Large African herbivores,bruchid beetles and their interactions with Acacia seeds

This study investigated the interactions of large African herbivores and bruchid seed beetles with Acacia seeds. The germination of bruchid-infested and uninfested seeds was compaed. The effects of pod consumption by large herbivores on bruchid infestation and seed germination was also assessed. Bruchid-infested seeds did germinate, and the germination of bruchid-infested and uninfested A. tortilis, A. nilotica and A. hebeclade seeds did not differ. Pod ingestion by large herbivores lowered the bruchid infestation of consumed and defaecated seeds compared to uningested seeds. Uninfested, ingested and voided A. tortilis seeds germinated seeds. Furthermore, infested A. tortilis seeds egested by giraffe, kudu and ostrich germinated better than infested, uningested seeds. Pod ingestion by large herbivores may reduce bruchid infestation, increase Acacia seed germination and therefore increase potential Acacia seedling recruitment.     

Seed protection through dispersal by African savannah elephants (Loxodonta africana africana) in northern Tanzania.

We examined effectiveness of African savannah elephant dung as a protective barrier for seeds of three tree species, Acacia tortilis Hayne, Tamarindus indica L. and Ximenia aegyptiaca L. Seeds were collected from dung and underneath fruiting trees in Tarangire National Park, Tanzania. Experimental treatments were established to test: (i) the efficacy of dung in protecting seeds of A. tortilis from bruchid beetle infestation and the role of animals larger than insects in removing seeds; (ii) the same tenets as in Experiment 1, using seeds of T. indica; and (iii) the effect of distance on survival of seeds of X. aegyptiaca. Sites were established during two field seasons underneath conspecific trees, where seed predation was likely highest. Repeated‐measures two‐way ANOVA indicated that there was no treatment effect for Experiment 1. For Experiments 2 and 3 in October 2013, seeds in dung experienced less beetle infestation than fresh seeds. Repeated‐measures two‐way ANOVA and Tukey's HSD indicated that treatment effect differed among all treatments. Passed seeds at distances ≥5 m experienced less beetle infestation than fresh seeds underneath conspecifics. African savannah elephants appear to be important seed dispersers of these three tree species.     

Effects of storage,mucilage presence,photoperiod, thermoperiod and salinity on germination of Farsetia aegyptia Turra (Brassicaceae) seeds: implications for restoration and seed banks in Arabian Desert

Seed viability and germination are key factors in the success of restoration efforts, especially when stored seeds are used. However, the effect of seed storage on germination of most of the native Arabian species is not well documented. We investigated the effect of storage time and role of the seed mucilage in regulating germination, dormancy, salinity tolerance and consequential survival strategy of F. aegyptia in an unpredictable arid desert setting. Effect of light and temperature during germination was studied under two photoperiods and two thermoperiods using intact and de-mucilaged seeds. Presence of mucilage and thermoperiod did not affect the germination. However, seed collection year and photoperiod had a highly significant effect on the germination. Increasing salinity levels decreased the germination of F. aegyptia but ungerminated seeds were able to germinate when salinity stress was alleviated. Seed storage at room temperature enhances the germination percentage, indicating that F. aegyptia seeds have physiological dormancy and it can be alleviated by after-ripening at dry storage. In addition, F. aegyptia seeds show ability to germinate at lower salinity concentration and remain viable even at higher saline conditions, indicating their adaptability to cope with such harsh environmental conditions.     

Bet‐hedging and germination in the Australian arid zone shrub Acacia ligulata

The diaspore of the Australian arid zone shrub Acacia ligulata is dispersed by birds and ants. To investigate the benefits of providing a dispersal structure attractive to both groups, we compared the germination response and viability of seeds eaten by birds, handled by ants or collected from trees to simulated precursors of germination: scarification, fire and rainfall were simulated. Seed germination and viability were related to the degree of preheating disturbance to the seed coat. Heating increased the germinability of seeds not scarified or eaten by birds. In the absence of heating, ingestion by birds increased germinability. Heating increased the mortality of seeds. Our results suggest that ingestion of seeds by birds may break seed dormancy and hence enable some seeds to germinate soon after dispersal. Alternatively, seeds not eaten by birds are likely to remain dormant until sufficiently scarified by soil or stimulated by fire. Consequently, in areas such as the Simpson Desert, A. ligulata may be able to use a range of seedling establishment ‘windows’ provided by monsoon rains, post‐fire environments and unseasonal winter rains, and also spread the risk of unsuccessful seedling establishment by retaining dormant seeds in the seedbank.     

