Responses of Pinus resinosa in Newfoundland to wildfire

Abstract. Natural Pinus resinosa (red pine) stands in Newfoundland are restricted to 22 small, dry, nutrient-poor sites. A short wildfire cycle (15 - 30 yr) of both surface and crown fire regulates stand perimeters and is the main factor in regulating stand development. At the nucleus of current stands < 100 yr old, a few trees > 200 yr occur, usually showing multiple fire scars. Stem char heights confirmed an increased flammability of the stand and tree mortality for fires moving in the upslope position, as well as for mixed Pinus resinosa-Picea mariana stands. All P. resinosa stands are severely nutrient-deficient. Leaf concentrations of N, P and K were below or near the reported critical values. Nutrient concentrations were highest three months after a surface fire, but dropped considerably one year later. A gradual increase to near post-fire levels is achieved four years after fire. Foliar nutrient concentrations were positively correlated with average width of the annual rings. Aspects of the ericaceous understory dynamics and its relation to P. resinosa regeneration are also discussed.     

Possible role of disturbance in shaping the northern distribution of Pinus resinosa

Abstract. The objective of this study is to determine the factors responsible for the distribution of Pinus resinosa (red pine) at its northern limit in northwestern Québec. Pinus resinosa is found only on islands and protected lake shores at its northern distribution boundary. The influence of climate on the germination of P. resinosa seed and on the phenology of P. resinosa was investigated in the Lake Duparquet region of northwestern Québec. The results indicate that P. resinosa seed readily germinates at island- and inland sites, which indicates that germination is not responsible for the distribution of P. resinosa in the Lake Duparquet region. Also, cones and seeds developed normally at island sites and an inland plantation, which suggests that seed production is not hindering the expansion of P. resinosa. These results, together with other studies in the literature, suggest that no climatic factor could explain the present distribution of P. resinosa nor its northern limit. The fire regime appears to be responsible for the restriction of P. resinosa to lake environments and hinders the northward expansion of P. resinosa. The typical crown fire regime of the boreal forest is not conducive for P. resinosa regeneration and restricts the species to fire-sheltered locations such as islands and protected lake shores.     

Burial of canopy-stored seeds in the annual psammophyte Agriophyllum squarrosum Moq. (Chenopodiaceae) and its ecological significance

Agriophyllum squarrosum Moq. is a dominant annual on sand dunes in the arid regions of central Asia. A high percentage of seeds is retained on dead plants which become covered by moving sand, but little is known about the ecological significance of burial of canopy-stored seeds. We investigated the size and dynamics of the buried canopy-stored seed bank and effects of burial on seed germination. In March (during the windy season), May (beginning of the germination season), and July (middle of the growing season), the number of seeds per square meter in sample plots in the dunes was 623, 223 and 22, respectively, with 54.6, 30.6 and 12.9% of the total seeds retained on buried plant canopies. In a controlled experiment, more seedlings emerged from released (dispersed) than from canopy-stored seeds when burial depth was the same. No viable ungerminated released seeds were found, but 45–80% of the ungerminated canopy-stored seeds were viable. In general, with an increase in applied water germination of released seeds buried at a depth of 1 or 2 cm and of canopy-stored seeds buried at 1 cm increased, but regardless of watering regime few or no released seeds at 4 cm or canopy-stored seeds at 2 or 4 cm germinated. Significantly more seedlings emerged from plants buried in a horizontal than in a vertical position. Seedlings originating from buried canopy-stored seeds on an active dune accounted for only 5.4% of the total seedlings emerging, and most of them emerged later than those from released seeds. Thus, seed release is more effectively postponed in buried than in exposed canopies, and burial of canopy-stored seeds is a mechanism that helps regulate seed germination and seedling emergence of A. squarrosum on active dunes.     

Comparison of selection methods for optimizing genetic gain and gene diversity in a red pine (<Emphasis Type="Italic">Pinus resinosa</Emphasis> Ait.) seedling seed orchard

