Effect of phylogeny, life history and habitat correlates on seed germination of 69 arid and semi-arid zone species from northwest China

We tested the effects of phylogeny, life history and habitat conditions for seedling establishment on seed germination of 69 arid/semi-arid zone species from northwest China. Final germination percentages had a bimodal distribution, whereas, days to first germination were skewed toward short periods. Stepwise multi-way ANOVAs showed that the amount of variance in final germination percentage among species for seeds incubated in light was explained by phylogeny (13.6%), dispersal mode (11.9%), seed mass (3.6%), and habitat (0.2%); in darkness by dispersal mode (17.5%), phylogeny (12.7%), seed mass (5.7%), and habitat (0%). A series of ANOVAs revealed that the variance in final germination percentages among species is largely dependent upon phylogeny and dispersal mode but that it is also influenced by seed mass and habitat. The effects of dispersal mode and seed mass on final germination percentage among species were phylogenetic group- and habitat-specific. Wind-dispersed seeds had higher germination percentages than unassisted and vertebrate-dispersed seeds. Compared to xerophytes, mesophytes tended to have smaller seed mass, higher mean germination percentages and a greater effect of seed mass on final germination fractions. Our results suggest that phylogeny and life history should be taken into account in studies on the role of natural selection in regulation of seed germination.     

Community-wide germination strategies in an alpine meadow on the eastern Qinghai-Tibet plateau: phylogenetic and life-history correlates

In this study, we built up a database of 633 species (48 families, 205 genera) from an alpine meadow on the eastern Qinghai-Tibet plateau. Our objective was to assess the effects of phylogenetic and life-history (life form, perenniality, seed size, dispersal strategy and period) background on the community-wide germination strategies. We found that the seeds of shrubs, perennials, and well-dispersed plants, and the smaller seeds germinated more and comparatively earlier. In one-way ANOVAs, phylogenetic groups explained 12% of the variance in GT (mean germination time for all seeds germinated of each species); life-history attributes, such as seed size, dispersal strategy, perenniality and life form explained 10%, 7%, 5%, and 1% respectively, and dispersal period had no significant effect on GT. Multifactorial ANOVAs revealed that the three major factors contributing to differences in GT were phylogenetic relatedness, seed size and dispersal strategy (explained 4%, 5% and 4% of the interspecific variation independently, respectively). Thus, seeds germination strategies were significantly correlated with phylogenetic and life-history relatedness. In addition, phylogenetic relatedness had close associations and interactions with seed size and dispersal strategy. Then, we think phylogeny and life-history attributes could not be considered mutual exclusively. Seed germination, like any other trait, is shaped by the natural history of the species and by the evolutionary history of the lineage. And a large percentage of the variance remained unexplained by our model, which suggested important selective factors or parameters may have been left out of this analysis. Electronic Supplementary Material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

青藏高原东缘61种常见木本植物种子萌发特性及其与生活史的关联

为了评价亚高山地带木本植物种子萌发对策及其与生活史关联, 该文对青藏高原东缘61种常见木本植物的种子萌发特征以及种子萌发与种子大小、扩散方式、海拔及生境的关系进行了调查分析。结果表明: 1) 61种植物的种子萌发率呈现偏斜、双峰分布。6种植物(9.8%)的种子萌发率大于80%, 13种(21.3%)的萌发率在60%~80%之间, 9种(14.8%)的萌发率在40%~60%之间, 8种(13.1%)的萌发率在20%~40%之间, 25种(41%)的萌发率小于20%, 其中8种植物的种子萌发率为零, 显示了该生境中木本植物的种子以休眠、低萌发率或连续稳定的萌发对策占优势, 以避免不利的环境因素。2)种子扩散方式是决定61种木本植物种间萌发率变异(R² = 24.1%, p < 0.001) 和萌发开始时间变异(R ²= 21.3%, p < 0.01) 的主要因素; 种子大小、母体植株的海拔和生境对种子萌发率和萌发开始时间几乎没有影响(p > 0.05); 风扩散的种子比脊椎动物扩散和无助扩散的种子有更高的萌发率(F = 9.219, p < 0.001) 和较早的萌发开始时间(F = 6.772, p < 0.01)。说明植物生活史特征如种子萌发与扩散方式之间存在固有而紧密的联系, 扩散能力强的种子(如风扩散)由于能散布到较远而空旷的生境, 可能避免了各种有害因素或个体竞争, 因而种子以较强的萌发能力进行拓殖; 扩散能力弱的种子(如无助扩散)由于其散布距离较近, 种子以休眠或降低萌发的方式来避免个体或同胞竞争以及各种有害因素, 以获得最大的生态利益并确保物种延续。     

