Light gradient partitioning by tropical tree seedlings in the absence of canopy gaps

To explore the importance of light availability for seedling growth in low light environments, we examined light-dependent growth, biomass allocation and mortality of tree seedlings growing in sites with 0.2-6.5% full sun, the range of light commonly encountered in the understory of closed canopy, lowland tropical forests. We transplanted seedlings of the canopy tree species, Dipteryx panamensis, Virola koschnyii, and Brosimum alicastrum into second-growth forest and native tree plantations at La Selva Biological Station, Costa Rica. We assessed seedling survival, growth, and seedling light environments bimonthly for 14 months. Plants were harvested at the end of the study to assess leaf area, total biomass, biomass partitioning and root growth. Survivorship of all species exceeded 60% across all microsites, although both D. panamensis and B. alicastrum had lower probabilities of survival in the darkest microsites. All species showed a strong positive relationship between light availability and growth, increasing in total biomass as light increased. However, the strength of the growth response differed among species causing a change in the rank order of species growth rates as light availability increased. Although D. panamensis showed the lowest growth rates in the darkest microsites, a strong response to increasing light led to a cross-over in performance, such that D. panamensis had the highest growth rate at the highest light levels studied. These data suggest that resource gradient partitioning could occur even in low light environments (0.2-6.5%). Given the limited range of light regimes sampled (i.e., non-gap microsites), our data demonstrate that growth of tropical tree seedlings beneath closed canopies is highly sensitive to light availability and that shade-tolerant species vary in these responses. Our results show that understory light heterogeneity, in the absence of canopy gaps, can significantly affect recruitment processes for shade-tolerant tree species.     

吉林东部天然林林下光质分布特征

天然林林下光质对乔木幼苗以及灌草的组成与更新具有重要的生态学意义。但目前对于林下光质的研究仍然有限。以吉林东部地区天然林为例,通过调查乔木数据和林下光质数据,基于移动窗口法分析不同空间尺度森林冠层结构与林下光质的关系。结果表明：不同林型下红光光子通量密度(R)与蓝光光子通量密度(B)存在差异。其中沙松-千金榆-花楷槭混交林林下蓝光光子通量密度最小,而沙松-紫椴-臭冷杉混交林和长白落叶松纯林林下最大。随着尺度的增大,天然林乔木胸高断面积与R/PFD(红光/光子通量密度比值)和B/PFD(蓝光/光子通量密度比值)的比值呈显著正相关(P<0.05)。并且随着尺度的增加,相关系数总体逐渐增大,在35m处达到峰值。在此基础上在南向、东向和西向各延伸10m时呈现显著正相关(P<0.05)。在该尺度下分析优势树种对林下R/PFD和B/PFD比值的影响时发现：R/PFD与B/PFD比值随着针叶林胸高断面积的增加而增加。相对于阔叶林来说,多数林型针叶林下的冠层结构与林下R/PFD和B/PFD比值之间显著正相关(P<0.05)。在不同树种下,乔木冠层结构对R/PFD和B/PFD比值的影响...     

Microenvironmental variability and species diversity in treefall gaps in a sub-tropical broadleaved forest

Microenvironmental variability and species diversity in gaps and forest understorey were studied to assess the role of treefall gaps in maintaining composition and patchy distribution in a broad-leaved sub-tropical climax forest, Mawphlang, Meghalaya, India. Photon flux density was higher in gaps than in the surrounding understorey. Relative humidity was low and the litter layer was relatively thin in gaps throughout the year. Soil moisture and photon flux density in the gaps significantly varied between seasons and gaps of different sizes. Relative humidity significantly varied between seasons but difference among gaps was insignificant. Among-gap and among-season variations in soil and air temperature were insignificant.The number of tree species in the gaps was positively correlated with gap area, and tree species abundance showed higher equitability in larger than in smaller gaps. In gaps, -diversity was highest for herbs and lowest for shrubs. -diversity was highest for shrubs and lowest for tree seedlings. -diversity of tree seedlings was higher in the gaps than in the forest understorey. Conversely, -diversity was higher in the understorey than in the gaps. Low species similarity for tree seedlings among the gaps could be an effect of patchy distribution of parent tree species in the forest. Thus a significant change in light and moisture regimes along the gap size gradient played an important role in influencing the composition and abundance of shade tolerant and intolerant tree species in gaps on one hand, and affected the overall species diversity of the forest, on the other.     