Effects of ingestion of seeds by sika deer (<Emphasis Type="Italic">Cervus nippon</Emphasis>) and dung presence on their germination in a herbaceous community

In a herbaceous community subjected to continual impacts of sika deer (Cervus nippon), I examined the effects of seed ingestion by deer on seeds by comparing the ripening and germination rates of seeds of two dominant species, Zoysia japonica and Hydrocotyle maritima, between seeds taken out of fecal pellets (deer-ingested seeds) and mature seeds collected directly from living plants (control seeds). Seeds of Z. japonica were likely to have tolerance to ingestion from earlier periods of seed maturity. In contrast, only ripened seeds of H. maritima may have tolerance to ingestion. When the seeds ripened, the germination rates of two species did not differ significantly between deer-ingested seeds and control seeds. Thus, although immature seeds may be crushed by ingestion, many mature seeds can be dispersed by sika deer with no alteration of germination rate. However, the other germination experiment showed that the germination rates were significantly higher for seeds of Juncus tenuis in artificially broken fecal pellets than for those kept confined in the pellets, and all seeds germinated from intact pellets were situated near the surface of the pellets. These results suggest that dung may physically prevent seeds inside from germinating and decomposition of dung enables herbaceous small seeds in the dung to germinate.     

Interaction between ungulates and bruchid beetles and its effect on <Emphasis Type="Italic">Acacia</Emphasis> trees: modeling the costs and benefits of seed dispersal to plant demography

Integrative studies of plant–animal interactions that incorporate the multiple effects of interactions are important for discerning the importance of each factor within the population dynamics of a plant species. The low regeneration capacity of many Acacia species in arid savannas is a consequence of a combination of reduction in seed dispersal and high seed predation. Here we studied how ungulates (acting as both seed dispersers and herbivores) and bruchid beetles (post-dispersal seed predators) modulate the population dynamics of A. raddiana, a keystone species in the Middle East. We developed two simulation models of plant demography: the first included seed ingestion by ungulates and seed predation by bruchids, whereas the second model additionally incorporated herbivory by ungulates. We also included the interacting effects of seed removal and body mass, because larger ungulates destroy proportionally fewer seeds and enhance seed germination. Simulations showed that the negative effect of seed predation on acacia population size was compensated for by the positive effect of seed ingestion at 50 and 30% seed removal under scenarios with and without herbivory, respectively. Smaller ungulates (e.g., <35 kg) must necessarily remove tenfold more seeds than larger ungulates (e.g., >250 kg) to compensate for the negative effect of seed predation. Seedling proportion increased with seed removal in the model with herbivory. Managing and restoring acacia seed dispersers is key to conserving acacia populations, because low-to-medium seed removal could quickly restore their regeneration capacity.     

Germination variability and the effect of various pre-treatment on germination in the perennial spurge Euphorbia nicaeensis All.

Patterns of seed germination of the perennial spurge Euphorbia nicaeensis were studied in three populations in south-western Spain. We investigated the variation in seed viability and germination among individuals, and among populations over 2 consecutive years. We also studied if diverse factors such as temperature, acid scarification, darkness or caruncle loss affect the germination of the seeds.

Interindividual and interpopulation variability in seed viability was found. E. nicaeensis seeds are nondormant, so differences in interpopulation viability translate into differences in final germination rates. The germination percentage of the seeds from each population was similar in the 2 years studied. The effects of diverse factors were homogeneous in the two populations studied. Darkness has no effect on seed germination, and ecarunculate seeds germinated in the same proportions as carunculate seeds; this could allow the seeds to germinate in the chambers of deserted anthills. Acid scarification significantly reduced the germination percentage in only one of the populations, but over 50% of the seeds germinated, which could allow herbivores to act as occasional dispersing agents. The seeds that were preheated at 100 °C for 1 and 5 min germinated in the same proportions as the control group. The seeds that were preheated at 120 °C for 5 min displayed a significant decrease in germination, but the percentage was over 40% for both populations, indicating that the seeds could still germinate after the passage of a fire.     