Impacts of growth form and phylogenetic relatedness on seed germination: A large‐scale analysis of a subtropical regional flora

Plant regeneration strategy plays a critical role in species survival and can be used as a proxy for the evolutionary response of species to climate change. However, information on the effects of key plant traits and phylogenetic relatedness on seed germination is limited at large regional scales that vary in climate. To test the hypotheses that phylogenetic niche conservatism plays a critical force in shaping seed ecophysiological traits across species, and also drives their response to climatic fluctuation, we conducted a controlled experiment on seed germination and determined the percentage and rate of germination for 249 species in subtropical China under two temperature regimes (i.e., daily 25°C; daily alternating 25/15°C for each 12 hr). Germination was low with a skewed distribution (mean = 38.9% at 25°C, and 43.3% at 25/15°C). One fifth of the species had low (<10%) and slow (4–30 days) germination, and only a few (8%) species had a high (>80%) and rapid (1.2–6.6 days) germination. All studied plant traits (including germination responses) showed a significant phylogenetic signal, with an exception of seed germination percentage under the alternating temperature scenario. Generalized linear models (GLMs) and phylogenetic generalized estimation equations (GEEs) demonstrated that growth form and seed dispersal mode were strong drivers of germination. Our experimental study highlights that integrating plant key traits and phylogeny is critical to predicting seed germination response to future climate change.     

Reserve mass and dispersal investment in relation to geographic range of plant species: phylogenetically independent contrasts

Recent studies have reported conflicting evidence about correlations between seed size and plant species geographic range sizes. Using phylogenetically independent contrasts (PICs) within genera, we found no consistent differences in reserve mass between species with similar dispersal morphology and «wide>> versus «narrow>> geographic ranges. There was also no tendency within genera for broad ranged species to be those that allocate a larger percentage of the resources invested in each diaspora to dispersal structures. PICs were also constructed between species having a tenfold difference in seed size. In these PICs, the larger seeded species often occupied a greater number of regions than species with smaller seed sizes. This result was generated primarily through the comparison of species from different genera, families or higher level taxa which differed not only in seed mass but also in dispersal modes and growth forms. Finally, comparing species within Acacia and Eucalyptus having similar seed size but different dispersal modes, we found that bird dispersal (in Acacia ) and possession of a wing for wind dispersal (in Eucalyptus ) was associated with wider geographic range compared to lower-investment dispersal modes. Taken together, these comparisons indicate that seed size is not itself important as a factor influencing breadth of geographic range. Dispersal mode and growth form may have an influence, however, and seed size differences may be associated with contrasts in dispersal mode or growth form.     

