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.     

Seed dispersal distances by ant partners reflect preferential recruitment patterns in two ant-dispersed sedges

An important feature of seed dispersal mutualism is the differentiation of dispersal-related seed traits (dispersal syndrome), which potentially contribute to partitioning of both seed dispersers and regeneration sites among sympatric plants. Yet, the selective factors underlying the diversity in dispersal syndromes are largely unknown. The differential requirements for seed dispersal distances are often proposed as a main factor in plant adaptations to disperser animals. Focusing on two sympatric ant-dispersed sedges Carex lanceolata and Carex tristachya (Cyperaceae), we tested the association of the adaptation to different dispersers with requirements for seed dispersal distances. We found that C. lanceolata was more frequently dispersed by the large ant Formica japonica (which had relatively long dispersal distances compared with other smaller ants) than by C. tristachya, and this was caused by the higher seed attractiveness of C. lanceolata to F. japonica. Pot experiments manipulating adult-to-seedling distances showed that isolation from conspecific adults only benefited C. lanceolata seedlings, and C. tristachya seedlings were not affected. These results support the importance of differential requirements for seed dispersal distances as a factor underlying the diversity in dispersal syndromes among animal-dispersed plants.     

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.     

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.     

Effect of cold stratification on seed germination in <Emphasis Type="Italic">Solidago</Emphasis> × <Emphasis Type="Italic">niederederi</Emphasis> (<Emphasis Type="Italic">Asteraceae</Emphasis>) and its parental species

In this study, we investigated the influence of cold stratification on seed germination in S. × niederederi, a hybrid between the North American S. canadensis and the European S. virgaurea, using fruit samples collected in 2016 in Poland. We aimed to test the hypothesis that the low temperature exposure decreases the final percentage and speed of seed germination in the hybrid and its parental species. For each species, sets of 100 achenes in three replications were mixed with dry sand and stored in Petri dishes in darkness for 12 weeks, at ?18 °C and?+?4 °C, and?+?25 °C. The seeds were incubated for 21 d at room temperature (+25 °C), under the 12 h photoperiod (630 lx). We showed a lack of significant differences in: (i) the final percentage of germinated seeds of studied species stored at the same conditions, (ii) the final percentage of germinated seeds between the applied stratification conditions in the hybrid and its parental species, and (iii) the mean values of Timson’s index, mean germination time, and coefficient of velocity of germination between the stratification conditions in each species. The statistically significant inter-specific differences in the mean germination time parameter after the +25 °C treatment suggest that the seeds of S. × niederederi are able to germinate faster than the seeds of its parental species. However, to improve our knowledge of naturalization and invasion abilities of S. × niederederi by sexual reproduction, the seed germination and seedling survival of the hybrid should be tested in the field.     

Environmental cues for germination of the invasive bunch grass <Emphasis Type="Italic">Pennisetum ciliare</Emphasis> (L.) Link

Responses of seed germination to air temperature, water potential, light, and smoke were studied in the laboratory for seeds of the invasive bunch grass Pennisetum ciliare (L.) Link (syn. Cenchrus ciliare L.; buffel grass). First introduced to North America during the mid-twentieth Century for establishing pastures, this African bunch grass has become an invasive species of concern. Across all the experiments conducted, a low germination was observed for P. ciliare fascicles that never exceeded 30 % at 21 days after sowing. Optimal day/night air temperatures for germination, controlled with an environmental chamber, were 25/15 and 30/20 °C, while extreme temperatures of 15/5 and 45/35 °C inhibited germination. By sowing seeds of P. ciliare under different water potentials, created with aqueous solutions of polyethylene glycol, an optimum of ?0.03 MPa led to the highest germination, while no germination was observed at ?1.0 MPa. Monochromatic optical filters were utilized to germinate seeds under various wavelengths, of which red (650 nm) and far red (730 nm) led to the highest germination. In addition, seeds that were incubated in the dark had higher germination than those incubated under white light. Incubation in smoke water, which can stimulate germination of pyrophytic species, resulted in a marginal inhibition of germination compared with imbibition with distilled water.     