Effects of bird ingestion on seed germination of Sorbus commixta

To determine the effects of ingestion by birds on seed germination, we performed germination experiments in the field and laboratory with Sorbus commixta. The germination of four groups of seeds was compared: ingested seeds, seeds defecated in feces after feeding of fruits to birds; extracted seeds, seeds deliberately extracted from the fruit pulp; juiced seeds, seeds plus the juice of the pulp after seeds had been deliberately extracted from the pulp; intact seeds, seeds in untreated intact fruits. In the laboratory, intact and juiced seeds hardly germinated, but ingested and extracted seeds germinated. Thus, the pulp and its juice appeared to inhibit germination, but seeds could germinate without ingestion by birds once the seeds had been manually extracted from the pulp. In the field, intact fruits did not germinate in the first spring, because the seed was still covered with pulp. The pulp of intact seeds decomposed during the first summer, and thus, the seeds had the potential to germinate during the second spring. In fact, most intact seeds do not germinate during the second spring either, since they lose their viability during the first summer. Thus, under natural conditions, most seeds of Sorbus commixta cannot germinate without bird ingestion. Received: 5 July 1997 / Accepted: 7 November 1997     

Establishment of pasture species on cattle dung: the role of endozoochorous seeds

Abstract. The process of colonization of cattle dung patches and its relation to endozoochorous dispersal is analysed in a Mediterranean pasture. Dung pats dropped in spring and winter were marked, and the vegetation developing on them during the first two years of colonization was followed. The influence of endozoochorous seeds on dung colonization is assessed through the study of: (1) the vegetation on spring and winter dung pats and the seed contents of both, which is abundant in the former and negligible in the latter; and (2) the vegetation on spring dung pats compared with that of the surrounding pasture. The origin of plants growing on dung (either seeds in the manure or in the soil seed bank) was traced by studying dung colonization under controlled conditions in pots with manure put on seed-free sand, and on pasture turf. The results indicate that endozoochorous seeds germinate in manure; they are the main source of recovery in gaps generated by dung pats. The micro-succession involved is independent of the type of pasture. A small-scale spatial pattern results in which gaps of old dung are dominated by endozoochorous species. Thus, dung patches enhance the similarity between grazed communities, and the variation within communities.     

Imbibition phases and germination response of Mimosa bimucronata (Fabaceae: Mimosoideae) to water submersion

Mimosa bimucronata is a pioneering tree that occurs predominantly in moist lowlands, floodplains and on margins of rivers and lakes in Latin America. The effect of submergence on seed germination in M. bimucronata was firstly studied. Patterns of water absorption by M. bimucronata seeds were investigated thereafter to assess the imbibition phases of scarified and unscarified seeds. The germination percentage was significantly higher in scarified than in unscarified seeds, and the velocity of seed germination also increased considerably in scarified seeds. Submergence duration did not significantly affect germination percentages of scarified and unscarified seeds. Therefore, seed viability after submersion suggests that M. bimucronata may display hydrochorous dispersal and also that seeds are able to germinate successfully in areas with frequent seasonal flooding. With respect to imbibition phases, phase II was very short or even absent for scarified and unscarified seeds; therefore, a plateau, where water absorption by seeds is established, was not observed. Finally, we verified that the passage from phase I to III was very tenuous and took a long time in seeds without scarification.     

不同畜禽粪便浸出液对青钱柳种子发芽特性的影响研究

本文选取猪粪、牛粪、羊粪、鸡粪等4种畜禽粪便堆肥为供试样品,分别采用不同浓度的浸提液,对青钱柳种子进行育苗,统计其发芽率、根数、叶片数并测量根长。结果表明,猪粪、羊粪、鸡粪粪便浸提液在1∶40浸提比浓度处理下对青钱柳种子的相对发芽率、根数、叶片数有明显的促进作用,鸡粪浸提液的促进作用最明显,且鸡粪在1∶50和1∶60浸提比浓度下处理对相对发芽率也有一定的促进作用,因此鸡粪能显著提高青钱柳种子的相对发芽率、根数、叶片数。而猪粪、羊粪、鸡粪粪便浸提液1∶20浸提比浓度对青钱柳相对根长具有显著的促进作用,其余浸提比对青钱柳相对根长则有一定的抑制作用,且随着浓度的下降其抑制作用越显著。因此,用鸡粪粪便浸提液在1∶40浸提比浓度处理下对青钱柳种子育苗是有积极作用的,为今后对青钱柳种子育苗提供参考。     