五蕊寄生种子萌发特性

采用室内与野外实验相结合的方法,对五蕊寄生（Dendrophthoe pentandra）种子萌发特性以及鸟类消化过程对其种子萌发的影响进行研究。结果表明：五蕊寄生种子对温度和光照较敏感,与黑暗条件相比,光照能显著提高种子的萌发率;20℃为种子萌发的最适温度,萌发率达53.8%,高温35℃及低温10℃均不利于种子萌发。研究发现纯色啄花鸟（Dicaeum concolor）是该植物的主要食果者,种子外的果肉不是限制种子萌发的关键因素,人为去除果肉并不能提高种子的萌发率,但可缩短种子萌发的时间,果实通过纯色啄花鸟类肠道消化后,随着粪便排出的种子萌发率显著提高（达69%）,暗示纯色啄花鸟在五蕊寄生的种子萌发过程中具有重要的作用。     

Cracking the case: Seed traits and phylogeny predict time to germination in prairie restoration species

Traits are important for understanding how plant communities assemble and function, providing a common currency for studying ecological processes across species, locations, and habitat types. However, the majority of studies relating species traits to community assembly rely upon vegetative traits of mature plants. Seed traits, which are understudied relative to whole‐plant traits, are key to understanding assembly of plant communities. This is particularly true for restored communities, which are typically started de novo from seed, making seed germination a critical first step in community assembly and an early filter for plant establishment. We experimentally tested the effects of seed traits (mass, shape, and embryo to seed size ratio) and phylogeny on germination response in 32 species commonly used in prairie grassland restoration in the Midwestern USA, analyzing data using time‐to‐event (survival) analysis. As germination is also influenced by seed dormancy, and dormancy break treatments are commonly employed in restoration, we also tested the effects of two pretreatments (cold stratification and gibberellic acid application) on time to germination. Seed traits, phylogeny, and seed pretreatments all affected time to germination. Of all traits tested, variables related to seed shape (height and shape variance) best predicted germination response, with high‐variance (i.e., pointier and narrower) seeds germinating faster. Phylogenetic position (the location of species on the phylogenetic tree relative to other tested species) was also an important predictor of germination response, that is, closely related species showed similar patterns in time to germination. This was true despite the fact that all measured seed traits showed phylogenetic signal, therefore phylogeny provided residual information that was not already captured by measured seed traits. Seed traits, phylogenetic position, and germination pretreatments were important predictors of germination response for a suite of species commonly used in grassland restoration. Shape traits were especially important, while mass, often the only seed trait used in studies of community assembly, was not a strong predictor of germination timing. These findings illustrate the ecological importance of seed traits that are rarely incorporated into functional studies of plant communities. This information can also be used to advance restoration practice by guiding restoration planning and seed mix design.     

Seed germination in temperate rain forest species of southern Chile: chilling and gap-dependency germination

Canopy gap and chilling requirements for seed germination were assessed in 61 and 44 species, respectively, in the temperate rain forest of southern Chile. Germination assays within canopy gap and understorey were carried out under natural conditions. Germination tests in cold stratified and non-stratified seeds were performed under laboratory conditions. Seeds were collected of common trees, shrubs, vines and perennial herbs of forests in southern Chile. Final percent germination was significantly enhanced under canopy gap conditions in 19 species, and significantly reduced in 11 species. Germination proved indifferent under gap vs. understorey conditions in half the species tested. Cold stratification affected germination in 11 species: significantly increasing final germination of four species, and significantly decreasing final percent germination of seven species. A Principal Components Analysis (PCA) was applied with the purpose of identifying groups of species with similar germination strategies. Four attributes were included in the PCA: final germination percentage in canopy gap, germination rate in the laboratory, and gap-and chilling-dependency indices (EGAP and STRAT, respectively). The first axis separated species mainly on EGAP variation while the second axis separated them mainly according to STRAT variation; the two axes together explaining 73% of the among-species variation. A small group of trees and vines germinating best in the understorey and neutral to chilling could clearly be distinguished from the remaining species analyzed. Multifactorial ANOVAs were used for evaluating the combined effect of successional status, seed mass, dispersal period, life form, phylogenetic categories, and dispersal syndrome on EGAP and STRAT variation. The EGAP value of secondary successional species proved significantly greater than that of primary successional species and species with endozoochorous seed dispersal were significantly less dependent on chilling (according to STRAT values) compared to species with mainly wind-dispersed seeds. The possible implications of these results for seedling establishment are discussed. This revised version was published online in August 2006 with corrections to the Cover Date.     

