Herbivores equalize the seedling height growth of three dominant tree species in an African tropical rain forest

Determining the impact of insect herbivores on forest tree seedlings and saplings is difficult without experimentation in the field. Moreover, this impact may be heterogeneous in time and space because of seasonal rainfall and canopy disturbances, or ‘gaps’, which can influence both insect abundance and plant performance. In this study we used fine netting to individually protect seedlings of Microberlinia bisulcata, Tetraberlinia bifoliolata and Tetraberlinia korupensis trees (Fabaceae = Leguminosae) from insects in 41 paired gap-understorey locations across 80 ha of primary rain forest (Korup, Cameroon). For all species, growth in height and leaf numbers was negligible in the understorey, where M. bisulcata had the lowest survival after c. 2 years. In gaps, however, all species responded positively with pronounced above-ground growth across seasons. When exposed to herbivores their seedling height growth was similar, but in the absence of herbivores, M. bisulcata significantly outgrew both Tetraberlinia species and matched their leaf numbers. This result suggests that insect herbivores might play an important role in maintaining species coexistence by mitigating sapling abundance of the more palatable M. bisulcata, which in gaps was eaten the most severely. The higher ratio in static leaf damage of control-to-caged M. bisulcata seedlings in gaps than understorey locations was consistent with the Plant Vigour Hypothesis. This result, however, did not apply to either Tetraberlinia species. For M. bisulcata and T. korupensis, but not T. bifoliolata (the most shade-tolerant species), caging improved relative seedling survival in the understory locations compared to gaps, providing restricted support for the Limiting Resource Model. Approximately 2.25 years after treatments were removed, the caged seedlings were taller and had more leaves than controls in all three species, and the effect remained strongest for M. bisulcata. We conclude that in this community the impact of leaf herbivory on seedling growth in gaps is strong for the dominant M. bisulcata, which coupled to a very low shade-tolerance contributes to limiting its regeneration. However, because gaps are common to most forests, insect herbivores may be having impacts upon functionally similar tree species that are also characterized by low sapling recruitment much more widely than currently appreciated. An implication for the restoration and management of M. bisulcata populations in forests outside of Korup is that physical protection from herbivores of new seedlings where the canopy is opened by gaps, or by harvesting, should substantially increase its subcanopy regeneration, and thus, too, its opportunities for adult recruitment.

Effects of seed origin latitude on the timing of height growth cessation and field performance of silver birch

The aim of this thesis was to increase our knowledge about the effects of seed origin on the timing of height growth cessation and field performance of silver birch from different latitudes, with special attention paid to the browsing damage by moose in young birch plantations. The effect of seed origin latitude and sowing time on timing of height growth cessation of first-year seedlings was studied in a greenhouse experiment with seven seed origins (lat. 58º - 67ºN). Variation in critical night length (CNL) for 50 % bud set within two latitudinally distant stands (60º and 67ºN) was studied in three phytotron experiments. Browsing by moose on 5-11 -year-old silver birch saplings from latitudinally different seed origins (53º - 67ºN) was studied in a field experiment in southern Finland. Yield and stem quality of 22-year-old silver birch trees of Baltic, Finnish and Russian origin (54º - 63ºN) and the effect of latitudinal seed transfers were studied in two provenance trials at Tuusula, southern and Viitasaari, central Finland. The timing of height growth cessation depended systematically on latitude of seed origin and sowing date. The more northern the seed origin, the earlier the growth cessation and the shorter the growth period. Later sowing dates delayed growth cessation but also shortened the growth period. The mean CNL of the southern ecotype was longer, 6.3 ± 0.2 h (95 % confidence interval), than that of the northern ecotype, 3.1 ± 0.3 h. Within-ecotype variance of the CNL was higher in the northern ecotype (0.484 h2) than in the southern ecotype (0.150 h2). Browsing by moose decreased with increasing latitude of seed origin and sapling height. Origins transferred from more southern latitudes were more heavily browsed than the more northern native ones. Southern Finnish seed origins produced the highest volume per unit area in central Finland (lat. 63º11'N). Estonian and north Latvian stand seed origins, and the southern Finnish plus tree origins, were the most productive ones in southern Finland (lat. 60º21'N). Latitudinal seed transfer distance had a significant effect on survival, stem volume/ha and proportion of trees with a stem defect. The relationship of both survival and stem volume/ha to the latitudinal seed transfer distance was curvilinear. Volume was increased by transferring seed from ca. 2 degrees of latitude from the south. A longer transfer from the south, and transfer from the north, decreased the yield. The proportion of trees with a stem defect increased linearly in relation to the latitudinal seed transfer distance from the south.

