Dung beetle activity affects rain forest seed bank dynamics and seedling establishment

Dung beetles relocate vertebrate feces under the soil surface, and this behavior has many ecological consequences. In tropical forests, for example, seeds defecated by mammals that are subsequently buried by dung beetles are less likely to suffer predation. While the effects of dung beetles on the fate of defecated seeds have been relatively well studied, their effect on seeds already buried in the soil has not. To contribute to fill this gap, we designed a study with three objectives: (a) Describe the vertical re‐distribution of soil seeds that occurs due to dung beetle activity; (b) Determine if beetle activity favors establishment of seedlings from the soil seed bank; and (c) Determine if the effect of dung beetles is stronger in sites of recurrent mammal defecation. We carried out three complementary field experiments, one with artificial seeds (plastic beads) of three sizes buried at known depths, one with two species of seeds buried at those same depths, and one with the natural soil seed bank in sites of single vs. recurrent defecation. Buried beads were moved by dung beetles along the vertical axis, both upwards (9.5%) and downwards (11.5%); smaller beads were more frequently moved downwards while the contrary occurred for larger beads. Dung beetle activity caused an increase in seedling establishment, both from experimentally buried seeds and from the natural seed bank. Defecation recurrence had no effect on seedling establishment. We conclude that dung beetle activity affects seed bank dynamics with important consequences for seedling establishment in tropical forests.Abstract in Spanish is available with online material.     

Litter effects on seedling establishment interact with seed position and earthworm activity

Seedling establishment is influenced by litter cover and by seed predators, but little is known about interactions between these two factors. We tested their effects on emergence of five typical grassland species in a microcosm experiment. We manipulated the amounts of grass litter, seed sowing position and earthworm activity to determine whether: (i) the protective effect of litter against seed predation depends on cover amount and seed sowing position, i.e., on top or beneath litter; (ii) seed transport by earthworms changes the effect of seed sowing position on seedling emergence; and (iii) seeds transported into deeper soil layers by earthworms are still germinable. Litter cover and presence of earthworms lowered seedling emergence. The impact of seed position increased with seed size. Emergence of large-seeded species was reduced when sown on the surface. Additionally, we found an important seed position × earthworm interaction related to seed size. Emergence of large-seeded species sown on top of the litter was up to three times higher when earthworms were present than without earthworms. Earthworms also significantly altered the depth distribution of seeds in the soil and across treatments: on average 6% of seeds germinated after burial. In contrast to the seed position effect, we found no size effect on mobility and germinability of seeds after burial in the soil. Nevertheless, the fate of different-sized seeds may differ. While burial will remove large seeds from the regeneration pool, it may enhance seed bank build up in small-seeded species. Consequently, changes in the amount of litter cover and the invertebrate community play a significant role in plant community composition.     

Effects of microsite conditions on seedling establishment on the foreland of Coleman Glacier,Washington

Abstract. Questions: How do physical microsite conditions of microsites affect germination and seedling survival in different successional stages? Do different species germinate in similar microsites in a given successional stage? Location: Coleman Glacier foreland, Mount Baker, Washington State, USA. Methods: Two methods were used to characterize safe sites. 1. Grids of 300 10 cm × 10 cm plots were located in four different age classes on the foreland. 2.105 pairs of plots, with and without seedlings of Abies amabilis, were located in each age class. For each plot we identified all seedlings and all individuals < 1 m tall. Microsite characteristics such as topography and presence of rocks or woody debris were noted for each plot. Microsite characteristics were compared between plots with and without each species. In addition we examined the effect of distance from seed sources on the presence of Alnus viridis seeds and seedlings in a newly disturbed area. Results: In early successional sites, seedlings of several species were positively associated with depressions and presence of rocks, and negatively associated with ridges. Patterns were generally consistent among species. In later succession, seedlings were not significantly associated with any microsite characteristics. For Alnus viridis, seed density decreased with distance from seed sources but seedling density did not. Conclusions: Because of harsh conditions in early succession, physical microsites are important, and most species have similar microsite requirements. In later succession, physical microsites characteristics are not as important and are more variable. Microsites appear to be more important than seed rain in controlling the distribution of Alnus viridis in early succession.     