Seed dispersal by the Florida box turtle (<Emphasis Type="Italic">Terrapene carolina bauri</Emphasis>) in pine rockland forests of the lower Florida Keys,United States

Seed dispersal by animals is one of the most important plant-animal mutualisms, but saurochory, the dispersal of seeds by reptiles, has received little attention. We investigated the role of the Florida box turtle (Terrapene carolina bauri) as a seed dispersal agent in pine rockland forests of the lower Florida Keys and examined the effect of turtle digestion on seed germination. We obtained seeds of 11 species with fleshy fruits and 2 species with non-fleshy fruits (a grass and legume) from the feces of 145 box turtles collected on Key Deer National Wildlife Refuge from 1999 to 2000. We planted the seeds of nine species and germination percentage (percentage of seeds that germinated during the experiment) varied from 10% to 80%. Comparative germination experiments were conducted with Thrinax morrissii, Serenoa repens, and Byrsonima lucida. We compared the germination percentage and germination rate (number of days from planting to seedling emergence) of seeds from three treatments (seeds recovered from feces, control seeds with pulp, and control seeds without pulp) and continued these experiments for up to 2 years. Passage through the box turtle digestive tract greatly enhanced the germination percentage and germination rate of S. repens, but decreased the germination percentage of B. lucida and T. morrissii, and decreased germination rate for T. morrissii. Subsequent destructive seed viability tests revealed that many ungerminated T. morrissii seeds remained viable, suggesting long-term seed dormancy may occur, even after passage through the turtle digestive system. In addition, the proportion of ungerminated seeds which remained viable was greater for seeds recovered from turtle feces than from control seeds with pulp. Furthermore, removal of fleshy pulp either manually or by the turtle digestive system may allow T. morrissii to escape insect predation.     

A reassessment of the role of simple seed traits in mortality following herbivore ingestion

Grazing mammals are regarded as major vectors in seed dispersal of grassland plants, through seed ingestion and subsequent excretion (endozoochory). The (evolutionary) ecology underlying this dispersal mode is relatively poorly understood because there are limited data, among others, on how seed attributes perform in this process (and could thus be selected for). For seed mortality following ingestion, contrasting patterns described for the role of simple seed traits seem partly due to inadequate comparative analytical methods. We conducted a feeding experiment in which controlled seed quantities from 48 grassland herb and grass species were fed to cattle. Seed mass, length and shape measurements were related to seed mortality rate using phylogenetically independent contrasts, which account for taxonomic interdependence. The proportion of seeds surviving the digestive tract was generally low, but it reached 100%, with increased germination for some species. Neither seed size nor shape correlated significantly with mortality. Structural traits are likely to overrule these simple seed traits, as illustrated by high survival of species having water-impermeable seeds. As this trait has interesting properties with respect to intraspecific variation and genetics, it could provide opportunities for inference on evolutionary consequences for this type of dispersal.     

异子蓬二型种子的萌发与休眠特性及其生态适应

异子蓬为中亚荒漠的特有种,其果实为胞果,花被片在结果时宿存包被果实形成浆果状的散布单位.对异子蓬的散布单位、果实形态进行比较观察,并在5/15℃、5/25℃、15/25 ℃(暗/光=12h/12h)3个温变周期下进行种子萌发实验,结果表明:(1)异子蓬的散布单位绿色、浆果状,有大小两种形态;大小两种散布单位内包被的果实在形状、大小、颜色上存在着明显差异.(2)大的散布单位内包被的果实圆形,扁平,褐色,无光泽,直径(2.64±0.03) mm,重量(3.68±0.04) mg;小的散布单位内包被的果实双凸镜形,黑色,有光泽,直径(2.40±0.04) mm,重量(2.86±0.06) mg.(3)褐色种子在3个温变周期下的萌发率都在81%以上,萌发速度快,而黑色种子萌发率低,萌发慢,具有休眠现象.(4)划破种皮及低温层积处理可显著提高黑色种子在3个温变周期下的萌发率,延长储存时间也可有效地促进黑色种子的萌发,表明黑色种子处于非深度生理休眠状态.异子蓬产生的两种不同类型的种子及其在时间和空间上的萌发差异对荒漠异质环境具有重要的适应意义.     