Tree seed germination and seedling establishment in treefall gaps and understorey in a subtropical forest of northeast India

Abstract Seed germination, and survival and growth of seedlings of four dominant tree species, Quercus dealbata, Quercus griffithii, Quercus glauca and Schima khasiana were studied in the treefall gaps and forest understorey of an undisturbed mature-phase humid subtropical broadleaved forest in northeast India. Three important microenvironmental factors namely photosynthetically active radiation (PAR), soil moisture and litter depth, were also measured in the forest understorey and gaps and correlated with seedling mortality. Seed germination of S. khasiana was significantly higher in the treefall gaps than in the understorey; among the tree species studied, it had the highest germination. Quercus seedlings were abundant in the understorey and small gaps, while S. khasiana seedlings were more numerous in the large gaps. The survivorship curves for the seedling populations revealed that the three Quercus species survived better in the understorey, while S. khasiana did so in the gaps. PAR and soil moisture were positively correlated with tree seedling mortality, which occurred mainly during the winter months. The Quercus seedlings grew better in the forest understorey and small gaps and S. khasiana seedlings in the large gaps. The differential performance of the tree seedlings to the conditions prevailing in the understorey and gaps of two sizes indicates that different species were adapted to different light environments depending upon their optimum requirements. This could be an effective mechanism for promoting species coexistence in the forest community.     

The effect of cactus spines on light interception and Photosystem II for three sympatric species of Opuntia from the Mojave Desert

Cactus spines reduce herbivory, direct water toward roots and reduce the impacts of high- and low-temperature extremes. Yet, shading of stems by spines reduces incident photosynthetic photon flux density (PFD), photosynthesis and growth. This study compared spinescence, PFD interception, stem temperature, Photosystem II (PSII) photochemistry and xanthophyll pigment composition for three species of cacti from the Mojave Desert, CA. The species vary in spinescence: Opuntia basilaris , which has no central or radial spines; Opuntia erinacea , which is densely covered with spines; and Opuntia phaeacantha , which has an intermediate coverage of spines. The role of spines was tested by removing spines from stems of O. erinacea . PFD interception was similar for both O. basilaris and O. phaeacantha , and about three times that for densely spined O. erinacea ; removal of spines increased incident PFD three-fold. There were no effects of spines on stem temperatures. Steady-state light-response curves of chlorophyll a fluorescence from PSII indicated that ΦPSII, photochemical quenching (qP) and electron flux within PSII were lower, and non-photochemical quenching was higher, for O. erinacea in comparison to the other two species with less spines. After 2 months, qP was higher and electron flux lower, and xanthophyll pigment pool size was higher, for stems from which spines had been removed compared with intact stems. These three species allocate different amounts of biomass to spines, resulting in species-specific PFD interception, PSII photochemistry and xanthophyll pigment pool size, which may help maintain rates of photosynthesis during the hot, dry Mojave Desert summer.     

The complexity of forest borders determines the understorey vegetation

Questions

What are the most important drivers of plant species richness (gamma‐diversity) and species turnover (beta‐diversity) in the field layer of a forest edge? Does the tree and shrub species richness structure and complexity affect the richness of forest and grassland specialist species?

Location

Southeast Sweden.

Methods

We sampled 50 forest edges with different levels of structural complexity in agricultural landscapes. In each border we recorded trees, shrubs and herb layer species in a 50‐m transect parallel with the forest. We investigated species composition and species turnover in relation to the proportions of gaps in the border and the diversity of trees and shrubs.

Results

Total plant species richness in the field layer was mainly explained by the proportion of gaps to areas with full canopy cover and tree diversity. Increasing number of gaps promoted higher diversity of grassland specialist species within the field layer, resulting in open forest borders with the highest overall species richness. Gaps did however have a negative impact on forest species richness. Conversely, increasing forest species richness was positively related to tree diversity, but the number of grassland specialist species was negatively affected by tree diversity.

Conclusions

Managing forest borders, and therefore increasing the area of semi‐open habitats in fragmented agricultural landscapes, provides future opportunities to create a network of suitable habitats for both grassland and deciduous forest specialist species. Such measures therefore have the potential to increase functional connectivity and support dispersal of species in homogeneous forest/agricultural landscapes.     

