Role of indehiscent pericarp in formation of soil seed bank in five cold desert Brassicaceae species

The dispersal and germination unit of some Brassicaceae species is the fruit, and we hypothesized that it could affect germination phenology and promote formation of a soil seed bank. We determined the effects of the indehiscent pericarp on germination and longevity of buried seeds of five Brassicaceae species native to cold deserts of central Asia. Germination phenology (seedling emergence) was monitored for intact dispersal units and isolated seeds of Chorispora sibirica, Goldbachia laevigata, Spirorrhynchus sabulosus, Tauscheria lasiocarpa (annuals), and Sterigmostemum fuhaiense (perennial) at natural temperatures in watered and non-watered (natural precipitation) soil. Intact dispersal units and isolated seeds were buried under natural conditions and exhumed at regular intervals for 35 months to monitor germination, viability and moisture content of isolated seeds, seeds in dispersal units, and seeds removed from dispersal units after burial. Isolated seeds of Goldbachia, Spirorrhynchus, and Tauscheria germinated only the first autumn and those of Chorispora and Sterigmostemum the first autumn and first spring, with higher germination percentages in all species in watered than in non-watered soil. A high percentage of seeds in buried dispersal units of Chorispora, Goldbachia, and Sterigmostemum was viable after 35 months, and seeds exhibited a 6-month dormancy cycle, being non-dormant only in autumn and spring. Seeds in buried dispersal units of Spirorrhynchus and Tauscheria germinated when exhumed in the first spring, but all non-germinated seeds were dead after 1 year. Thus, the presence of the pericarp allows Chorispora, Goldbachia, and Sterigmostemum to form a persistent seed bank but not Spirorrhynchus and Tauscheria.     

Composition of harvested seeds and seed selection by the invasive tropical fire ant, <Emphasis Type="Italic">Solenopsis geminata</Emphasis> (Hymenoptera: Formicidae) in Taiwan

Solenopsis geminata (F.) is an invasive ant that is widely distributed in weedy areas and agricultural fields in Taiwan. Previous studies have found that S. geminata harvests the seeds of numerous plants. In the present study, we further investigated the composition of harvested seeds in ant nests and seed selection by workers. The seed caches in S. geminata nests sampled in eight areas in Taiwan suggest that the seeds harvested by workers were diverse and belonged to 52 plant species in 17 plant families. Twenty-three species (44%) belong to the family Gramineae, and most of the seeds weighed from 0.02 to 2.29 mg, which might suggest that these are the main seeds harvested by S. geminata from their habitat. Ten common species with similar seed weights were used to compare the seed preferences of workers from two experimental sites. The results suggest that the seed preference was different between the two experimental sites. The seeds of Casuarina equisetifolia showed the most obvious difference in seed removal speed, which might suggest that S. geminata potentially prefers the encountered seed species in the habitat. The various plant species in the ant nests and seed preference suggest that fire ants easily accept newly encountered plant species. As more than half of the plant species (52%) and the total seed number (63%) belonged to exotic species, the role of S. geminata might be negative because it tends to harvest exotic seeds and has a high opportunity to improve the establishment of exotic seedlings.     

Is induction ability of seed germination of <Emphasis Type="Italic">Phelipanche ramosa</Emphasis> phylogenetically structured among hosts? A case study on Fabaceae species

Phelipanche ramosa is a major root-holoparasitic damaging weed characterized by a broad host range, including numerous Fabaceae species. In France, the agricultural threat posed by P. ramosa has increased over two decades due to the appearance of a genetically differentiated pathovar presenting a clear host specificity for oilseed rape. The new pathovar has led to a massive expansion of P. ramosa in oilseed rape fields. The germination rate of P. ramosa seeds is currently known to vary among P. ramosa pathovars and host species. However, only a few studies have investigated whether phylogenetic relatedness among potential host species is a predictor of the ability of these species to induce the seed germination of parasitic weeds by testing for phylogenetic signal. We focused on a set of 12 Fabaceae species and we assessed the rate of induction of seed germination by these species for two pathovars based on in vitro co-cultivation experiments. All Fabaceae species tested induced the germination of P. ramosa seeds. The germination rate of P. ramosa seeds varied between Fabaceae species and tribes studied, while pathovars appeared non-influential. Considering oilseed rape as a reference species, we also highlighted a significant phylogenetic signal. Phylogenetically related species therefore showed more similar rates of induction of seed germination than species drawn at random from a phylogenetic tree. In in vitro conditions, only Lotus corniculatus induced a significantly higher germination rate than oilseed rape, and could potentially be used as a catch crop after confirmation of these results under field conditions.     