Propagation of Rare Plants from Historic Seed Collections: Implications for Species Restoration and Herbarium Management

Herbaria are potentially important repositories of living seeds that could be useful for recovery of rare plant species. To examine this capacity, we tested seed germination of rare milkweed (Asclepias) and milkvetch (Astragalus) species representing different collection dates and different herbaria. These groups have contrasting seed characteristics, with greater potential for longevity in the nonpermeable hard-coated milkvetch seeds. Twelve-year-old Asclepias lanuginosa seeds failed to germinate. However, we achieved 45% germination from three-year-old Asclepias meadii seeds, but germination dropped to 0% after ages of four to five years. Astragalus neglectus seeds germinated from 97-, 48-, and 28-year-old herbarium specimens, and Astragalus tennesseensis seeds germinated from a four-year-old collection. Seedlings produced from these experiments were incorporated into ex situ garden populations for recovery or restoration of rare species populations. Different herbarium pest control techniques may have significant bearing on the viability of seeds stored on herbarium specimens. Microwaving can cause precipitous loss of seed viability, while deep-freezing appears to allow some seeds to remain viable. Potentially live seeds of rare species should be stored under conditions that enhance their long-term viability.     

黄土丘陵沟壑区主要物种植冠种子库动态及其生态策略

植冠种子库是植物适应环境并应对外界干扰的种子生态策略之一,研究了黄土丘陵沟壑区12种主要植物植冠种子库动态,结果表明:杠柳不具有植冠种子库,其他11种植物均具有植冠种子库;除了黄刺玫种子在翌年5月达到脱落高峰,其他植物大部分种子在冬季脱落,其中杠柳、达乌里胡枝子、茭蒿、黄柏刺和水栒子的大部分种子脱落集中偏早,铁杆蒿和土庄绣线菊的大部分种子脱落集中偏晚;植冠宿存对大部分植物种子的萌发特性表现为促进作用;但不同植物种子的萌发时滞对植冠宿存响应差异较大;9种植物种子在植冠上宿存至翌年2月底,其种子活力仍能维持达60%以上;该区植物表现出不同的植冠种子库策略,通过不同的方式来减少干扰的威胁,提高成功萌发与更新的几率,它们或具有较大规模的宿存量、或调控种子萌发特性、或提高种子维持活力的百分比。此外,全面了解该区植物形成植冠种子库的机理及对应的生态策略还有待于全面、深入的研究。     

Efficiency of endozoochorous seed dispersal in six dry-fruited species (Cistaceae): from seed ingestion to early seedling establishment

We combined laboratory and nursery experiments to analyse the effectiveness of sheep as endozoochorous seed dispersers of six native shrubby Cistaceae species collected in SE Spain (Helianthemum apenninum (L.) Mill., H. violaceum (Cav.) Pers., Fumana ericoides (Cav.) Grand., F. thymifolia (L.) Spach, Cistus monspeliensis L. and C. laurifolius L.), considering the main stages after seed ingestion, i.e. seed recovery, seed germination, seedling emergence and early seedling establishment. Seed recovery after gut passage was high (around 40%) for all the species, except F. thymifolia (12%). Most seeds (ca. 90%) were recovered within 48 h after ingestion for all the species, although seeds were still recovered up to 96 h after ingestion. Gut passage increased germination up to seven-fold compared to non-ingested seeds. Furthermore, seedling emergence from seeds contained in pellets was overall similar (intact pellets) to or higher (crumbled pellets) than emergence from seeds without dung. Survival of emerged seedlings and mass of seedlings after 20 days were not reduced by dung. Sheep act therefore as effective dispersers of these Cistaceae species by scattering seeds and promoting germination, while faeces do not hamper seedling establishment. We conclude that the interaction between herbivorous ungulates and these dry-fruited species may be considered a mutualism qualitatively similar to the mutualism between frugivorous vertebrates and fleshy-fruited plants.     