Limits to tree species invasion in pampean grassland and forest plant communities

Factors limiting tree invasion in the Inland Pampas of Argentina were studied by monitoring the establishment of four alien tree species in remnant grassland and cultivated forest stands. We tested whether disturbances facilitated tree seedling recruitment and survival once seeds of invaders were made available by hand sowing. Seed addition to grassland failed to produce seedlings of two study species, Ligustrum lucidum and Ulmus pumila, but did result in abundant recruitment of Gleditsia triacanthos and Prosopis caldenia. While emergence was sparse in intact grassland, seedling densities were significantly increased by canopy and soil disturbances. Longer-term surveys showed that only Gleditsia became successfully established in disturbed grassland. These results support the hypothesis that interference from herbaceous vegetation may play a significant role in slowing down tree invasion, whereas disturbances create microsites that can be exploited by invasive woody plants. Seed sowing in a Ligustrum forest promoted the emergence of all four study species in understorey and treefall gap conditions. Litter removal had species-specific effects on emergence and early seedling growth, but had little impact on survivorship. Seedlings emerging under the closed forest canopy died within a few months. In the treefall gap, recruits of Gleditsia and Prosopis survived the first year, but did not survive in the longer term after natural gap closure. The forest community thus appeared less susceptible to colonization by alien trees than the grassland. We conclude that tree invasion in this system is strongly limited by the availability of recruitment microsites and biotic interactions, as well as by dispersal from existing propagule sources.     

Potential causes of arrested succession in Kibale National Park, Uganda: growth and mortality of seedlings

Recent studies suggest that regeneration following large-scale disturbance in Kibale National Park, Uganda, is slow or possibly arrested. Here, data is provided on the growth and mortality of seedlings in the forest understory, treefall gaps, and in large gaps that suggest that this pattern of arrested succession can be attributed partially to the fact that this East African community lacks aggressive colonizing tree species. Growth and mortality rates were contrasted for seedlings of six tree species planted in the understory, small gaps, and large gaps for 36 months. Data suggest that species are adapted to gaps of particular sizes. For example, Uvariopsis congensis grew faster in the understory than in small gaps, whereas Warburgia ugandensis had the lowest mortality rate and highest growth rate in large gaps. Seedlings (n=170) of 15 species were transplanted to assess the response of the tree community to large gap conditions. The limited survival of seedlings in large gaps relative to the understory suggests that only a small proportion of the tree community in this forest regenerates best in gaps larger than those created by the collapse of a single tree. These findings differ from a number of studies conducted in other geographical regions, and suggests that tree communities differ with respect to the proportion of tree species adapted to gaps of particular sizes. This may relate to variation among regions in their history of disturbance and thus frequency of gap formation, size of gaps, and the duration of periods of release. Such variation could imply the existence of a corresponding pattern among tropical forests of differential vulnerability to human disturbance, which tends to create many large gaps.     

卧龙自然保护区亚高山暗针叶林树种更新研究

在卧龙亚高山暗针叶林中选取有代表性的18个林窗斑块以及对照的林下样方进行群落学调查,记录乔木树种和灌木树种的相关数量特征。按照树种在林窗内外的重要值差异结合树种本身生态学特性将群落中出现的乔木层树种划分为先锋组和耐荫组两类生态种组。林窗内外乔木树种的组成明显不同,两类生态种组树种幼苗在林窗与林下环境中的更新表现出差别,这与树种本身的生态学特性以及所处林窗的环境有关,同时亚高山暗针叶林中灌木层优势种小径竹的生长对树种更新方式也产生一定的影响。     

Phylogenetic Structure of Tree Species across Different Life Stages from Seedlings to Canopy Trees in a Subtropical Evergreen Broad-Leaved Forest