Soil-borne seed pathogens: contributors to the naturalization gauntlet in Pacific Northwest (USA) forest and steppe communities?

Soil-borne seed pathogens are omnipresent but are often overlooked components of a community’s biotic resistance to plant naturalization and invasion. Using multi-year greenhouse experiments, we compared the seed mortality of single invasive, naturalized, and native grass species in sterilized and unsterilized soils collected from Pacific Northwest (USA) steppe and forest communities. Native Pseudoroegneria spicata displayed the greatest seed mortality, naturalized Secale cereale displayed intermediate seed mortality, and invasive Bromus tectorum was least affected by soil pathogens. Seed mortality across all three species was consistently greater in soils collected from steppe than soils collected from forest; seeds sown into sterilized steppe soil experienced half the overall seed mortality compared to seeds sown into unsterilized steppe soil. Soil sterilization did not affect grass seed mortality in forest soils. We conclude that (1) removing soil-borne pathogens with sterilization does increase native and non-native grass seed survival, and (2) soil-borne pathogens may influence whether an introduced species becomes invasive or naturalized within these Pacific Northwest communities as a result of differential seed survival. Soil-borne pathogens in these communities, however, have the greatest negative effect on the survival of native grass seeds, suggesting that the native microbial soil flora more effectively attack seeds of native plants than seeds of non-native species.     

Potential allelopathic effects of the tropical legume <Emphasis Type="Italic">Sesbania virgata</Emphasis> on the alien <Emphasis Type="Italic">Leucaena leucocephala</Emphasis> related to seed carbohydrate metabolism

Allelopathy has been considered a key mechanism to explain the invasiveness of some species. It is well known that invasive plants can affect native plants by producing novel allelochemicals but some exotic plant species may be also sensitive to allelochemicals released by native species, providing a tool to reduce growth and impacts of invasive exotic species. Here, using growth chamber experiments we tested the mutual potential allelopathic effects of Sesbania virgata (a native dominant species) and the alien Leucaena leucocephala seeds. S. virgata was unaffected by seed leachates of L. leucocephala, indicating that, under lab conditions, this legume presents resistance to the phytotoxic compounds produced by seeds of this alien species. In contrast, germination and seedling growth of L. leucocephala were strongly affected by the phytochemicals produced by seeds of S. virgata. A delay in endospermic mobilization of storage carbohydrates (raffinose-family oligosaccharides and galactomannan) was observed in the alien species. These potential allelopathic effects could not be attributed sole to the presence neither of the phytoxic catechin nor of ABA in seed leachates of S. virgata. Our findings indicate that the in vitro behavior of S. virgata is consistent with its aggressiveness in natural environment and suggest sesbanimide as a potential candidate as implicated in the noxious effects of S. virgata on the alien species.     

Diminishing importance of elaiosomes for acacia seed removal in non-native ranges

Myrmecochorous plants produce seeds with lipid-rich appendages (elaiosomes) which act as a reward for seed-dispersing ants. Seed dispersal is important for exotic species, which often need to establish new mutualistic interactions in order to colonize new non-native habitats. However, little is known about the importance of elaiosomes for seed removal in many of their non-native ranges. We studied ant–seed interactions of elaiosome-bearing and elaiosome-removed seeds of the Australian trees Acacia dealbata and Acacia longifolia in order to assess the relative importance of elaiosomes for seed removal between their native (Australia) and non-native (Portugal) ranges. In Portugal, we also studied the co-occurring native plant species with myrmecochorous seeds, Pterospartum tridentatum and Ulex europaeus, across three contiguous levels of acacia invasion: control (i.e. no acacia), low, and high acacia tree density. Acacia seeds were successfully removed by ants in their non-native region by a diversified assemblage of ant species, even in sites where native plants interacted with only one specialized ant species. In the invaded range, diminishing relative importance of elaiosomes was associated with changes in the ant community due to acacia invasion, and for A. dealbata, ant species richness decreased with increasing acacia tree density. Although seed removal was high for both acacia species, the importance of elaiosomes was proportionally lower for A. dealbata in the non-native region. Native plant species experienced significant reductions in seed removal in areas highly invaded by acacia, identifying another mechanism of displacement of native plants by acacias.     