Effect of mango weevil (Coleoptera: Curculionidae) damage on mango seed viability in Hawaii

The mango weevil, Cryptorhynchus (= Sternochetus) mangiferae (F.), is a federally quarantined pest that prevents shipment of mangos from Hawaii into the continental United States. Although this monophagous weevil allegedly causes reduced seed germination, damage to the fruit pulp, and premature fruit drop in mangos, there are few studies examining these potential sources of crop loss. We conducted studies to assess the effect of mango weevil infestation on seed viability while making observations on the frequency of pulp feeding. Naturally infested seeds from mature fruit were planted in pots and scored for successful germination. Germination rates for infested seeds were equal to those of uninfested control seeds in a polyembryonic cultivar ('Common'), whereas germination was significantly reduced for infested seeds of a monoembryonic cultivar ('Haden') compared with uninfested control seeds but germination of infested seeds was still > 70%. To assess seed tolerance of damage, seeds were artificially damaged by cutting away 25, 50, or 75% of the cotyledon before planting and scored for germination. None of the damage treatments was significantly different from the undamaged controls, indicating that mango seeds can withstand substantial damage and still germinate successfully. Over the 2-yr period we conducted experiments, only four of 3,602 mango fruits (0.11%) showed evidence of direct feeding damage to the pulp. Results suggest that C. mangiferae is a less serious pest of mangos than previously thought.     

THE NINETY-YEAR PERIOD FOR DR. BEAL'S SEED VIABILITY EXPERIMENT

In 1879 Dr. W. J. Beal selected seeds of 23 different species of locally common plants, mixed 50 seeds of each species with moist sand in unstoppered one-pint bottles, and buried the bottles in a sandy knoll to be unearthed and the viability of the seeds tested periodically. The year 1970 marked the ninetieth year the seed had been buried, and the thirteenth bottle was recovered to test for seed viability. Of the three species which had germinated in the 1960 test (curly dock, Rumex crispus; evening primrose, Oenothera biennis; and moth mullein, Verbascum blattaria), only V. blattaria had viable seed with 20% germination. No other species germinated. All ten seedlings of V. blattaria were grown to maturity, and seeds were then harvested to study the possible deviations from normality and the requirements for seed germination. All seedlings emerging from the first progeny seed appeared normal. The most prominent requirement for germination was light, and this is a possible explanation of why the seeds had remained viable but dormant for so long a period. One-third of the freshly harvested seed germinated in darkness and, furthermore, redrying of dark-moistened seed in the absence of light induced additional germination. Germination of dark-moistened seed was not completely restored when the still moist seeds were subsequently exposed to light. However, when dark-moistened seeds were dried and then remoistened in the light, germination was about 95 %. About 5 % of the seed did not germinate under the conditions used. We find that 5 % of the population of V. blattaria seeds are dormant for unknown reasons, that 30 % will germinate if supplied only with moisture, and that 65 % are inhibited and require light and moisture simultaneously for germination. Supplying this 65 % of the population with moisture in darkness results in the development of a second type of inhibition which is no longer light reversible. It appears that the simultaneous requirement for light and moisture is an important factor in permitting V. blattaria seeds to remain dormant during prolonged burial.     

Seed germination traits of desert perennials

While understanding that seed germination is crucial for ecological restoration activities, the seed traits of desert perennials are understudied. We experimentally determined germination traits of 43 species from 14 families from Hummock grasslands in the Great Sandy Desert, Australia. We defined morphological and physiological seed traits of framework species required for restoration and investigated the effects of fire and temperature on seed germination. We classified dormancy and explored the effect of Karrikinolide, a fire cue derived from smoke, on germination. Seeds of 38 (88%) out of 43 species were dormant: 13 (30%) with physical and 25 (58%) with physiological dormancy. Karrikinolide promoted seed germination of 9 (21%) species across all life-forms except trees, and widened the range of germination temperatures and increased germination rate of one species. Although high germination percentages were obtained over a wide temperature range, germination rate was affected by temperature. Non-dormant seeds and seeds pre-treated to overcome physical dormancy germinated quickly, with times to 50% germination of 1–5 days. Dormancy class differed between life-forms and families. Fast germination of non-dormant seeds is a trait that allows seeds to germinate during short periods of moisture availability. An absence of under-developed embryos is consistent with the global trends for hot deserts. A response to Karrikinolide shows that seed germination is related to a fire cue. These results will inform land managers of effective seed pre-treatments prior to seed broadcasting for restoration, and information on seed germination temperatures and rates will improve the understanding of when and where seeds could germinate in restored sites.     

