before perennial growth began. However, water potentials for the perennials were not significantly reduced, suggesting that direct competition for water may not be the major mechanism driving reduced perennial growth. The impact of Bromus on Larrea was lower than for the two perennial grasses, likely because Larrea maintains low growth rates throughout the year, even after Bromus has completed its life cycle. This result contrasts with the perennial grasses, whose phenology completely overlaps with (Achnatherum) or closely follows (Pleuraphis) that of Bromus. In comparison, Bromus plants established in spring were smaller than those established in winter and thus did not effectively reduce growth of the perennials. Growth of perennials with mixed annuals as neighbors also did not differ from those with Bromus neighbors of equivalent biomass, but stands of these native annuals did not achieve the high biomass of Bromus stands that were necessary to reduce perennial growth. Seed dormancy and narrow requirements for seedling survivorship of native annuals produce densities and biomass lower than those achieved by Bromus; thus, impacts of native Mojave Desert annuals on perennials are expected to be lower than those of Bromus.     

Patterns of Seed Dispersal and Dispersal Failure in a Hawaiian Dry Forest Having Only Introduced Birds

Dry forests are among the most endangered natural communities in the Hawaiian Islands. Most have been reduced to isolated trees and small forest fragments in which native tree species reproduce poorly. The replacement of native birds by introduced generalists may be contributing to dry forest decline through modification of seed dispersal patterns. To document seed dispersal by introduced birds, we conducted foraging observations on fleshy-fruited trees and measured seed rain under trees and in adjacent open areas for 1 year in a dry forest dominated by native trees. Although trees covered only 15.2 percent of the study area, 96.9 percent of the bird-dispersed seeds were deposited beneath them. The Japanese white-eye (Zosterops japonicus) was the principal dispersal agent. Among bird-dispersed seeds, those of the invasive tree Bocconia frutescens accounted for 75 percent of all seeds collected beneath trees (14.8 seeds/m²/yr) and the invasive shrub Lantana camara accounted for 17 percent. Although nearly 60 percent of the reserve's native woody species possess fleshy fruits, introduced birds rarely disperse their seeds. Native trees accounted for <8 percent of all bird-dispersed seeds and are consequently experiencing dispersal failure by falling directly under parent trees. Smaller-seeded non-native plants, in contrast, may be benefiting from dispersal by introduced birds. Current dispersal patterns suggest that these readily disseminated non-native plants may eventually replace the remaining native flora.     

Interactions between annuals and woody perennials in a Western Australian nature reserve

Abstract. We studied the interactions between woody perennial species and native and non-native annual species in a number of vegetation types within a nature reserve in the Western Australian wheatbelt. In particular, we examined the responses of annuals to perennial canopy removal, fire, soil disturbance and nutrient additions, and the effects of removal of annuals on perennial seedling regeneration. Experimental shrub removal significantly increased the abundance of annuals in a dense shrubland dominated by Allocasuarina campestris, but had no effect in a more open species-rich sandplain heath. Soil disturbance and nutrient addition in the heath area had no significant influence on annual abundance until three years after treatment. Fire had no clear effect on annual abundance in the heath within the reserve, but promoted a large increase in non-native species within an adjacent roadverge. A pattern of increased soil nutrient levels was accompanied by greatly increased non-native annual abundance beneath individual trees of Santalum spicatum. Exploratory laboratory bioassay experiments indicated that several woody perennials produced leachates that were capable of reducing the germination or growth of the introduced grass Avena fatua, indicating that allelopathy may be an important component of the interaction between the annual and perennial components. Within a woodland community, fire temporarily reduced the abundance of annual species and increased the establishment of perennial seedlings. Field experiments showed that annuals significantly reduced the survival of seedlings of the shrub Allocasuarina campestris. Our results indicate that intact native vegetation canopies effectively prevent invasion by non-native annuals, and that regeneration by native perennials is likely to be inhibited by the presence of an abundant annual cover.     

科尔沁草甸草地放牧和割草条件下土壤种子库研究

研究了科尔沁沙地草甸草地在放牧、割草两种利用方式下的土壤种子库的大小、组成及其与地上植被的关系．结果表明,在自由放牧下,土壤种子库密度为6158±1647粒·m^-2,在割草利用下,土壤种子库密度为8312±2540粒·m^-2．放牧干扰下,种子库组成以矮小、短命的一年生植物为主,其中一二年生植物占81．66％,占比例最大的前4种植物分别为虎尾草(38．55％)、灰绿藜(15．42％)、毛马唐(14．95％)和狗尾草(9．83％),多年生植物种子密度仅为1129±302粒·m^-2;割草干扰下,一二年生植物占68．08％,其中狗尾草占52．7％;而割草地多年生植物种子密度为2653±811粒·m^-2．放牧地土壤种子库密度与地上植物相关不显著;割草地土壤种子库密度与地上植物多度显著相关(r=0．76,P<0．01)．放牧地种子库的Shannon—Wiener指数、丰富度指数分别为2．96和2．98,明显小于割草地的3．10和5．09,表明自由放牧更易使物种多样性下降．     

Restoring prairie pothole wetlands: does the species pool concept offer decision‐making guidance for re‐vegetation?