Variation in mycorrhizal performance in the epiphytic orchid <Emphasis Type="Italic">Tolumnia variegata in vitro</Emphasis>: the potential for natural selection

Symbiotic seed germination is a critical stage in orchid life histories. Natural selection may act to favor plants that efficiently use mycorrhizal fungi. However, the necessary conditions for natural selection – variation, heritability, and differences in fitness – have not been demonstrated for either orchid or fungus. With the epiphytic orchid Tolumnia variegata as a model system, we ask the following questions: (1) Do seeds from different individuals in a population differ in germination and seedling development in the presence of the same fungi? (2) Do different mycorrhizal fungi (Ceratobasidium spp.) differ in ability to stimulate seed germination and growth in T. variegata? And (3) are the Ceratobasidium isolates that best induce seed germination and seedling development more closely related to each other than to isolates that are less effective? We performed symbiotic seed germination experiments in vitro. The experiments were done using mycorrhizal fungi isolated from T. variegata; relationships among the fungi were inferred from nuclear ribosomal ITS sequences. We found significant variation for both symbiotic germination and seedling growth among biparental seed crops obtained from a population of T. variegata plants. Differences among Ceratobasidium fungi in seed germination were significant. The fungi that induced highest seed germination and seedling development belonged to two of four clades of Ceratobasidium. The two experiments show that there is potential for natural selection to act on orchid–fungus relationships. Given that orchids vary in performance, and that mycorrhizal fungi are not geographically distributed homogeneously, mycorrhizae may affect population size, distribution and evolution of orchids.     

Effects of invasive Green Iguanas (<Emphasis Type="Italic">Iguana iguana</Emphasis>) on seed germination and seed dispersal potential in southeastern Puerto Rico

Green Iguanas (Iguana iguana) are invasive in Puerto Rico due to a variety of negative economic effects, yet we know very little about their ecological impacts. Because they are herbivorous, defecate intact seeds, move through the forest, and have long gut-passage times, Green Iguanas may affect seed germination and seed dispersal. In summer 2013, a total of 258 Green Iguana scat samples were collected at the Humacao Natural Reserve in southeastern Puerto Rico. Seeds extracted from scat and collected from fruit were planted under common garden conditions using experimental treatments designed to tease apart the effects of feces, fruit, and ingestion on seed germination. Green Iguanas decreased the time for seeds to germinate in Ficus spp. by removing fruit pulp, but had no effect on germination of native Annona glabra seeds. For non-native P. pterocarpus and Pterocarpus spp., Green Iguanas produced conflicting results, decreasing the percentage of seeds germinating, but at the same time, reducing the time for seeds to germinate. Green Iguanas likely disperse most seeds beyond the canopies of parental tree at our site. Government and economic resources are being used to eradicate Green Iguana populations in Puerto Rico, but the lack of consistent effects of Green Iguanas on seed germination for the plant species consumed at our site complicates generalizing about their ecological effects and developing management plans that minimize negative effects for native plant communities. We recommend additional studies that target both species of particular concern, such as threatened native or invasive species, as well as studies of sensitive habitats in Puerto Rico.     

Interaction between a threatened endemic plant (<Emphasis Type="Italic">Anchusa crispa</Emphasis>) and the invasive Argentine ant (<Emphasis Type="Italic">Linepithema humile</Emphasis>)