Mating structure and inbreeding and outbreeding depression in the rare plant Gentianella germanica (Gentianaceae)

Isolation and small size of populations as a result of habitat destruction and fragmentation may negatively affect plant fitness through pollinator limitation and increased levels of inbreeding. To increase genetic variation in small populations of rare plants artificial gene flow has been suggested as a management tool. We investigated whether pollinator limitation and inbreeding depression could reduce fitness in Gentianella germanica, an endangered biennial of increasingly fragmented calcareous grasslands in Central Europe. We experimentally excluded pollinators and generated progenies by hand-pollinating flowers with pollen from different distances. G. germanica was highly selfing. Pollinator exclusion strongly reduced seed set, indicating that pollinator limitation could potentially reduce plant fitness. Germination rate as well as number of leaves and rosette size of progeny from 10-m crosses was higher than that of progeny from open pollinations, self-, 1-m, and interpopulation crosses. After 6 mo of growth differences in the number of surviving plants persisted, whereas differences in plant size did not. The results suggest that inbreeding depression may reduce plant performance in G. germanica. Outbreeding depression in the performance of progeny from interpopulation crosses indicates that caution is necessary in using artificial interpopulation gene flow as a management tool.     

Sand burial effects on seed germination, seedling emergence and establishment of Panicum virgatum

Studies in the field and in a greenhouse were conducted to examine the effects of sand burial on seed germination, seedling emergence and establishment of Panicum virgatum L. on the foredunes of Lake Erie. Under natural conditions, the seedlings emerged from sand burial depths ranging from 0 to 11 cm, with a mean ± SD of 4.73 ± 1.82 cm. The frequency distribution of the depth of emergence of seedlings in the field was significantly skewed to the right and platykurtic. In the greenhouse, some seedlings emerged from a burial depth of as much as 16 cm. Although percent germination of seeds was not affected by sand burial, the percent emergence and the rate of emergence of seedlings were significantly reduced by excessive sand burial. Seedling mortality was found only among seedlings that emerged from sand burial depths of 10 cm or more. In the field, all the seedlings established in one growing season had originally emerged from sand burial depths of less than 12 cm. Within this burial range, seedlings from shallower burial depths had lower chances of establishment than expected, whereas those from deeper burial depths had higher probabilities of establishment than expected.     

围封沙质草甸土壤种子库与幼苗库的特征

采用野外定点取样和萌发实验相结合的方法,研究了科尔沁沙地围封沙质草甸土壤种子库和幼苗库的特征.结果表明:土壤种子库总密度为13854±1651有效种子数/m2,而幼苗库的总密度为808±164株/m2,仅为土壤种子库总密度的6%,土壤种子库与幼苗库总密度之间存在显著的差异 F=36.0,n=219,P<0.001 .土壤种子库与幼苗库的植物种数分别为37和44,其中一、二年生草本植物分别占种子库与幼苗库植物种数的62%和50%,而多年生草本植物所占的比例分别为24%和41%,灌木所占的比例很小,分别为14%和9%.土壤种子库与幼苗库种类组成上的相似性系数为0.667,表现出较小的异质性.土壤种子库密度与幼苗库密度之间存在显著的相关性,其间关系可用三次曲线来描述.     

Effects of grazing on seedling establishment: the role of seed and safe-site availability

Abstract. The first objective of this paper was to assess the effects of grazing on seedling establishment of two species whose relative abundance at the adult stage is affected by grazing in a contrasting fashion. Second, we evaluated the relative importance of seed versus safe-site availability in explaining the effect of grazing on seedling establishment. We monitored seedling establishment on a grazed area, on two areas which had not been grazed for two and seven years, and on plots which had been experimentally defoliated. The species compared were Dan-thonia montevidensis, a native perennial grass which dominates both grazed and ungrazed communities, and Leontodón taraxacoides, an invading exotic rosette species from the Compositae family. Continuous grazing enhanced seedling establishment of both species through its effect on the availability of safe sites. Seed availability accounted for only one, but very important, grazing effect: the lack of response by L. taraxacoides to the defoliation in the seven-year old exclosure. Its seed supply was depleted by exclusion of grazing and, consequently, its short-term regeneration capacity after disturbance was lost.     