Question: Do regional species pools, landscape isolation or on‐site constraints cause plants from different guilds to vary in their ability to colonize restored wetlands? Location: Iowa, Minnesota, and South Dakota, USA. Methods: Floristic surveys of 41 restored wetlands were made three and 12 years after reflooding to determine changes in local species pools for eight plant guilds. The effect of landscape isolation on colonization efficiency was evaluated for each guild by plotting local species pools against distance to nearby natural wetlands, and the relative importance of dispersal vs. on‐site constraints in limiting colonization was explored by comparing the local species pools of restored and natural wetlands within the region. Results: Of the 517 wetland plant taxa occurring in the region, 50% have established within 12 years. The proportion of the regional species pool represented in local species pools differed among guilds, with sedge‐meadow perennials, emergent perennials and floating/submersed aquatics least represented (33‐36%) and annual guilds most represented (74‐94%). Colonization‐to‐extinction ratios suggest that floating/submersed aquatics have already reached a species equilibrium while sedge‐meadow and emergent perennials are still accumulating species. Increasing distance to nearest wetlands decreased the proportion of the regional species pool present in local pools for all guilds except native annuals and woody plants. The maximum proportion predicted, assuming no distance constraint, was comparable to the lowest‐diversity natural wetlands for most perennial guilds, and also lower than what was achieved in a planted, weeded restoration. Conclusions: A biotic constraints seem to limit the colonization of floating/submersed aquatics into natural or restored wetlands, whereas all other guilds are potentially constrained by dispersal or biotic factors (i.e. competition from invasive species). Using species pools to evaluate restoration progress revealed that immigration potential varies considerably among guilds, that local species richness does not necessarily correspond to immigration limitations, and that some guilds (e.g. sedge‐meadow perennials) will likely benefit more than others from being planted at restoration sites.     

Indirect effects of granivory by harvester ants: plant species composition and reproductive increase near ant nests

Summary Of 36 plant species surveyed, 6 were significantly associated with nests of the desert seed-harvester ant Veromessor pergandei or Pogonomyrmex rugosus; two other plant species were significantly absent from ant nests. Seeds of two common desert annuals, Schismus arabicus and Plantago insularis, realize a 15.6 and 6.5 fold increase (respectively) in number of fruits or seeds produced per plant growing in ant nest refuse piles compared to nearby controls. Mass of individual S. arabicus seed produced by plants growing in refuse piles also increased significantly. Schismus arabicus, P. insularis and other plants associated with ant nests do not have seeds with obvious appendages attractive to ants. Dispersal and reproductive increase of such seeds may represent a relatively primitive form of ant-plant dispersal devoid of seed morphological specializations. Alternatively, evolution of specialized seed structures for dispersal may be precluded by the assemblage of North American seed-harvester ants whose workers are significantly larger than those ants normally associated with elaiosome-attached seed dispersal. Large worker size may permit consumption of elaiosome and seed.     

Morphological dispersal structures in relation to growth habit in theCompositae

The relationship between gross morphological dispersal structures, growth habit, and weediness is reported for a worldwide sample of 5893 Compositae species from 18 regional floras. A significantly smaller percentage of annual species than perennial species has dispersal structures. Among species with dispersal structures, plumed types are more frequent in perennials while adhesive types are more frequent in annuals. Differences in the occurrence and nature of dispersal structures between perennial herbs, shrubs and trees were minor. Weeds are less likely to have well developed morphological dispersal structures than non-weeds. The data support the idea that dispersal in space may be more important for perennial or non-weedy plants than for annual and weedy plants.     

Are introduced rats (<Emphasis Type="Italic">Rattus rattus</Emphasis>) both seed predators and dispersers in Hawaii?

Invasive rodents are among the most ubiquitous and problematic species introduced to islands; more than 80% of the world’s island groups have been invaded. Introduced rats (black rat, Rattus rattus; Norway rat, R. norvegicus; Pacific rat, R. exulans) are well known as seed predators but are often overlooked as potential seed dispersers despite their common habit of transporting fruits and seeds prior to consumption. The relative likelihood of seed predation and dispersal by the black rat, which is the most common rat in Hawaiian forest, was tested with field and laboratory experiments. In the field, fruits of eight native and four non-native common woody plant species were arranged individually on the forest floor in four treatments that excluded vertebrates of different sizes. Eleven species had a portion (3–100%) of their fruits removed from vertebrate-accessible treatments, and automated cameras photographed only black rats removing fruit. In the laboratory, black rats were offered fruits of all 12 species to assess consumption and seed fate. Seeds of two species (non-native Clidemia hirta and native Kadua affinis) passed intact through the digestive tracts of rats. Most of the remaining larger-seeded species had their seeds chewed and destroyed, but for several of these, some partly damaged or undamaged seeds survived rat exposure. The combined field and laboratory findings indicate that many interactions between black rats and seeds of native and non-native plants may result in dispersal. Rats are likely to be affecting plant communities through both seed predation and dispersal.     

Internal transport of alien and native plants by geese and ducks: an experimental study

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Host plant predictability and the feeding patterns of monophagous,oligophagous, and polyphagous insect herbivores

Summary Host plant preferences for 34 insect herbivore species are reported. Most polyphagous herbivores feeding on annuals, herbaceous perennials, and woody perennials show distinct preferences for the least abundant plant species among their various host plants. In addition, some populations of widely distributed polyphagous species are much more specialized in their diet than host plant lists alone would suggest. The high level of polyphagy on annuals and herbaceous perennials is suggested to be strongly influenced by the unpredictability of the host plant that is, in turn, controlled by environmental variability. Oligophagous herbivores preferred the least abundant woody perennials on the study sites. Ten of the 22 monophagous herbivores preferred the rarest of all the plant species on the same sites.