Ant interactions with native and exotic seeds in the Patagonian steppe: Influence of seed traits,disturbance levels and ant assemblage

Invasive plants may establish strong interactions with species in their new range which could limit or enhance their establishment and spread. These interactions depend upon traits of the invader and the recipient community, and may alter interactions among native species. In the Patagonian steppe we studied interactions of native ant assemblages with seeds of native and exotic plants, and asked whether ant–seed interactions differ with seed types and disturbance levels and whether the amount and type of ant–seed interactions can be predicted if both ant and seed traits are known. To characterize and quantify ant–seed interactions, we offered baits with large seeds of Pappostipa speciosa (native) and medium-sized elaiosome-bearing seeds of Carduus thoermeri (exotic), near and far from a road (high vs. low disturbed areas), and compared ant abundance and composition between areas. Interaction frequency was the highest for C. thoermeri seeds far from the road. Composition of ants interacting with C. thoermeri in these areas differed from that near the road and from that interacting with native seeds. Ant composition and abundance were similar between areas, but some species interacted more with exotic seeds in low disturbed areas. Ant foraging type predicted ant–seed interactions since the abundance of seed harvesters was positively correlated to interactions with P. speciosa, and that of generalists and predators, with interactions with C. thoermeri. The high interaction of ants with exotic seeds in low invaded areas suggests that ant activity could influence plant invasion, either by predating or dispersing seeds of invasive plants.     

What we don’t seed: the role of long-lived seed banks as hidden legacies of invasive plants

Acacia dealbata is an invasive south-eastern Australian tree that produces a persistent soil seed bank. In order to characterize the seed bank in plots invaded by A. dealbata and to understand its implications for management and re-invasion risk, density, germinability, and viability of the seed bank were evaluated in five sites in central Portugal. Soil samples were collected in A. dealbata invaded plots and adjacent areas and screened for seeds, which were then quantified and germinated at 25 °C. A subset of seeds was first exposed to 60 °C to assess the effect of high soil temperature on dormancy breaking. Variables influencing differences between sites were explored with generalized linear mixed models with a Poisson distribution. Inside A. dealbata invaded plots the seed bank averaged 4608 seeds/m², reaching up to 62,747 seeds/m²; in adjacent areas, up to 14 m from the plots, only 9 seeds/m² were found. Seed bank densities were mostly influenced by stoniness, number of fires in the last 10 years, and density of trees and roots. Almost 90% of seeds were viable, but only 8.6% germinated without treatment. Nearly 70% of seeds exposed to 60 °C germinated without any physical stimulation, suggesting that high soil temperatures can effectively break seed dormancy. The high density and viability of A. dealbata’s seed bank and its ability to disperse seeds far from the parent plants contribute to the species’ invasive success. These features combined with heat events that can overcome seed dormancy need to be considered in the management of this species.     

Ants Associated with <Emphasis Type="Italic">Turnera subulata</Emphasis> (Turneraceae): Elaiosome Attraction,Seed Dispersion and Germination

Symbiosis between plants and ants include examples in which the plant provides shelter and/or food for ants that, in turn, act in the defense or in the dispersion of seeds from the host plant. Although traditionally referred as mutualistic, the results of these interactions may vary with the ecological context in which patterns are involved. A range of species have facultative association with Turnera subulata (Turneraceae). Here, using behavioral bioassays, we investigated the effects of the most frequent ant species associated with T. subulata (Brachymyrmex sp.1, Camponotus blandus (Smith), Dorymyrmex sp.1, Crematogaster obscurata Emery, and Solenopsis invicta Buren) in the dispersion of plant host seeds and in the number of seedlings around the associated ant nests. We also evaluated the effects of these ant species in the germination of T. subulata seeds, in the consumption of elaiosome, and in the attractiveness to elaiosome odor. Our results showed that the ant species associated with T. subulata presented variation in the attraction by the odor and in the rate of consumption of the elaiosomes. However, none of the ant species studied contributed significantly to the increase of seed germination and seedling growth. Our results suggest that the consumption of the elaiosome by ant species is not a determinant factor to the success of germination of T. subulata. However, such species could contribute indirectly to seed germination by carrying seeds to sites more fertile to germination. In general, our results help to elucidate the results of ecological interactions involving ants and plants.     