RAPD variation in relation to population size and plant fitness in the rare Gentianella germanica (Gentianaceae)

We investigated the distribution of genetic variation and the relationship between population size and genetic variation in the rare plant Gentianella germanica using RAPD (random amplified polymorphic DNA) profiles. Plants for the analysis were grown from seeds sampled from 72 parent plants in 11 G. germanica populations of different size (40-5000 fruiting individuals). In large populations, seeds were sampled from parents in two spatially distinct subpopulations comparable in area to the total area covered by small populations. Analysis of molecular variance revealed significant genetic variation among populations (P <0.001), while genetic variation among subpopulations was marginally significant (P <0.06). Average molecular variance within subpopulations in large populations did not differ significantly from whole-population values. There was a positive correlation between genetic variation and population size (P <0.01). Genetic variation was also positively correlated with the number of seeds per plant in the field (P <0.02) and the number of flowers per planted seed in a common garden experiment (P <0.051). We conclude that gene flow among natural populations is very limited and that reduced plant fitness in small populations of G. germanica most likely has genetic causes. Management should aim to increase the size of small populations to minimize further loss of genetic variation. Because a large proportion of genetic variation is among populations, even small populations are worth preserving.     

Population size and the nature of genetic load in Gentianella germanica

Abstract Theory predicts a significant relationship between the size of a population and the magnitude and composition of its genetic load, but few natural populations have been investigated. We examined the magnitude of genetic load due to recessive deleterious alleles (GL) both segregating and fixed within Gentianella germanica populations of varying size by selfing and reciprocally crossing plants within and between natural populations according to a partial diallel design and by comparing the performance of the experimental progeny in a common-garden experiment. The results show that GL for total fitness in small populations (fewer than 200 plants) was mainly due to fixed recessive deleterious alleles, whereas GL for total fitness in larger populations (more than 200 plants) appeared to be mainly due to segregating deleterious recessive alleles. The total fitness of selfed plants increased with decreasing population size, indicating some purging of deleterious alleles associated with declining population sizes. The magnitudes of GL due to fixed deleterious alleles in small populations and segregating deleterious alleles in large populations, however, were overall similar, suggesting that purging selection was an insignificant force when compared to genetic drift in determining the magnitude of GL in small natural populations in this species. The results of this study highlight the importance of population size in determining the dynamics of genetic loads of natural populations and are overall in line with a large body of theoretical work indicating that small populations may face higher extinction risks due to the fixation and accumulation of deleterious alleles of small effect.     

Irrigation and fertilization effects on seed number,size, germination and seedling growth: implications for desert shrub establishment

Plants with limited resources adjust partitioning among growth, survival, and reproduction. We tested the effects of water and nutrient amendments on seed production, size, and quality in Sarcobatus vermiculatus (greasewood) to assess the magnitude and importance of changes in reproductive partitioning. In addition, we assessed interactions among the environment of seed-producing plants (adult plant scale), seed size, and seedling microenvironment (seedling scale) on successful seedling establishment. Interactions of these factors determine the scale of resource heterogeneity that affects seedling establishment in deserts. Both total number of seeds produced per plant and seed quality (weight and germination) increased significantly in the enriched treatment in a 3-year field experiment. Seedling length 3 days after germination and seed N concentration, other measures of seed quality, were higher for seed from both irrigated and enriched plants than for seed from control plants. Field S. vermiculatus seed production and quality can be substantially increased with irrigation and nutrient enrichment at the adult plant scale and this allows management of seed availability for restoration. However, based on a greenhouse study, seedling environment, not the environment of the seed-producing plant or seed size, was the most important factor affecting seedling germination, survival, and growth. Thus it appears that production of more seed may be more important than improved seed quality, because higher quality seed did not compensate for a low-resource seedling environment. For both natural establishment and restoration this suggests that heterogeneity at the scale of seedling microsites, perhaps combined with fertilization of adult shrubs (or multi-plant patches), would produce the greatest benefit for establishing seedlings in the field.     

生物土壤结皮对种子萌发和幼苗建植的影响研究进展

生物土壤结皮（BSC）在陆地生态系统中具有重要的生态地位,尤其是旱地生态系统中,BSC占据了种子植物之间的广阔地面。因此,BSC的发展必然影响种子植物更新过程与植被空间格局;但其作用方式、影响程度等因相关研究涉及多气候要素、土壤类型、BSC组成物种和种子植物物种的差异及其不同组合,导致目前的研究结论存在广泛争议。研究综合论述了BSC改变地表微形态对种子传播过程的影响;BSC改变土壤特性（物理、化学、生物学）对种子萌发和幼苗存活与建植等关键环节的影响;并结合种子形态特征及种子萌发、幼苗建植的性状等,综合分析了BSC对种子传播、种子萌发与幼苗建植等更新过程的潜在影响机理;探讨了目前研究矛盾性结论产生的原因。总体来说,深入研究并全面揭示BSC对种子植物更新过程的影响,应加强学科交叉,将分子生物学、植物生理学、生物化学等微观研究,与遥感、野外生态因子过程监测、控制实验等宏观、中观研究结合,从机理到过程方面动态研究BSC对种子植物更新过程的影响,并引入水文模型、气候模型、种群动态模型等模型预测方法,研究气候变化、各类干扰频发的情景下,BSC对种子传播、萌发及幼苗建植过程的潜在影响,以期促进对BSC与种子植物间相互关系的研究,加深对干旱脆弱生态系统植被发展规律的认识。     