The effect of temperature on height growth of Scots pine in northern Finland

The effect of temperature on height growth of Scots pine in the northern boreal zone in Lapland was studied in two different time scales. Intra-annual growth was monitored in four stands in up to four growing seasons using an approximately biweekly measurement interval. Inter-annual growth was studied using growth records representing seven stands and five geographical locations. All the stands were growing on a dry to semi-dry heath that is a typical site type for pine stands in Finland. The applied methodology is based on applied time-series analysis and multilevel modelling. Intra-annual elongation of the leader shoot correlated with temperature sum accumulation. Height growth ceased when, on average, 41% of the relative temperature sum of the site was achieved (observed minimum and maximum were 38% and 43%). The relative temperature sum was calculated by dividing the actual temperature sum by the long-term mean of the total annual temperature sum for the site. Our results suggest that annual height growth ceases when a location-specific temperature sum threshold is attained. The positive effect of the mean July temperature of the previous year on annual height increment proved to be very strong at high latitudes. The mean November temperature of the year before the previous had a statistically significantly effect on height increment in the three northernmost stands. The effect of mean monthly precipitation on annual height growth was statistically insignificant. There was a non-linear dependence between length and needle density of annual shoots. Exceptionally low height growth results in high needle-density, but the effect is weaker in years of average or good height growth. Radial growth and next year s height growth are both largely controlled by current July temperature. Nevertheless, their growth variation in terms of minimum and maximum is not necessarily strongly correlated. This is partly because height growth is more sensitive to changes in temperature. In addition, the actual effective temperature period is not exactly the same for these two growth components. Yet, there is a long-term balance that was also statistically distinguishable; radial growth correlated significantly with height growth with a lag of 2 years. Temperature periods shorter than a month are more effective variables than mean monthly values, but the improvement is on the scale of modest to good when applying Julian days or growing-degree-days as pointers.

Nutrient enrichment effects on mangrove sediments from differing tree height zones in Bocas del Toro, Panama

Red mangrove (Rhizophora mangle L.) forests have distinct tree-height zones, with tall trees fringing the ocean and shorter trees in interior stands. A long-term nitrogen (N) and phosphorus (P) fertilization experiment in Almirante Bay, Bocas del Toro Province, Panama has shown that tree-height zonation is primarily related to nutrient limitation. This experiment was used to test the effects of in-situ nutrient additions and tree zonation on mangrove sediments. The sediments underlying the experimental R. mangle trees were sampled and N₂ fixation, ¹⁵N, chlorophyll a, percent N and P, and percent organic biomass were quantified. Both N and P additions significantly affected almost every parameter measured in both zones within this experiment. These results are likely to have implications for management since N and P inputs are predicted to increase throughout the tropics and subtropics worldwide.

Tree height integrated into pantropical forest biomass estimates

Aboveground tropical tree biomass and carbon storage estimates commonly ignore tree height (H). We estimate the effect of incorporating H on tropics-wide forest biomass estimates in 327 plots across four continents using 42 656 H and diameter measurements and harvested trees from 20 sites to answer the following questions: 1. What is the best H-model form and geographic unit to include in biomass models to minimise site-level uncertainty in estimates of destructive biomass? 2. To what extent does including H estimates derived in (1) reduce uncertainty in biomass estimates across all 327 plots? 3. What effect does accounting for H have on plot- and continental-scale forest biomass estimates? The mean relative error in biomass estimates of destructively harvested trees when including H (mean 0.06), was half that when excluding H (mean 0.13). Power- and Weibull-H models provided the greatest reduction in uncertainty, with regional Weibull-H models preferred because they reduce uncertainty in smaller-diameter classes (< 40 cm D) that store about one-third of biomass per hectare in most forests. Propagating the relationships from destructively harvested tree biomass to each of the 327 plots from across the tropics shows that including H reduces errors from 41.8 Mg ha(-1) (range 6.6 to 112.4) to 8.0 Mg ha(-1) (-2.5 to 23.0).