Seed Dispersal by Primates and Implications for the Conservation of a Biodiversity Hotspot,the Atlantic Forest of South America

Primates play a fundamental role as seed dispersers, particularly in tropical rainforests. Because defaunation and fragmentation are leading several primate species to local extinction, it is fundamental to understand the role of primates as effective seed dispersers. Here we present a systematic review of studies of seed dispersal by primates in a biodiversity hotspot, the Atlantic Forest of South America, to 1) highlight gaps in our knowledge, 2) determine species richness and proportion of seed species dispersed, and 3) test the relationship between primate body size and the size of dispersed species. Our review found 79 studies of the diet of six ecospecies (Callithrix, Leontopithecus, Callicebus, Sapajus, Alouatta, Brachyteles) but only 20 of these report information on seed dispersal, and none of these are on Callithrix or Callicebus. We found a strong bias in the distribution of species and regions, with most of the studies concentrated in southeastern Brazil. All ecospecies dispersed a large proportion of the seed species they handled (72.1–93.6%). Brachyteles dispersed the highest diversity of plants (N = 73), followed by Sapajus (N = 66), Leontopithecus (N = 49), and Alouatta (N = 26). Although we found no significant relationship between primate body size and the size of seeds dispersed, Brachyteles disperse a higher diversity of large-seeded species than smaller-bodied primates. These results suggest that the local extinction of large primate species may lead to dramatic changes in the plant community, as many large-seeded plants are inaccessible to smaller arboreal frugivores. We propose guidelines for future research on primate seed dispersal to enable the evaluation of seed dispersal effectiveness and to improve our understanding of the fundamental role of primates in this key ecosystem process.     

Host specificity of Asian <Emphasis Type="Italic">Chrysochus</Emphasis> Chevr. in Dej. (Coleoptera,Chrysomelidae: Eumolpinae) and their potential use for biological control of invasive <Emphasis Type="Italic">Vincetoxicum</Emphasis> species

Three Asian leaf beetles of the genus Chrysochus were investigated as potential biological control agents of Vincetoxicum rossicum and V. nigrum, invasive weeds in northeastern North America. Chrysochus chinensis and Ch. globicollis were collected from a field host in a different genus and subtribe. Preliminary no-choice laboratory tests with Ch. goniostoma showed that its physiological host range is too broad. Based on these data, we are not considering these three species as potential biological control agents of invasive Vincetoxicum species.     

The seeds of success: release from fungal attack on seeds may influence the invasiveness of alien <Emphasis Type="Italic">Impatiens</Emphasis>

Although closely related, Impatiens glandulifera and Impatiens balfourii differ in their invasiveness in Europe; only the former is highly invasive there. Following the assumptions of the enemy release hypothesis (ERH), we tested whether these differences may be explained by the levels of seed infestation by pathogenic fungi. Using seeds collected along the Swiss-Italian border, we recorded four true pathogens of seeds: Fusarium culmorum, F. oxysporum, F. sporotrichoides, and Giberella avenacea. In Italy the seeds of I. balfourii were infected by fungal pathogens more often than those of I. glandulifera, while in Switzerland both species were under the same level of pressure. However, the overall differences in pathogen abundance were consistent with the ERH: seeds of the more invasive species were attacked less. This could be a result of differences between the communities of fungal pathogens attacking the seeds of both species in each country. The number of colonies of secondary pathogens (Cladosporium cladosporioides, Alternaria alternata) correlated negatively with the number of colonies of true pathogens; we suggest that the secondary pathogens may have prevented the occurrence of the true pathogens. The reason for the between-country differences in the fungal pathogen communities is unclear. A possible explanation is that Italy and Switzerland differ in their road and green-area maintenance work schemes, which may have influenced pathogen pressure on seeds. This study is one of the few that offers results indicating that release from enemies may be crucial to the invasion success of plants as early as the seed stage.     