Seed dispersal mutualisms are essential to ensure the survival of diverse plant species and communities worldwide. Here, we investigated whether the invasive Argentine ant can replace native ants by fulfilling their functional role in the seed dispersal of the rare and threatened endemic myrmecochorous plant, Anchusa crispa, in Corsica (France). Our study addressed the potential of Linepithema humile to disperse elaiosome-bearing seeds of A. crispa, examining L. humile’s effects on (1) the composition of communities of ants removing seeds, (2) the number of seed removals, (3) seed preference, (4) the distance of seed dispersion, and (5) seed germination. We caught seven native species at the control site, but only the Argentine ant at invaded sites. L humile removed A. crispa seeds in greater numbers than did native ants, respectively 66 and 23%, probably due to their higher worker density. The invader was similar to native ants with respect to distance of seed transport. Finally, rates of seed germination were not significantly different between seeds previously in contact with either Argentine ants or not. Taken all together, these results suggest that the Argentine ant is unlikely to pose a threat to A. crispa population. These results have important implications for the management of this rare and threatened endemic plant and provide an example of non-negative interactions between invasive and native species.     

Proteomic approach to address low seed germination in <Emphasis Type="Italic">Cyclobalnopsis gilva</Emphasis>

Seeds play essential roles in plant life cycle and germination is a complex process which is associated with different phases of water imbibition. Upon imbibition, seeds begin utilization of storage substances coupled with metabolic activity and biosynthesis of new proteins. Regeneration of organelles and emergence of radicals lead to the establishment of seedlings. All these activities are regulated in coordinated manners. Translation is the requirement of germination of seeds via involvements of several proteins like beta-amylase, starch phosphorylase. Some important proteins involved in seed germination are discussed in this review. In the past decade, several proteomic studies regarding seed germination of various species such as rice, Arabidopsis have been conducted. We face A paucity of proteomic data with respect to woody plants e.g. Fagus, Pheonix etc. With particular reference to Cyclobalnopsis gilva, a woody plant having low seed germination rate, no proteomic studies have been conducted. The review aims to reveal the complex seed germination mechanisms from woody and herbaceous plants that will help in understanding different seed germination phases and the involved proteins in C. gilva.     

Correction to: Neighbourhood stories: role of neighbour identity,spatial location and order of arrival in legume and non-legume initial interactions

Background and aims

We characterized fungal endophytes of seeds of invasive, non-native Phragmites from three sites in the Great Lakes region to determine if fungal symbiosis could contribute to invasiveness through their effects on seed germination and seedling growth.

Methods

Field-collected seeds were surface sterilized and plated on agar to culture endophytes for ITS sequencing. Prevalence of specific endophytes from germinated and non-germinated seeds, and from seedlings, was compared.

Results

One-third of 740 seeds yielded endophyte isolates. Fifteen taxa were identified with Alternaria sp. representing 54% of all isolates followed by Phoma sp. (21%) and Penicillium corylophilum (12%). Overall germination of seeds producing an isolate (36%) was significantly higher than seeds not producing an isolate (20%). Penicillium in particular was strongly associated with increased germination of seeds from one site. Sixty-three isolates and 11 taxa were also obtained from 30 seedlings where Phoma, Penicillium and Alternaria respectively were most prevalent. There was a significant effect of isolating an endophyte from the seed on seedling growth.

Conclusions

These results suggest that many endophyte taxa are transmitted in seeds and can increase seed germination and seedling growth of invasive Phragmites. The role of fungal endophytes in host establishment, growth and invasiveness in nature requires further research.

     

Non-specific symbiotic germination of <Emphasis Type="Italic">Cynorkis purpurea</Emphasis> (Thouars) Kraezl., a habitat-specific terrestrial orchid from the Central Highlands of Madagascar