Effects of temperature and storage length on seed germination and the effects of light conditions on seedling establishment with respect to seed size in Ligularia virgaurea

Ligularia virgaurea is widely distributed in the alpine meadows of the eastern Qinghai-Tibet plateau. We studied the effects of temperature and seed storage length on seed germination in a laboratory experiment, the effects of seed mass and light intensity on seed emergence in a pot experiment, and the effect of meadow disturbance intensity on seed emergence in a field experiment. Our results showed that seeds of L. virgaurea germinated well under a wide range of temperatures. Germination percentage decreased with increased seed storage length. When seeds were stored for either 6 or 12 months the germination percentage increased with decreasing temperature. The emergence percentage of large seeds was higher than that of small seeds. Seedling survival of large seeds was greater than that of small seeds in 75% and 50% of natural irradiance. Large seeds of L. virgaurea were more successful colonizers in the alpine meadows. Seeds of L. virgaurea germinated in both disturbed meadows and non-disturbed meadows, but seed emergence and seedling establishment were higher in the disturbed meadows, that is, disturbance could facilitate the recruitment and establishment of L. virgaurea .     

黄帚橐吾种子生产的花序位置效应及其对幼苗建植的影响

关于开花植物花序内变异原因的研究有很多,而关于花序内资源分配的位置效应产生影响的研究并不常见。通过对青藏高原高寒草甸退化草地一种花序轴基向发育的优势毒杂草黄帚橐吾(Ligularia virgaurea)的野外调查和方差分析发现,其种子生产存在显著的位置效应。结实率和种子平均重表现出顶部头状花序基部头状花序的变化趋势,胚珠平均数则基部顶部,而中部种子平均数最多。通过室内萌发实验和野外盆栽实验研究了位置效应对幼苗建植能力的影响,发现基部种子的萌发开始时间比顶部种子早,萌发率和出苗率在部位间的差异不显著,而部位间幼苗存活率表现一致;幼苗的根冠比和光合速率表现出顶部基部的变化趋势,而幼苗总生物量的差异不显著,这些幼苗特性在两种肥力条件下表现一致,这说明不同花序部位种子的差异是该种植物所固有的特征,且部位间种子的萌发和幼苗生长对策不同。但相对于基部种子,顶部种子扩散距离更远,拓展到新生境的能力更强,相对较高的幼苗根冠比和净光合速率增强了其幼苗建植能力,对自然种群的更新补充有着更重要的生态学意义,对扩大种群和植被更新有更大的贡献。     

Effects of seed size and emergence time on tree seedling establishment: importance of developmental constraints

Arafura filesnakes (Acrochordus arafurae) are large (up to 2.5 m, 5 kg) aquatic nonvenomous snakes that feed entirely on fishes. A 10-year field study in the Australian wet-dry tropics revealed strong correlations between rainfall patterns, fish abundance, and snake population dynamics. All of these characteristics showed considerable annual variation. High rainfall late in the wet season (February–March) caused prolonged inundation of the floodplain. Following such years, dry-season sampling revealed that fishes were abundant, filesnakes were in good body condition, and a high proportion of adult female filesnakes were reproductive. Annual variation in recruitment to the population (as judged by the relative abundance of yearling snakes) was also correlated with fish abundance and thus, with rainfall patterns in the late-wet season. Our results fit well with those from other studies on a diverse array of aquatic and terrestrial species within the wet-dry tropics. Annual variation in rainfall patterns, via its effects on prey abundance, may drive the population dynamics of many tropical predators. Received: 28 June 1999 / Accepted: 29 February 2000     

Effect of seed predation on seed bank size and seedling recruitment of bush lupine (Lupinus arboreus)