Host-specificity of symbiotic mycorrhizal fungi for enhancing seed germination,protocorm formation and seedling development of over-collected medicinal orchid, <Emphasis Type="Italic">Dendrobium devonianum</Emphasis>

All orchids maintain an obligate relationship with mycorrhizal symbionts during seed germination. In most cases, germination-enhancing fungi have been isolated from roots of mature plants for conservation and cultivation purposes. To understand the germination biology of Dendrobium devonianum, an over-collected medicinal orchid, the seeds of D. devonianum were inoculated with a fungal strain (FDd1) isolated from naturally occurring protocorms of D. devonianum and two other germination-enhancing fungal strains (FDaI7 and FCb4) from D. aphyllum and Cymbidium mannii, respectively. The fungal strain was isolated from five protocorms of D. devonianum and identified as a species of the genus Epulorhiza. In germination trials, treatments with all of the three fungal strains showed a significant promoting effect on seed germination and protocorm formation, compared with the control treatment (no inoculation). However, FDd1 fungal strain showed the greatest effectiveness followed by FDaI7 and FCb4. For all inoculation and control treatments, seeds developed to protocorms regardless of the presence of illumination, whereas protocorms did not develop to seedlings unless illumination was provided. The results of our manipulative experiments confirmed the hypothesis that mycorrhizae associated with orchid seedlings are highly host-specific, and the degree of specificity may be life stagespecific under in vitro conditions. The specific mycorrhizal symbionts from protocorms can enhance restoration efforts and the conservation of orchids such as D. devonianum.     

From species to individuals: combining barcoding and microsatellite analyses from non-invasive samples in plant ecology studies

By means of DNA barcoding and microsatellite analyses, we studied the species and individuals of legitimate seed dispersers of the Mediterranean shrub Pistacia lentiscus, a keystone species that represents the main source of food in winter for frugivorous birds. We collected dropping of birds containing seeds, and after DNA extraction we amplified and sequenced a fragment of the mitochondrial COI gene. Through BLASTN queries of the sequenced fragments against registered sequences in the GenBank database we identified the bird species that are currently dispersing P. lentiscus seeds. Further, through the amplification of specific nuclear microsatellite loci we calculated standard genetic diversity parameters of the population of birds from the genus Sylvia (the blackcap and Sardinian warbler), the most important dispersers of P. lentiscus. Five bird species were identified as seed dispersers through their barcode match. Further, we found that S. melanocephala displayed slightly lower levels of genetic diversity than S. atricapilla. In this study we show how the genetic analyses of environmental faecal samples can be a useful and convenient tool for the study of plant-frugivore interactions through the ascertainment of the identity of the species involved and through the analyses of genetic variability of their populations.     

Factors determining species richness of soil seed banks in lowland ancient woodlands

The demise of coppicing in UK ancient woodlands, combined with the planting of non-native, fast-growing conifers in the twentieth century, heightens the potential recharge value of ground flora seed banks. Soil cores from adjoining semi-natural and conifer-containing stands in four lowland ancient woods in central England were removed to establish seed bank species richness. During a fourteen-month germination trial soil from two depths yielded 6554 seedlings from 81 species, ten of which showed a strong affinity for ancient woodland conditions. Juncus effusus accounted for 80% of emergent seeds whilst 23 other species, including Lysimachia nummularia and Potentilla sterilis, were represented by only one individual. Species richness is described by a model that explains 40% of observed variance (P < 0.00001). The model has three significant variables: species richness increases as soil pH rises, and decreases with both depth and increasing time since the most recent planting/disturbance event. No difference was found in the density of seeds from species common to paired semi-natural and conifer-containing stands that were separated only by a woodland ride, suggesting prior management and environmental conditions have a greater influence on seed banks than current stand type. Sørensen similarity index values revealed poor congruence between above-ground vegetation and species in the seed bank. Taking pH measurements in conifer stands identified as younger in terms of planting/disturbance may help locate areas where greater numbers of species (including woodland specialists) are located. Caution is required, however, as these seed banks may also contain non-target, competitive species that may swamp the regeneration of woodland specialists.     