Frugivory and Seed Dispersal by Northern Pigtailed Macaques (Macaca leonina), in Thailand

Tropical rain forest conservation requires a good understanding of plant–animal interactions. Seed dispersal provides a means for plant seeds to escape competition and density-dependent seed predators and pathogens and to colonize new habitats. This makes the role and effectiveness of frugivorous species in the seed dispersal process an important topic. Northern pigtailed macaques (Macaca leonina) may be effective seed dispersers because they have a diverse diet and process seeds in several ways (swallowing, spitting out, or dropping them). To investigate the seed dispersal effectiveness of a habituated group of pigtailed macaques in Khao Yai National Park, Thailand, we examined seed dispersal quantity (number of fruit species eaten, proportion in the diet, number of feces containing seeds, and number of seeds processed) and quality (processing methods used, seed viability and germination success, habitat type and distance from parent tree for the deposited seeds, and dispersal patterns) via focal and scan sampling, seed collection, and germination tests. We found thousands of seeds per feces, including seeds up to 58 mm in length and from 88 fruit species. Importantly, the macaques dispersed seeds from primary to secondary forests, via swallowing, spitting, and dropping. Of 21 species, the effect of swallowing and spitting was positive for two species (i.e., processed seeds had a higher % germination and % viability than control seeds), neutral for 13 species (no difference in % germination or viability), and negative (processed seeds had lower % germination and viability) for five species. For the final species, the effect was neutral for spat-out seeds but negative for swallowed seeds. We conclude that macaques are effective seed dispersers in both quantitative and qualitative terms and that they are of potential importance for tropical rain forest regeneration.     

光质及种子大小对普洱地区14种植物种子萌发的影响

以普洱地区14种常见植物种子为材料,在实验室条件下研究了其在白光、黑暗、红光和蓝光条件下的萌发特性,并分析了种子大小与萌发率、萌发速率、萌发开始时间的关系。结果表明:光质对四方蒿、沙针、尖子木、藿香蓟种子萌发率和萌发速率均有显著影响(P0.05)。光质对大叶斑鸠菊、云南山枇花、臭灵丹、车桑子、光萼猪屎豆、葫芦茶、云南地桃花、西南宿苞豆、岗柃、中国宿苞豆10个物种的种子萌发率和萌发速率均没有显著影响(P0.05),以上物种中除中国宿苞豆外,其他物种种子萌发率均在20%以下,处于休眠状态。四方蒿种子在白光(89.9%)和红光(84.7%)下萌发率最高,红光下种子萌发最快(4.93),蓝光下种子萌发开始时间最晚(11.3 d);沙针种子在白光下萌发率最高(80.4%)、萌发速率最快(2.71),在黑暗和蓝光下萌发率较低(43.9%和38%)、萌发速率最慢(0.73和0.85),白光、红光下萌发开始最早(11 d),黑暗条件下萌发开始最晚(21.7 d);尖子木种子萌发率在白光、黑暗、蓝光下均在86%以上,而红光下仅32%且萌发速率最慢(1.29),在蓝光下萌发开始时间最晚(13 d);藿香蓟种子萌发率和萌发速率在红光下最高(分别为71.3%和6.46),黑暗条件下最低(分别为42.5%和2.62);大叶斑鸠菊萌发开始时间在黑暗条件下最早(6 d),其次是白光下(7 d),蓝光和红光下较晚,分别为8 d和7.7 d。14个物种种子的萌发率与种子大小间均有显著负相关关系;种子萌发速率、萌发开始时间与种子大小间也有负相关关系,但不显著;种子大小与萌发率、萌发速率和萌发开始时间的关系不会随着光质的变化而发生变化。     