Whether seed consumers affect plant establishment is an important unresolved question in plant population biology. Seed consumption is ubiquitous; at issue is whether seedling recruitment is limited by safe-sites or seeds. If most seeds inhabit sites unsuitable for germination, post-dispersal seed consumption primarily removes seeds that would otherwise never contribute to the population and granivory has minimal impacts on plant abundance. Alternatively, if most seeds ultimately germinate before they lose viability, there is greater potential for seed consumption to affect plant recruitment. Of the many studies on seed consumption, few ask how seed loss affects seedling recruitment for species with long-lived seed banks. We examined post-dispersal seed predation and seedling emergence in bush lupine (Lupinus arboreus), a woody leguminous shrub of coastal grasslands and dunes in California. We followed the fate of seeds in paired experimental seed plots that were either protected or exposed to rodent granivores in grassland and dune habitats. Significantly more seeds were removed by rodents in dunes than grasslands. In dunes, where rodent granivory was greatest (65% and 86% of seeds removed from plots by rodents in two successive years), there is a sparse seed bank (6.6 seeds m⁻²), and granivory significantly reduced seedling emergence (in the same two years, 18% and 19.4% fewer seedlings emerged from exposed versus protected plots), suggesting seed rather than safe-site limited seedling recruitment. In contrast, rodents removed an average of 6% and 56% of seeds from grassland plots during the same two years, and the grassland seed bank is 43-fold that of the dunes (288 seeds m⁻²). Even high seed consumption in the second year of the study only marginally influenced recruitment because seeds that escaped predation remained dormant. Burial of seeds in both habitats significantly reduced the percentage of seeds removed by rodents. Results suggest that granivores exert strong but habitat-dependent effects on lupine seed survival and seedling emergence. Received: 24 October 1996 / Accepted: 4 February 1997     

Role of non-microbial biostimulants in regulation of seed germination and seedling establishment

Plant Growth Regulation - The application of biostimulants from natural origin represents one of the most innovative and promising strategies to promote plant growth and improve crop productivity....     

Cowpea ribonuclease: properties and effect of NaCl-salinity on its activation during seed germination and seedling establishment

Pitiúba cowpea [Vigna unguiculata (L.) Walp] seeds were germinated in distilled water (control treatment) or in 100 mM NaCl solution (salt treatment), and RNase was purified from different parts of the seedlings. Seedling growth was reduced by the NaCl treatment. RNase activity was low in cotyledons of quiescent seeds, but the enzyme was activated during germination and seedling establishment. Salinity reduced cotyledon RNase activity, and this effect appeared to be due to a delay in its activation. The RNases from roots, stems, and leaves were immunologically identical to that found in cotyledons. Partially purified RNase fractions from the different parts of the seedling showed some activity with DNA as substrate. However, this DNA hydrolyzing activity was much lower than that of RNA hydrolyzing activity. The DNA hydrolyzing activity was strongly inhibited by Cu²⁺, Hg²⁺, and Zn²⁺ ions, stimulated by MgCl₂, and slowly inhibited by EDTA. This activity from the most purified fraction was inhibited by increasing concentrations of RNA in the reaction medium. It is suggested that the major biological role of this cotyledon RNase would be to hydrolyze seed storage RNA during germination and seedling establishment, and it was discussed that it might have a protective role against abiotic stress during later part of seedling establishment.     

海草床种子扩散过程及种子库形成机制研究进展

海草是唯一一类可以完全生活在海水中的高等被子植物,具有重要的生态服务功能和巨大的经济价值。但受气候变化和人类活动的双重影响,海草床退化趋势日益严峻。海草床生态系统受到外界胁迫后的稳定性和恢复能力很大程度上依赖于有性繁殖即种子繁殖,当胁迫造成海草死亡等不可逆转的伤害时,通过沉积物种子库能够进行种群维持和自我更新,因此研究海草种子扩散过程及种子库形成机制对海草生态系统稳定性的维持具有重要意义。综述了海草生活史类型、种子繁殖特征、种子扩散过程及影响因素、种子库形成机制等。在此基础上总结了目前研究存在的几方面不足和未来展望：1)不同环境胁迫条件对海草有性繁殖努力的影响研究;2)海草种子二次扩散的影响因素和扩散机制研究;3)沉积物沉降和再悬浮对种子扩散和截留的影响研究;4)环境因素变化下种子库的潜在分布和海草适宜生境预测与模拟。本研究以期为海草床生态系统的保护恢复研究提供理论参考。