Four selection methods, individual selection (IS), family selection (FS), family plus within-family selection (FWFS) and combined selection (CS), were used to estimate genetic gain [E(g)] for stem volume and gene diversity (GD) for ten theoretical selection intensities in a 108-family, 12-year-old red pine seedling seed orchard. Estimated genetic gain for stem volume ranged from 4.6% to 11.8% across all selection methods and intensities with CS consistently having the highest gains and FS the lowest for any given selection intensity. Genetic diversity ranged from 0.9797 to 0.9954 across all selection methods and intensities. Individual selection was the best selection method for retaining GD, especially at the higher selection intensities, while FWFS was more efficient at the lowest selection intensity. An optimization point, which maximized E(g) and GD relative to each other, was calculated for each selection method. In all cases the optimization point indicated that both E(g) and GD would be favorably high when optimized relative to each other. The implications for volume gain, genetic diversity and potential inbreeding in red pine, a species with inherently low levels of genetic variation, are discussed.Communicated by D.B. Neale     

Comparative studies of ectomycorrhiza formation in Alnus glutinosa and Pinus resinosa with Paxillus involutus

 Mycorrhiza ontogeny and details of Hartig net and mantle structure were compared in ectomycorrhizas synthesized in growth pouches between the broad host range fungus Paxillus involutus and the tree species European black alder (Alnus glutinosa) and red pine (Pinus resinosa). In Alnus glutinosa, a paraepidermal Hartig net was restricted to the proximal (basal) portion of first-order laterals; the hypodermal layer appeared to be a barrier to fungal penetration. Phi-thickenings were present in some cortical cells but these were not related to lack of fungal ingress into the cortex. The mantle was often present close to the root apex but in many roots it was loosely organized and patchy. In several instances, the mantle formed around the root apex was only temporary; renewed root growth occurred without the formation of a mantle. In Pinus resinosa, the Hartig net developed between cortical cell layers of monopodial and dichotomously branched first–order laterals. Fungal hyphae in the Hartig net exhibited a complex labyrinthine mode of growth. The mantle had a pseudoparenchymatous structure and covered the root, including apices of dichotomously branched roots. The Paxillus–Pinus resinosa interaction had all the characteristics of a compatible ectomycorrhizal association. The Paxillus–Alnus glutinosa interaction, however, showed only aspects of superficial ectomycorrhizas, including the presence of a minimal (sometimes absent) and mostly proximal Hartig net and variable mantle development. Sclerotia were produced in the extraradical mycelium of Paxillus involutus when associated with either Alnus glutinosa or Pinus resinosa. Accepted: 22 October 1998     

浙江天童灌丛植被的土壤种子库与幼苗库特征

采用野外样点取样和萌发实验相结合的办法,研究浙江天童灌丛植被的幼苗库和种子库。结果表明：种子库中优势种类主要是淡竹叶、芒、苔草等草本植物,其次为山矾、Ji木、白栎等灌木种类。在厚度为10cm的表层土壤中,枯枝落叶层、0．2cm层、2－4cm、4－10cm层的种子密度分别为125．0粒／m^2、134．3粒／m^2、84．6粒／m^2、62．7粒/m^2。幼苗库的优势种类不太集中,数量较多的种类有苦槠、紫金牛、淡竹叶等。     

Application of the chloroform fumigation-incubation method to the estimation of soil microbial biomass in burned and unburned Japanese red pine forests

Abstract Estimation of soil microbial biomass in burned and unburned Japanese red pine forests was attempted using the chloroform fumigation-incubation method. As the amount of CO₂-C evolved from the fumigated soil for 10–20 days after fumigation (designated as F') was always lower than that from the unfumigated soil during the same period (UF'), the formula, microbial biomass-C(M) = the amount of CO₂-C evolved from the fumigated soil for 0–10 days after fumigation, F) − F'/ k _c, was proposed instead of Jenkinson's conventional formula, M = (F − UF')/ k _c. The k _c value was also determined as 0.30 using 3 fungal and 3 bacterial cultured species as internal standards. Microbial biomass-C calculated by (F − F')/0.30 decreased with soil depth at both the burned (Nenoura, 3.5 years after fire) and unburned (Ato) sites, showing the significant correlation with the decrease of soil respiration and organic C content along soil depth. Microbial biomass-C in the 0–2 cm soil layer at the burned site at Nenoura was 130 mg/100 g dry soil and those in the HF horizon and 0–2 cm soil layer at the unburned site at Ato were 686 and 146 mg/100 g dry soil, respectively.     