Effects of seed colour heterogeneity on germination behaviour of the desert plant Lotononis platycarpa (Fabaceae)

We aimed to determine the ecological role of three seed morphs observed for the first time in a desert population of Lotononis platycarpa (Fabaceae), with respect to their germination requirements. Seeds sorted by seed coat colour (olive green, orange and brown) were germinated under laboratory conditions under two photoperiods (12/12‐h light and continuous dark) and three alternating temperature regimes (15/25, 20/30, 25/35°C). We found that the three distinct seed types differ in their seed mass, germination percentage and speed of germination. Overall, the light‐incubated seeds germinated with higher percentages than seeds in the total darkness. Furthermore, seeds with orange coat germinated with higher percentages at 15/25 and 25/35°C (up to 60%, for both) and significantly faster than the other two colour morphs. Our results suggest an adaptive significance of seed colour heterogeneity in the harsh desert habitat inhabited by the study species.     

CONVERGENT AND CORRELATED EVOLUTION OF MAJOR LIFE‐HISTORY TRAITS IN THE ANGIOSPERM GENUS LEUCADENDRON (PROTEACEAE)

Natural selection is expected to cause convergence of life histories among taxa as well as correlated evolution of different life‐history traits. Here, we quantify the extent of convergence of five key life‐history traits (adult fire survival, seed storage, degree of sexual dimorphism, pollination mode, and seed‐dispersal mode) and test hypotheses about their correlated evolution in the genus Leucadendron (Proteaceae) from the fire‐prone South African fynbos. We reconstructed a new molecular phylogeny of this highly diverse genus that involves more taxa and molecular markers than previously. This reconstruction identifies new clades that were not detected by previous molecular study and morphological classifications. Using this new phylogeny and robust methods that account for phylogenetic uncertainty, we show that the five life‐history traits studied were labile during the evolutionary history of the genus. This diversity allowed us to tackle major questions about the correlated evolution of life‐history strategies. We found that species with longer seed‐dispersal distances tended to evolve lower pollen‐dispersal distance, that insect‐pollinated species evolved decreased sexual dimorphism, and that species with a persistent soil seed‐bank evolved toward reduced fire‐survival ability of adults.     

Seed dispersal by ants in the semi-arid Caatinga of North-East Brazil

BACKGROUND AND AIMS: Myrmecochory is a conspicuous feature of several sclerophyll ecosystems around the world but it has received little attention in the semi-arid areas of South America. This study addresses the importance of seed dispersal by ants in a 2500-km(2) area of the Caatinga ecosystem (north-east Brazil) and investigates ant-derived benefits to the plant through myrmecochory. METHODS: Seed manipulation and dispersal by ants was investigated during a 3-year period in the Xingó region. Both plant and ant assemblages involved in seed dispersal were described and ant behaviour was characterized. True myrmecochorous seeds of seven Euphorbiaceae species (i.e. elaiosome-bearing seeds) were used in experiments designed to: (1) quantify the rates of seed cleaning/removal and the influence of both seed size and elaiosome presence on seed removal; (2) identify the fate of seeds dispersed by ants; and (3) document the benefits of seed dispersal by ants in terms of seed germination and seedling growth. KEY RESULTS: Seed dispersal by ants involved one-quarter of the woody flora inhabiting the Xingó region, but true myrmecochory was restricted to 12.8 % of the woody plant species. Myrmecochorous seeds manipulated by ants faced high levels of seed removal (38-84 %) and 83 % of removed seeds were discarded on ant nests. Moreover, seed removal positively correlated with the presence of elaiosome, and elaiosome removal increased germination success by at least 30 %. Finally, some Euphorbiaceae species presented both increased germination and seedling growth on ant-nest soils. CONCLUSIONS: Myrmecochory is a relevant seed dispersal mode in the Caatinga ecosystem, and is particularly frequent among Euphorbiaceae trees and shrubs. The fact that seeds reach micro-sites suitable for establishment (ant nests) supports the directed dispersal hypothesis as a possible force favouring myrmecochory in this ecosystem. Ecosystems with a high frequency of myrmecochorous plants appear not to be restricted to regions of nutrient-impoverished soil or to fire-prone regions.     