Investigating patterns of phylogenetic structure across different life stages of tree species in forests is crucial to understanding forest community assembly, and investigating forest gap influence on the phylogenetic structure of forest regeneration is necessary for understanding forest community assembly. Here, we examine the phylogenetic structure of tree species across life stages from seedlings to canopy trees, as well as forest gap influence on the phylogenetic structure of forest regeneration in a forest of the subtropical region in China. We investigate changes in phylogenetic relatedness (measured as NRI) of tree species from seedlings, saplings, treelets to canopy trees; we compare the phylogenetic turnover (measured as β_NRI) between canopy trees and seedlings in forest understory with that between canopy trees and seedlings in forest gaps. We found that phylogenetic relatedness generally increases from seedlings through saplings and treelets up to canopy trees, and that phylogenetic relatedness does not differ between seedlings in forest understory and those in forest gaps, but phylogenetic turnover between canopy trees and seedlings in forest understory is lower than that between canopy trees and seedlings in forest gaps. We conclude that tree species tend to be more closely related from seedling to canopy layers, and that forest gaps alter the seedling phylogenetic turnover of the studied forest. It is likely that the increasing trend of phylogenetic clustering as tree stem size increases observed in this subtropical forest is primarily driven by abiotic filtering processes, which select a set of closely related evergreen broad-leaved tree species whose regeneration has adapted to the closed canopy environments of the subtropical forest developed under the regional monsoon climate.     

Photosynthetic induction in saplings of three shade-tolerant tree species: comparing understorey and gap habitats in a French Guiana rain forest

The photosynthetic induction response under constant and fluctuating light was examined in naturally occurring saplings (about 0.5-2 m in height) of three shade-tolerant tree species, Pourouma bicolor spp digitata, Dicorynia guianensis, and Vouacapoua americana, growing in bright gaps and in the shaded understorey in a Neotropical rain forest. Light availability to saplings was estimated by hemispherical photography. Photosynthetic induction was measured in the morning on leaves that had not yet experienced direct sunlight. In Dicorynia, the maximum net photosynthesis rate (A_max) was similar between forest environments (ca 4 µmol m^-2 s^-1), whereas for the two other species, it was twice as high in gaps (ca 7.5) as in the understorey (ca 4.5). However, the time required to reach 90% of A_max did not differ among species, and was short, 7-11 min. Biochemical induction was fast in leaves of Pourouma, as about 3 min were needed to reach 75% of maximum carboxylation capacity (V_cmax); the two other species needed 4-5 min. When induction continued after reaching 75% of V_cmax, stomatal conductance increased in Pourouma only (ca 80%), causing a further increase in its net photosynthesis rate. When fully induced leaves were shaded for 20 min, loss of induction was moderate in all species. However, gap saplings of Dicorynia had a rapid induction loss (ca 80%), which was mainly due to biochemical limitation as stomatal conductance decreased only slowly. When leaves were exposed to a series of lightflecks separated by short periods of low light, photosynthetic induction increased substantially and to a similar extent in all species. Although A_max was much lower in old than in young leaves as measured in Dicorynia and Vouacapoua, variables of the dynamic response of photosynthesis to a change in light tended to be similar between young and old leaves. Old leaves, therefore, might remain important for whole-plant carbon gain, especially in understorey environments. The three shade-tolerant species show that, particularly in low light, they are capable of efficient sunfleck utilization.     

Acclimation of tropical tree seedlings to excessive light in simulated tree-fall gaps

Acclimation to periodic high‐light stress was studied in tree seedlings from a neotropical forest. Seedlings of several pioneer and late‐succession species were cultivated under simulated tree‐fall gap conditions; they were placed under frames covered with shade cloth with apertures of different widths that permitted defined periods of daily leaf exposure to direct sunlight. During direct sun exposure, all plants exhibited a marked reversible decline in potential photosystem II (PSII) efficiency, determined by means of the ratio of variable to maximum Chl a fluorescence (F_v/F_m). The decline in F_v/F_m under full sunlight was much stronger in late‐succession than in pioneer species. For each gap size, all species exhibited a similar degree of de‐epoxidation of violaxanthin in direct sunlight and similar pool sizes of xanthophyll cycle pigments. Pool sizes increased with increasing gap size. Pioneer plants possessed high levels of β‐carotene that also increased with gap size, whereas α‐carotene decreased. In contrast to late‐succession plants, pioneer plants were capable of adjusting their Chl a/b ratio to a high value in wide gaps. The content of extractable UV‐B‐absorbing compounds was highest in the plants acclimated to large gaps and did not depend on the successional status of the plants. The results demonstrate a better performance of pioneer species under high‐light conditions as compared with late‐succession plants, manifested by reduced photoinhibition of PSII in pioneer species. This was not related to increased pool size and turnover of xanthophyll cycle pigments, nor to higher contents of UV‐B‐absorbing substances. High β‐carotene levels and increased Chl a/b ratios, i.e. reduced size of the Chl a and b binding antennae, may contribute to photoprotection in pioneer species.     