Comparative seed biology and co-existence of two fynbos shrub species

Abstract. Fire-prone fynbos communities include numerous species which are difficult to distinguish in terms of conventional niche axes since they are morphologically similar and share the same broad life-history traits. These species compete for space after each fire and lottery theory predicts that those with dissimilar per capita regeneration would not co-exist in the long-term. We studied seed production, soil seed bank dynamics, germination and post-fire seedling establishment and mortality of two obligate reseeding, morphologically similar shrubs which co-occur in dune fynbos throughout the southwestern Cape. Passerina paleacea produced 30 x more plump seeds and maintained soil seed banks between 1.5 x (pre-dispersal) and 2.6 x (post-dispersal) times larger than Phylica ericoides. Seeds of both species showed a high degree of dormancy although germination was stimulated by direct fire-related cues for Phylica only. Approximately 4 x more Passerina seedlings than Phylica seedlings were counted at the end of the winter germination period after a mild autumn fire. However, seedling mortality in the first five months of the dry summer was more than twice as high for Passerina than for Phylica. We suggest that the long-term co-existence of these two species does not result from similar post-fire regeneration success but rather from population instabilities arising from differential regeneration in relation to fire regime. The results predict that hot fires followed by dry summers would favour Phylica over Passerina as the former has fire-stimulated germination and greater drought tolerance; cool fires followed by moist summers would favour Passerina.     

The effects of Sasa and canopy gap formation on tree regeneration in an old beech forest

Abstract. We examined the response of tree seedling emergence and survival to the dieback of Sasa and canopy gap formation in an old‐growth forest near Lake Towada, northern Japan. Synchronous death of Sasa occurred in 1995. We established four types of sampling sites differing in forest canopy conditions (Closed or Gap) and Sasa status (Dead or Live). Gap‐Dead sites had the highest light levels and the greatest fluctuation in soil temperatures. The death of Sasa alone facilitated the emergence (Acer japonicum, Fagus crenata, Fraxinus lanuginosa, and Tilia japonica) and survival (Acanthopanax sciadophylloides, F. crenata, F. lanuginosa, Kalopanax pictus, and Sorbus commixta) of species with a seedling bank strategy. Cercidiphyllum japonicum grew at all sites at a higher density than other species, but survived well only in Gap‐Dead sites. This behaviour was associated with a seed rain strategy. The additive effects of Sasa death and canopy gap formation promoted seedling emergence of pioneer tree species (Betula maximowicziana, Lindera umbellata, and Magnolia obovata), probably through break of dormancy by the large temperature fluctuation. In addition, the scarcity of advance regeneration in canopy gaps due to Sasa cover facilitates the regeneration of pioneer species. The dominance and dieback cycle of Sasa contributes to species diversity in this forest.     

西天山野果林不同居群黑果小檗土壤种子库及幼苗更新研究

通过野外调查及采集土样,对新疆西天山野果林内黑果小檗(Berberis atrocarpa Schneid.)3个自然居群(霍城、新源和特克斯)的土壤种子库及幼苗更新情况进行研究。结果显示,3个居群内黑果小檗土壤种子质量无显著差异,霍城、新源和特克斯居群黑果小檗土壤种子库中完整种子分别占种子总数的51.6%、49.4%和54.1%。霍城、新源和特克斯居群黑果小檗完整种子均聚集在枯枝落叶层,分别为98.4%、97.4%和100%,3个居群5~10 cm土层均未发现黑果小檗种子。土壤种子水平扩散距离随着坡度的增加逐渐增大。霍城、新源和特克斯居群黑果小檗幼苗均为根蘖苗,根蘖繁殖可使幼苗扩散到母株周围100 cm范围或更远处。Ⅱ级苗(20~40 cm)数量最多,占幼苗总数的50%;3个居群黑果小檗的Ⅰ级苗(0~20 cm)向Ⅱ级苗的转化率均为100%;霍城和新源居群Ⅱ级、Ⅲ级(40~60 cm)苗间的转化率较低,分别为29%和32%,特克斯居群Ⅱ级、Ⅲ级苗间的转化率较高,为78%。研究结果表明西天山野果林黑果小檗种子易滞留于地表,难以进入土壤深层且土壤中种子霉变率高,这可能是黑果小檗自然更新困难的原因。     

Composition and seasonal dynamics of seed rain in broad-leaved Korean pine (Pinus koraiensis) mixed forest in Changbai Mountain, China