Tree height integrated into pantropical forest biomass estimates

Aboveground tropical tree biomass and carbon storage estimates commonly ignore tree height (H). We estimate the effect of incorporating H on tropics-wide forest biomass estimates in 327 plots across four continents using 42 656 H and diameter measurements and harvested trees from 20 sites to answer the following questions: 1. What is the best H-model form and geographic unit to include in biomass models to minimise site-level uncertainty in estimates of destructive biomass? 2. To what extent does including H estimates derived in (1) reduce uncertainty in biomass estimates across all 327 plots? 3. What effect does accounting for H have on plot- and continental-scale forest biomass estimates? The mean relative error in biomass estimates of destructively harvested trees when including H (mean 0.06), was half that when excluding H (mean 0.13). Power- andWeibull-H models provided the greatest reduction in uncertainty, with regional Weibull-H models preferred because they reduce uncertainty in smaller-diameter classes (?40 cm D) that store about one-third of biomass per hectare in most forests. Propagating the relationships from destructively harvested tree biomass to each of the 327 plots from across the tropics shows that including H reduces errors from 41.8Mgha?1 (range 6.6 to 112.4) to 8.0Mgha?1 (?2.5 to 23.0). For all plots, aboveground live biomass was ?52.2 Mgha?1 (?82.0 to ?20.3 bootstrapped 95%CI), or 13%, lower when including H estimates, with the greatest relative reductions in estimated biomass in forests of the Brazilian Shield, east Africa, and Australia, and relatively little change in the Guiana Shield, central Africa and southeast Asia. Appreciably different stand structure was observed among regions across the tropical continents, with some storing significantly more biomass in small diameter stems, which affects selection of the best height models to reduce uncertainty and biomass reductions due to H. After accounting for variation in H, total biomass per hectare is greatest in Australia, the Guiana Shield, Asia, central and east Africa, and lowest in eastcentral Amazonia, W. Africa, W. Amazonia, and the Brazilian Shield (descending order). Thus, if tropical forests span 1668 million km2 and store 285 Pg C (estimate including H), then applying our regional relationships implies that carbon storage is overestimated by 35 PgC (31?39 bootstrapped 95%CI) if H is ignored, assuming that the sampled plots are an unbiased statistical representation of all tropical forest in terms of biomass and height factors. Our results show that tree H is an important allometric factor that needs to be included in future forest biomass estimates to reduce error in estimates of tropical carbon stocks and emissions due to deforestation.

Tree height reduction for parallel processing of blocks of Fortran assignment statements.

Thesis (M.S.)--University of Illinois.

Jay Sah measuring tree height in a mangrove forest impacted by Hurricane Wilma, Harney River

This material is based upon work supported by the National Science Foundation through the Florida Coastal Everglades Long-Term Ecological Research program under Cooperative Agreements #DBI-0620409 and #DEB-9910514. This image is made available for non-commercial or educational use only.

Left to right: Mike Ross and Nilesh Timilsina measuring mangrove tree height after Hurricane Wilma, Harney River

This material is based upon work supported by the National Science Foundation through the Florida Coastal Everglades Long-Term Ecological Research program under Cooperative Agreements #DBI-0620409 and #DEB-9910514. This image is made available for non-commercial or educational use only.

Nilesh Timilsina measuring tree height in a mangrove forest impacted by Hurricane Wilma, Harney River

This material is based upon work supported by the National Science Foundation through the Florida Coastal Everglades Long-Term Ecological Research program under Cooperative Agreements #DBI-0620409 and #DEB-9910514. This image is made available for non-commercial or educational use only.

Preliminary evaluation of macadamia rootstocks for yield and tree height

Rootstock has profound effects on traits such as yield and tree size in various horticultural industries, however relatively little is known about rootstock effects for macadamia. In this study, 12 cultivars were propagated as open-pollinated seedling and clonal rootstocks, and own-rooted cuttings. The same cultivars were also used as scions, and grafted to a subset of rootstocks, then planted at four trial locations. In this preliminary analysis, rootstock accounted for 19% of the variance in yield compared with 72% for scion, and 23% in height compared with 72% for scion. There was no interaction between rootstock and scion for yield, and only a small effect for height. The interaction between rootstock and propagation method (seedling, clonal, own roots) was not significant for height. A small effect was observed for yield, with the own roots treatment producing significantly lower yield than grafted trees for all rootstock cultivars except 'HAES 849'. 'H2' seedling rootstock produced a cumulative yield to age 10 years of 11.1 kg tree -1 compared to the highest yield of 13.6 kg tree -1 for 'Beaumont' clonal rootstocks. 'H2' seedling rootstock produced 4.8 m trees at age 11 years, compared to the smallest grafted tree which was 'HAES 849' seedling at 4.7 m.