How are the phenologies of ripening and seed release affected by species' ecology and evolution?

The phenology of seed ripening and release are important for dispersal, reproductive success and survival of plants. Most phenological studies, however, consider early phenological phases. Here, we examined the ecological and evolutionary basis of ripening and seed release phenology. We monitored single flower phenology for 104 plant species from 30 families and three life forms from central Europe. Further, we undertook an associate monitoring study along an elevational gradient over two years. We calculated temperature demands (as growing degree days) for ripening and seed release and examined them with respect to the species’ seed mass, life form, dispersal mode and phylogeny. We found a strong correlation between species’ seed mass and temperature demands for ripening. For both variables seed mass and temperature demands for seed ripening, we found a strong effect of the species phylogeny. These phylogenetic signals indicate that the evolutionary history of the species’ lineage affects its seed mass and the temperature demands for seed ripening. Among the studied life forms, shrub species showed the most efficient ripening process. Anemochorous species showed lower relative humidity during seed release than epizoochorous species. For anemochorous species, the synchronisation of release timing with periods that show favourable environmental conditions for wind dispersal could be interpreted as a phenological adaptation to increase dispersal distances. According to the monitoring along the elevational gradient, individuals from higher altitudes showed lower temperature demands for ripening than individuals from lower altitudes. This might tentatively indicate physiological adaptations to lower temperature demands for locations with a shorter growing season. Our study provides basic insights into the ecological, environmental and evolutionary constraints that shape the ripening and seed release phenology of plants. We introduce data that can be used to advance existing models of ripening phenology, seed release and plant spread.     

Ecological genetics of seed germination regulation in Bromus tectorum L.

Regulation of seed germination phenology is an important aspect of the life history strategy of invading annual plant species. In the obligately selfing winter annual grass Bromus tectorum, seeds are at least conditionally dormant at dispersal in early summer and lose dormancy through dry-afterripening. Patterns of germination response at dispersal vary among populations and sometimes across years within populations. To assess the relative contribution of genotype and maturation environment to this variation, we grew progeny of ten parental lines from each of six contrasting populations in a common greenhouse environment. We then tested the germination responses of recently harvested seeds of the putative full-sib progeny at five incubation temperatures. Significant germination response differences among populations were observed in greenhouse cultivation, and major differences among full-sib families were evident for some populations and traits. Among-population variation accounted for over 90% of the variance in each trait, while within-family variance accounted for 1% or less. Germination responses of greenhouse-grown progeny were positively correlated with the responses of wild-collected seeds, but there was a tendency for lowered dormancy at higher incubation temperatures. This tendency was more marked in populations from cold desert, foothill, and plains habitats, suggesting a genotype-maturation environment interaction. Differences among populations in the amount of among-family variance were more evident at lower incubation temperatures, while among-family variance was more uniformly low at summer incubation temperatures. Populations from predictable extreme environments (subalpine meadow and warm desert margin) showed significantly less variation among families than populations from less predictable cold desert, foothill, and plains environments. Low among-family variance was not specifically associated with small population size or marginality of habitat, as small marginal populations from unpredictable environments showed variance as high as that of large populations. In populations with high among-family variance for germination traits, germination responses tended to be correlated across incubation temperatures, making it possible to characterize families in terms of their general dormancy status. The results indicate that seed germination regulation in this species is probably under strong genetic control, and that habitats with temporally varying selection are occupied by populations that tend to be more polymorphic in terms of their germination response patterns. Received: 19 May 1998 / Accepted: 27 January 1999     