To explore the composition and spatio-temporal dynamics of seed rain in broad-leaved Korean pine (Pinus koraiensis) mixed forest, 150 seed traps were set up in a 25 hm2 plot in Changbai Mountain. Seeds, fruits, anthotaxy and others in seed traps were collected, identified and divided into 4 types. From 2005 to 2006, we collected 47 different types. Total number of seeds and fruits was 121291, including 23147 mature seeds and fruits (19.1% of the total). Tilia amurensis and Fraxinus mandshurica, with the most seeds and fruits, accounted for 90% of the total. The analysis on seasonal dynamics of seed rain showed that there were the largest number of seeds and fruits between July and October, which were composed of immature seeds and fruits. In mid-October, mature seeds and fruits reached their peak, but immature seeds and fruits still accounted for high proportion. There were 91 traps that contained 100–200 mature seeds and fruits, and one trap without any mature seed or fruit. The largest number of species found in a trap was 7, and usually 3 or 4 species were found in most of the traps. There were obvious relationships between spatial patterns of mature seeds and fruits and their parent trees, indicating that their mature seeds and fruits were not dispersed far from their parent trees.     

长白山阔叶红松(Pinus koraiensis)林种子雨组成及其季节动态

为了解阔叶红松(Pinus koraiensis)林的种子雨组成及其在时间和空间上的动态变化,在长白山阔叶红松林25 hm2样地内,设置了150个种子收集器,收集掉落于种子收集器中的果实、种子等.所有收集到的样品分别鉴定其种类并分为成熟种实、未成熟种实、花序和果实或种子碎片及其附属物4类,计算各类别的数量,而后分别烘干秤重.从2005年6月到2006年11月,共收集到隶属11科12属20种的种实及其生殖器官残体.累计收集到121291粒种实,其中成熟种实23147粒,仅占所有种实总个体数的19.1%.种实数最多的树种是紫椴(Tilia amurensis)和水曲柳(Fraxinus mandshurica),两个树种种实的个体数占总个体数的90%.对2006年5月～11月种子雨季节动态的分析发现:种实在7月中旬至10月下旬数量极大,但主要由未成熟种实组成;在10月中旬出现成熟种实散落高峰,但未成熟种实仍占一定的比例.按每个收集器收集到的成熟种实数统计,成熟种实数量在100～200之间的收集器数量最多.按每个收集器收集到的树种数统计,收集器中最多收到的树种数为7,树种数为3和4的收集器个数最多.对6个主要树种成熟种实所在收集器的空间分布进行分析发现,紫椴和水曲柳的成熟种实在整个样地都有分布,春榆(Ulmus japonica)、糠椴(T. mandshurica)、色木槭(Acer mono)和假色槭(A. pseudo-sieboldianum)则只在样地的局部区域收集到成熟种实.成熟种实的空间分布与母树的空间分布大都表现出明显的相关性,表明这些树种的成熟种实并没有扩散到离母树较远的距离.     

Altitudinal gradients in seed dynamics of Calluna vulgaris in eastern Scotland

Abstract. Relationships between seed deposition, size of soil seed banks and some of the environmental factors affecting them were assessed for Calluna vulgaris throughout its altitudinal range (150–960 m a.s.l.) in eastern Scotland. Seed rain was assessed using pitfall traps, collected every 5–10 wk for 3 yr; germinability was determined by laboratory incubation. Seed bank size was estimated, once only, by counting seedlings emerging from soil cores kept for 50 wk in a glasshouse. Seed deposition varied annually, was related to parent plant cover but always declined with altitude, falling sharply above 600 m a.s.l. Seed bank size was more closely correlated with the proportion of organic matter in the soil than with the amount of seed rain. Seed bank sizes declined gradually with altitude but did not differ significantly between four altitudinal zones. The mean density of buried seeds was less than half the mean annual seed rain at sites below 300 m a.s.l. but was over 200 × greater than annual seed rain above 800 m, suggesting that seeds buried at high altitudes remain viable for much longer than those at lower altitudes.     

Early post-fire regeneration of a Pinus halepensis forest on Mount Párnis,Greece

Abstract. The post-fire regeneration of a 45-yr-old Pinus halepensis (Aleppo pine) forest, burned in July 1989, has been studied on Mount Párnis, Attiki, Greece. Four experimental plots at various slopes and exposures were established at altitudes of 400 - 450 m, and monitored for 3 yr at 3-month intervals. Early regeneration took place abundantly, through both resprouting and seed germination of mostly hard-seeded herbs and shrubs; the floristic richness was high with 80 taxa. Pine seedling emergence took place during the winter of the first post-fire year. The mean pine seedling density by the end of the recruitment period (March 1990) was 5–6 seedlings/m². This density decreased slightly during late spring and considerably during summer. During the second post-fire year only a relatively slight decline was observed; thereafter the density was stabilized to 1 - 2 seedlings/m². Mortality follows a negative exponential curve that levels off at ca. 20 %. Height distributions throughout the three post-fire years were all positively skewed as a result of the presence of few very tall saplings. A considerable fraction (20 %) of very short (5–15 cm) saplings were still alive 39 months after the fire; these may constitute the sapling bank. Based on the analysis of height distribution curves, it is concluded that the taller seedlings survived significantly better than the shorter ones.     