Non-redundancy in seed dispersal and germination by native and introduced frugivorous birds: implications of invasive bird impact on native plant communities

Seed dispersal by vertebrate animals is important for the establishment of many fleshy-fruited plant species. Different frugivorous species can provide different seed dispersal services according to their specific dietary preferences as well as behaviour and body traits (e.g. body size and beak size of birds). Our aim was to study redundancies and complementarities in seed dispersal and germination between the two main native seed disperser birds and the introduced silver pheasant Lophura nycthemera in the temperate Patagonian forests. For this, we collected fresh droppings from the studied species and analyzed seed content. We conducted germination trials for four plant species common in bird droppings; two native species (Aristotelia chilensis and Rhaphithamnus spinosus) and two invasive non-native species (Rubus ulmifolius and Rosa rubiginosa). Both native frugivorous birds and the silver pheasant dispersed fruits of non- native fleshy-fruited plants, but their roles were non-redundant in terms of species dispersed and effect on seed germination. The silver pheasant dispersed a proportionally high number of non-native seeds, while native birds dispersed a high number of native seeds. In addition, the effect of gut treatment in seed germination differed between seed dispersers. Native birds promoted the germination for the two native plant species studied, while the silver pheasant promoted the germination of one non-native plant. This suggests that seed dispersal by the silver pheasant may contribute to the spread of some invasive fleshy-fruited plants in the ecosystems that otherwise would not be dispersed by any other bird. The understanding of redundancies and complementarities on seed dispersal and germination between native and introduced birds will allow improving the management of fleshy-fruited non-native plants.     

Sexual reproduction of clonal dwarf shrubs in a forest–tundra ecotone

Successful sexual reproduction may be more important for regeneration of clonal species in high-latitude and -altitude areas than has been previously suggested. We investigated the potential of Vaccinium myrtillus, V. vitis-idaea and Empetrum nigrum ssp. hermaphroditum (E. hermaphroditum) for sexual reproduction at three sites in the forest–tundra ecotone in Finnish Lapland. We studied whether the potential differs between plant communities, whether disturbance enhances germination, and whether seedling emergence is limited by seed availability. We established a field experiment with disturbance and sowing treatments, and monitored seed and seedling numbers and survival rates for two years. The number of mature seeds of V. myrtillus was higher in plants from the tundra heath than in those from the coniferous and mountain birch forests. The number of mature seeds and seedlings emerging from the seed bank of E. hermaphroditum tended also to be higher in the tundra heath. Disturbance marginally increased the seedling emergence of V. myrtillus and E. hermaphroditum, whereas sowing generally increased the seedling numbers. The seedling number of V. myrtillus was lower in the tundra heath and that of E. hermaphroditum was lower in the coniferous forest than at the other sites. Seedling survival was equal for all plant species at all sites. We conclude that the capacity for sexual reproduction varies among plant communities, and seed availability is a stronger constraint than microsite availability for the studied species. Once the crucial early phase of seedling establishment is overcome, seedling survival enables successful recruitment of V. myrtillus, V. vitis-idaea and E. hermaphroditum in the subarctic area.     

Abiotic barriers limit tree invasion but do not hamper native shrub recruitment in invaded stands

The interplay between the invasion of alien plant species and re-colonization of native plant species is important for conservation. Sandy coastal plains (called restinga in Brazil) were used as a model system to explore the abiotic barriers that potentially limit the initial establishment of alien and native woody plants in invaded and non-invaded areas. The study tested the influence of light availability, soil type and litter layer on recruitment of a wind-dispersed alien tree (Casuarina equisetifolia) and two bird-dispersed native shrubs under a Casuarina stand and in the preserved restinga. The effect of soil type and the physical and allelopathic effects of Casuarina litter on seedling emergence of the three species were also evaluated under greenhouse conditions. Low dispersal associated with low seedling emergence and zero survival of young plants (mainly due to microhabitat conditions) apparently prevents the spread of Casuarina in the preserved restinga. The main cause of low recruitment of native species in the Casuarina stand was the physical barrier of the litter. However, if seeds overcome this physical barrier, the presence of litter improves seedling emergence and survival of young plants, mainly because the litter increases soil moisture. Sowing seeds below litter and planting young plants of native shrubs on litter can improve the re-colonization of native plants in invaded areas. In conclusion, Casuarina invasion on sandy coastal plains is strongly limited by abiotic barriers, but anthropogenic disturbances are altering the key processes that naturally make the restinga resistant to invasion.