Germination behaviour of annual plants under changing climatic conditions: separating local and regional environmental effects

The role of local adaptation and factors other than climate in determining extinction probabilities of species under climate change has not been yet explicitly studied. Here we performed a field experiment with annual plants growing along a steep climatic gradient in Israel to isolate climatic effects for local trait expression. The focus trait was seed dormancy, for which many theoretical predictions exist regarding climate-driven optimal germination behaviour. We evaluated how germination is consistent with theory, indicating local adaptation to current and changing climatic conditions, and how it varies among species and between natural and standardised soil conditions. We reciprocally sowed seeds from three or four origins for each of three annual species, Biscutella didyma, Bromus fasciculatus and Hymenocarpos circinnatus, in their home and neighbouring sowing locations along an aridity gradient. Our predictions were: lower germination fraction for seeds from more arid origins, and higher germination at wetter sowing locations for all seed origins. By sowing seeds in both local and standard soil, we separated climatic effects from local conditions. At the arid sowing location, two species supported the prediction of low germination of drier seed origins, but differences between seed origins at the other sites were not substantial. There were no clear rainfall effects on germination. Germination fractions were consistently lower on local soil than on standard soil, indicating the important role of soil type and neighbour conditions for trait expression. Local environmental conditions may override effects of climate and so should be carefully addressed in future studies testing for the potential of species to adapt or plastically respond to climate change.     

Reduced germination success of temperate grassland seeds sown in dung: consequences for post‐dispersal seed fate

• Endozoochory is one of the main drivers shaping temperate grassland communities by maintaining plant populations of its constituents and enabling plants to colonize new habitats. Successful endozoochorous dispersal implies that seeds not only get consumed and survive the digestive tract but are also able to develop into viable seedlings in a dung environment.
• We experimentally assessed the germination probability and timing of 15 annual and perennial temperate European grassland species in cattle and horse dung and in different climatic conditions (greenhouse and outdoor conditions).
• Interspecific variation in germinability and germination timing are found, while life strategy had only an effect on germination timing. We found adverse effects of both cattle and horse dung on the germination characteristics of all tested grassland species, but the effects of cattle dung were more pronounced. In comparison with the control treatment, fewer seeds emerged in dung and more time was needed to germinate. Also, germination metrics clearly differed between the artificial greenhouse and outdoor conditions, with generally a lower germinability in outdoor conditions.
• According to our results, a large cost seems to be associated with endozoochorous dispersal in this stage of the life cycle, as seed dispersal effectiveness strongly depends on the quality of the deposition site with a lowered survival and germination probability when seeds are deposited in dung.

     

Seed hydration memory in Sonoran Desert cacti and its ecological implication

Cactus seeds on the soil surface in the desert are subjected to periods of drought that last for up to a few months, and thus they are typically under discontinuous hydration (or discontinuous dehydration). Apparently, they can tolerate long periods of dehydration after single or multiple hydration events and subsequently germinate in accordance with the previous hydration experience. This was verified in three cactus species from the Sonoran Desert. Seeds of Stenocereus thurberi hydrated for 72 or 80 h followed by a dehydration period lasting for 4, 14, 70, 120 or 181 d germinated 2–3 d earlier and had 1.4–2 times shorter mean germination time (MGT) than untreated seeds. Seeds given shorter hydration periods also began to germinate sooner than the controls. MGT was shorter only when the hydration period was 48 h or longer. Final germination percentages were not affected by these treatments, only the MGT. Except for differences in germination percentages, similar results were found for Pachycereus pecten-aboriginum and Ferocactus peninsulae. When the cycle of 24 h hydration followed by 4 d dehydration was repeated one or two times, the effect was cumulative: MGT was equal to 48 and 72 h hydration, respectively. These results suggest a phenomenon of “seed hydration memory,” the ability of seeds to retain during dehydration periods those physiological changes that result from seed hydration. Thus, treated seeds subsequently germinated earlier then untreated seeds, regardless of the duration of dehydration period. This led to a greater biomass accumulation and thus to higher survival in seedlings from treated than from untreated seeds.