Internode length in Pisum. I. The effect of the Le/le gene difference on endogenous gibberellin-like substances

The allelopathic potential of the dry fruits of Washingtonia filifera (L. Linden) H. Wendl. was investigated. Leachates from fruits inhibited the germination of lettuce, wheat, red cabbage and cucumber seeds. The inhibitory effect was partly neutralized by kinetin (20 mg 1⁻¹) and gibberellic acid (50 mg 1⁻¹). The effect of kinetin was more pronounced at 25°C than at 20°C. Substances inhibiting germination were localized in the pericarp of the fruit and were resistant to high temperature.     

Stimulation of seed germination in an abandoned hay meadow

Abstract. In order to stimulate germination from the soil seed bank in a formerly species-rich hay meadow for conservation purposes, eight gaps were made in each of four habitat types: (1) Dry/open, (2) Dry/shaded, (3) Moist/open and (4) Moist/ shaded; the meadow was abandoned 12 yr before the experiment started. The mean tree cover of the gaps was reduced from 50 % to 22 %. Light was estimated as distance to nearest tree towards south and shade as tree cover. Soil moisture content was estimated from topography and measured from soil samples. There were no significant differences in total herb number between strata before the experiment started. The number of the target group herb indicators of old, traditional management, was however higher in dry, open areas than in shaded areas whether moist or dry. After the treatment, 15 herbs appeared anew in the gaps, nine of these were indicators of traditional management. 80 % of the indicators present after treatment were either new or had increased in frequency in the gaps. More indicators of traditional management appeared in gaps in the open, on dry soils in slopes than in shaded gaps, on moist soils and flat surface. In order to stimulate germination from the seed bank of as many herb indicators of traditional management as possible in abandoned hay meadows of the region, dry, sloping areas, situated as far as possible from large trees towards south and in the open, should be chosen for making gaps for germination.     

Role of ethylene and auxin in regenerative differentiation and orientation of tracheids in Pinus pinea seedlings

A low auxin concentration (0.1% naphthalene acetic acid) induced tracheids with longitudinal polarity parallel to the hypocotyl axis in young Pinus pinea seedlings. Application of 0.1% ethrel laterally and 0.1% naphthalene acetic acid apically disturbed axial tracheid polarity and promoted the differentiation of tracheids with a lateral orientation. Ethrel by itself, with no auxin background, did not affect tracheid differentiation. Apical application of 1% gibberellic acid with the low auxin, reversed the polarity disorder induced by ethrel. Disturbance of axial tracheid polarity was observed under a high auxin concentration (0.5% naphthalene acetic acid) which was similar to the combined effect of ethrel and auxin. The high auxin concentration increased tracheid number significantly. This effect was curtailed following treatment with inhibitors of ethylene formation (Co²⁺; 1-aminoethoxy- vinylglycine) and action (Ag²⁺). The role of ethylene in controlling the differentiation of radial tracheids, which characterize the vascular rays of pines, is discussed.     

Seed bank classification and its importance for the restoration of species-rich flood-meadows

Abstract. This paper focuses on the persistence of seeds in the seed bank of a formerly fertilized flood-meadow into which seeds from an original Alopecurus pratensis-Sanguisorba officinalis community have been introduced. The longevity of seeds in the seed bank of the original community is also estimated. The established vegetation was compared with the seed bank, as divided into two layers (0 - 5 cm and 5–10 cm); this allowed a classification of species into three groups, with (1) transient, (2) short-term persistent and (3) long-term persistent seed bank. The majority of the species of the unfertilized flood-meadow community have a transient or short-term persistent seed bank with seeds showing a large variance in shape i.e. flattened or elongate. Because of this, the characteristic flood-meadow species will disappear soon after the beginning of fertilizer application and will not re-establish from the seed bank, once the fertilizer application is ceased. The formerly fertilized flood-meadow contains many ruderal and arable weed species, the seeds of which tend to be compact or round. Significantly more seeds were found in the seed bank of the formerly fertilized flood-meadow under cattle-grazing than under sheep-grazing and no grazing.