The Light-Sensitivity of Germination of Ficus benjamina var. nuda (Moraceae) Seeds and Its Ecological Significance

In this paper , Ficus benjamina L . var . nuda seeds were used as experimental materials, effects of temperature, light , phytohormone ( gibberellin , 6-benzyladenine and ethylene ) , nitrogenous compounds ( sodium nitroprusside,nitrate , and nitrite ) on seed germination were studied , and ecological significance of light in seed germination was discussed. Under alternating photoperiod (14 h light , 10 h dark , 12μmol m- 2 s - 1 ) , final germination percentage of seeds was 87. 5% , 100% , 100% , 100% , 98% , 89% and 100% , respectively, at 15℃ , 20℃ , 25℃ , 30℃ , 35℃ , 40℃ and 30℃􊄯20℃ , mean germination time of seeds was 34 . 7 d , 16 . 3 d , 5. 6 d , 4 .8 d , 6 . 4 d , 9 d and 6 .3 d , respectively. Germination percentage of seeds which were incubated for 35 d at 15℃ , 20℃ , 25℃ , 30℃ , 35℃ , 40℃ , 30℃􊄯20℃ and dark was zero , and these seeds rapidly germinated after addition of alternating photoperiod . Light intensity of 0 . 5 - 20μmol m- 2 s - 1 markedly increased final germination percentage of seeds . However , different light intensity influencedhardly final germination percentage of seeds, but altered germination rate of seeds . Under photoperiod of 24 h , final germination percentage of seeds notably increased with increasing length of light duration . The seeds treated for 24 h ,36 h , and 48 h by continuous light could germinate at subsequent dark. Different concentrations of exogenous gibberellin, 6-benzyladenine, ethylene, sodium nitroprusside , nitrate and nitrite could not replace light and not accelerate seed germination, and these seeds rapidly gained germinablity after addition of alternating photoperiod . Germination behavior of F. enjamina L . var . nuda seeds was closely related to long-term adaptation for tropical forest .     

Seed mass,hardness, and phylogeny explain the potential for endozoochory by granivorous waterbirds

Field studies have shown that waterbirds, especially members of the Anatidae family, are major vectors of dispersal by endozoochory for a broad range of plants lacking a fleshy fruit, yet whose propagules can survive gut passage. Widely adopted dispersal syndromes ignore this dispersal mechanism, and we currently have little understanding of what traits determine the potential of angiosperms for endozoochory by waterbirds. Results from previous experimental studies have been inconsistent as to how seed traits affect seed survival and retention time in the gut and have failed to control for the influence of plant phylogeny. Using 13 angiosperm species from aquatic and terrestrial habitats representing nine families, we examined the effects of seed size, shape, and hardness on the proportion of seeds surviving gut passage through mallards (Anas platyrhynchos) and their retention time within the gut. We compiled a molecular phylogeny for these species and controlled for the nonindependence of taxa due to common descent in our analyses. Intact seeds from all 13 species were egested, but seed survival was strongly determined by phylogeny and by partial effects of seed mass and hardness (wet load): species with seeds harder than expected from their size, and smaller than expected from their loading, had greater survival. Once phylogeny was controlled for, a positive partial effect of seed roundness on seed survival was also revealed. Species with seeds harder than expected from their size had a longer mean retention time, a result retained after controlling for phylogeny. Our study is the first to demonstrate that seed shape and phylogeny are important predictors of seed survival in the avian gut. Our results demonstrate that the importance of controlling simultaneously for multiple traits and relating single traits (e.g., seed size) alone to seed survival or retention time is not a reliable way to detect important patterns, especially when phylogenetic effects are ignored.