Orchids, particularly terrestrial taxa, rely mostly on basidiomycete fungi in the Cantharellales and Sebacinales that trigger the process of seed germination and/or initiate the full development of the seedling. During the course of development, orchids may associate with the same fungus, or they may enlist other types of fungi for their developmental needs leading to resilience in a natural setting. This study examined in vitro seed germination and seedling developmental behavior of Cynorkis purpurea, a terrestrial orchid from the Central Highlands of Madagascar. This species is mostly restricted to gallery forests in the Itremo Massif, in moist substrate between rocks bordering streams. The main objective was to understand the influence of diverse mycorrhizal fungi on seed germination and further development of C. purpurea. The study aims to compare symbiotic versus asymbiotic germination and seedling development with seeds and fungi collected from a 13-km² area in the Itremo region. Seeds collected from the wild were sown with diverse orchid mycorrhizal fungi (OMF) spanning 12 operational taxonomic units (OTUs) in three genera (Tulasnella, Ceratobasidium, and Sebacina) acquired from different habitats. Treatments were assessed in terms of the percentage of germinated seeds and fully developed seedlings against those in asymbiotic control media treatments. Overall, OMF significantly improved seedling development within the 12-week experiment period. Sebacina as a genus was the most effective at promoting seedling development of C. purpurea, as well as having the ability to enter into successful symbiotic relationships with orchids of different life forms; this new knowledge may be especially useful for orchid conservation practiced in tropical areas like Madagascar. A Sebacina isolate from an epiphytic seedling of Polystachya concreta was the most effective at inducing rapid seedling development and was among the five that outperformed fungi isolated from roots of C. purpurea. C. purpurea was found to be a mycorrhizal generalist, despite its specific habitat preference, highlighting the complex interaction between the plant, fungi, and the environment. The potential impact on conservation strategies of understanding the requirements for orchid seed germination and development by identifying and using OMF from diverse sources is discussed in detail.     

Fire severity drives variation in post-fire recruitment and residual seed bank size of <Emphasis Type="Italic">Acacia</Emphasis> species

Very high-severity fires are a component of many fire-prone ecosystems, yet are often viewed as detrimental to vegetation. However, species in such systems are likely to have adapted to persist under a fire regime that includes high-severity fires. We examined how fire severity affects post-fire recruitment and residual seed banks of Acacia species and whether severity may affect plant responses to fire intervals. Nine sites of either high or low burn severity were identified after a large-scale mixed-severity fire in Warrumbungle National Park, south-eastern Australia. Transects were used to sample above-ground woody plant density. Seed bank size was surveyed by soil extraction from two depths and manual searching for seeds. Residual soil seed bank and recruitment were compared across the two burn severities. Acacia seedling density was higher in areas burnt at high severity, indicating that increased severity triggers increased germination from the seed bank. Size of residual seed bank was smaller after high-severity fire, but varied between species, with few Acacia cheelii seeds remaining despite high above-ground abundance. In contrast, A. penninervis retained a small residual seed bank. There was little evidence of negative effects on populations of Acacia species after high-severity burns. However, we found that high fire severity may impact on the ability of a species to persist in response to a subsequent short fire interval. Fire management for maintaining biodiversity needs to consider other key aspects of the fire regime, including severity and season, rather than focusing solely on fire frequency.     

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.     

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.     

First evidence of epizoochorous seed dispersal by golden snub-nosed monkeys (<Emphasis Type="Italic">Rhinopithecus roxellana</Emphasis>) in temperate forest

Although many primates are identified as effective endozoochorous seed dispersers in forest ecosystems, epizoochorous seed dispersal by primates has generally been overlooked. In this study, we report epizoochorous seed dispersal in a group of free-ranging golden snub-nosed monkeys (Rhinopithecus roxellana) residing in the temperate forests of central China by identifying seeds carried by 12 well-habituated individuals. The results showed a total of 1920 seeds (five families, seven genera) belonging to eight plant species attached to the monkeys’ fur; the three most abundant species were Geum aleppicum (54% of the total), Torilis japonica (17%), and Agrimonia pilosa (14%). The majority of seeds (95.3%) that attached to the monkeys’ fur had special morphologies such as hairs, hooks, or awns. We also found that the quantity of seeds attached to the fur was not significantly different by age-sex class, even though adult males have very long fur. Our study provides the first empirical evidence of epizoochorous seed dispersal by primates and suggests the role of primates in seed dispersal may be greater than previously assumed.