华山松（Pinus armandi）地理变异的生态生理学研究

华山松(Pinus armandi)是我国西部高海拔温凉山区的主要树种，自然分布大约在北纬23° 30'-36°30'，东经88°50'-113°30'之间，横跨13个省（区）包括两个季风区，六个气候带。由于水平空间区域广阔，分布区中间地带又为川藏高原的高山峡谷区，群体疏散，基因的迁移和交流困难，致使南北群体出现深刻的遗传分化，并在形态上、生理上充分表达出来。研究结果表明: 1)华山松树高生长出现明显的地理渐变模式，在同一适宜的立地条件下，15年生的南方群体的树高会超出北方种源的2-3倍。针叶长度也同样具有明显的南北差异，南方群体较长。 2)地理因子相关分析表明纬度因子是主导因子，根据种源的地理坐标，利用中科院植物所植被数量生态学开放室有关数据库和生态信息系统(EIS)进行有关气候数据统计分析表明大多数环境气候因子主要是由纬度决定的。在环境因子当中，年平均温度和极端最低气温，在华山松地理变异过程中起主要作用。利用聚类分析方法，可把华山松大致分为南北两个地理型，正好与气候因子聚类而成的两个环境类型相对应，反映着两个自然地理区的自然生态因子的综合区别。 3)利用已有的幼林生长数据和我们在华山松进入杆材期(15年生)的树高调查结果，作相关分析，发现二者相关关系显著。因此，利用早期树高作生长预测在林木选育上是可行的。 4)根据六个种源测定结果，发现南方温暖湿润气候条件下华山松种子较大，种壳较薄，而北方种源种子相对较小，种皮要厚一些。 5)虽然种群之间中性盐及碱性溶液提取的种子蛋白含量差异明显，但没有地理变异趋势。分析多肽组成，中性盐溶种子蛋白也没有地理特征。但碱性溶液提取的种子蛋白多肽组成却有明显地域性。其中36KD多肽仅出现在北方种群中，南方种群则没发现有同样多肽。 6)西藏种源表现出了一种独特的生物学特性，其种子较大有南方种源特征，但种皮厚又带有北方种源的特点，除此之外，它的碱性溶液提取的种子蛋白中含有36KD多肽，因此它可能是南北两个类群中的中间类型。 7)用活体针叶作测材料时，没能检测出光谱特性的地理变异。但改用分离的叶绿体作试验材料，发现南方种群的光能吸收4阶导数光谱在680nm处的峰值较大，670nm峰值较小，而北方种群中出现了670nm峰值较大的类群，推测北方种群反应中心活力有下降趋势。另外，低温荧光发射光谱及低温荧光激发光谱也有明显的地域分化，表明体内色素状态有一定的地理变异。 8)时绿素a荧光诱导动力学测定表明，南方种群可变荧光(Fv/Fo)与光能转换效率(Fv/Fm)较高，北方种群则相对要小一些，与四阶导数吸收光谱测定结果吻合．利用饱和脉冲重复激发技术，发现光化学荧光淬灭(Qp)和非光化学荧光淬灭(Qn)在北方种群中较大，南方种群较小，其中，Qn变化非常明显，而Qp变化较小． 综合研究结果，除树高生长．针叶和种子形态外，36KD碱溶性种子蛋白多肽680nm处4阶导数吸收光谱峰值大小，可变荧光强度(Fv/Fo)光能转换效率(Fv/Fm)以及非化学荧光淬灭系数都带有明显的地理特征，与华山松生长和适应能力紧密相关，可作为生理指标应用于生态地理型的区分或华山松的早期选择。

Growth performance and management of a mixed rainforest tree plantation

Monoculture plantations of Pinus, Eucalyptus and Acacia have been established oil rainforest lands throughout the world. However, this type of reforestation generally supplies low quality timber and contributes to landscape simplification. Alternatives to exotic monoculture plantations are now beginning to gain momentum with farmers and landholders attempting to establish a variety of rainforest trees in small plantations. When compared to the well studied commercial species, knowledge concerning the growth and management of many of these rainforest species is in its infancy. To help expand this limited knowledge base an experimental plantation of 16 rainforest tree species in a randomised design was established near Mt. Mee, in south-eastern Queensland, Australia. Changes in growth, form (based on stem straightness, branch size and branchiness), crown diameters and leaf area of each species were examined over 5 years. Patterns of height growth were also measured monthly for 31 months. Species in this trial could be separated into three groups based on their overall growth after 5 years and their growth patterns. Early successional status, low timber density, high maximum photosynthetic rates and large total leaf areas were generally correlated to rapid height growth. Several species (including Araucaria cunninghamii, Elaeocarpus grandis, Flindersia brayleyana, Grevillea robusta and Khaya nyasica) had above average form and growth, while all species in the trial had considerable potential to have increased productivity through tree selection. As canopy closure occurred at the site between years four and five, growth increments declined. To reduce stand competition a number of different thinning techniques could be employed. However, simple geometric or productivity based thinnings appear to be inappropriate management techniques for this mixed species stand as they would either remove many of the best performing trees or nearly half the species in the trial. Alternatively, a form based thinning would maintain the site's diversity, increase the average form of the plantation and provide some productivity benefits.