Mechanisms of competition: thermal inhibition of tree seedling growth by grass

The relative importance of thermal interference and competition for below-ground resources in the inhibition of tree seedling growth by grass was determined under field conditions. Snow gum (Eucalyptus pauciflora) seedlings were grown in bare soil or soil covered with either live grass or straw. Covering soil with straw produced thermal conditions in soil and air that were indistinguishable from those associated with live grass. In contrast, seedlings grown in bare soil experienced more rapid increase in soil temperatures during late winter and spring, less frequent and less severe frosts, and temperature maxima that more closely followed those of the atmosphere than seedlings growing in live grass or straw. After 1 year, seedlings in bare soil had four times the biomass of those grown in grass or straw. Inhibition of seedling growth by grass was attributed to alteration of the thermal environment which caused (1) seedlings to have a short growing season largely restricted to summer, (2) temporal separation in competition for resources with consumption of below-ground resources by grass in spring reducing availability of resources to support tree seedling growth in early summer, and (3) seedlings to be more subject to stress from temperature extremes. These results show that thermal interference plays a major role in interactions between plants.     

Effects of seedling size, El Niño drought, seedling density, and distance to nearest conspecific adult on 6-year survival of Ocotea whitei seedlings in Panamá

We present an analysis of the long-term survival of two cohorts of seedlings of the tropical canopy tree Ocotea whitei (Lauraceae) on a 1-ha plot of mature, lowland moist forest on Barro Colorado Island, Panamá. In 1980, we counted an even-aged cohort of seedlings that germinated in 1979, then measured and tagged survivors in 1981. We also measured and tagged a second, smaller cohort of seedlings that germinated in 1981. We followed the subsequent survival of all seedlings through 1985. Seedling mortality was phenotypically, temporally, and spatially non-random. Important correlates of non-random mortality included: (1) seedling size and age, (2) an El Niño drought, and (3) biotic neighborhood. Larger and older seedlings survived better than smaller and younger seedlings, respectively, and the El Niño-related drought of 1982-1983 was associated with elevated mortality rates. Seedling density, which was strongly correlated with the proximity to the nearest conspecific adult, increased mortality. The observed mortality patterns suggest that processes consistent with the Janzen-Connell hypothesis operate during the recruitment phase of O. whitei population dynamics. However, the processes causing the observed density- and distance-dependent mortality may vary with factors such as total seed number, seedling size, and climatic variation, making it difficult to determine whether time-integrated seedling-to-adult spacing mechanisms other than self-thinning operate on a given plant population. After 6 years in the hectare studied, survivors remained densest and most numerous underneath the adult trees. We conclude that only long-term demographic data, collected at a variety of scales on a variety of species, will ultimately answer the question: do Janzen-Connell effects contribute substantially to structuring tropical forests?     

Positive and negative plant interactions contribute to a north-south-patterned association between two desert shrub species

Abiotic factors are often thought to be the predominant forces shaping desert plant communities. But both positive and negative interactions between plants are frequently observed in deserts, and it is an open question whether they can strongly affect the spatial structure of a desert community. The goal of this study was to answer this question for a plant community in the North American Mojave Desert. Two semi-shrub species, Ambrosia dumosa and Acamptopappus sphaerocephalus, were the focus of this study. At the study site, seedlings emerged predominantly on the northern side of shrubs, indicating positive effects of canopy shading on emergence, but survival of Ambrosia seedlings was much higher in open areas than at the edge of conspecific shrubs. Negative intraspecific interactions also affected Ambrosia shrubs, which did not increase in size over a 4-year period unless the nearest conspecific neighbor had been removed. These negative intraspecific interactions among different life stages of Ambrosia appear to contribute to spatial segregation observed among shrubs of this species. In contrast, Acamptopappus shrubs and their seedlings were aggregated with Ambrosia shrubs, and occurred more often on the northern side of Ambrosia than expected by chance. Removal of Ambrosia neighbors positively affected growth of Acamptopappus, but only when the neighbor was removed on the northern side. For Acamptopappus, an Ambrosia neighbor on the southern side may have some positive effects, which appear to neutralize the negative effects found for northern neighbors. These positive effects were likely at least partly due to shading. Removal of Ambrosia neighbors negatively affected predawn xylem pressure potentials of Acamptopappus, but this effect was only found during one growing season and was briefly reversed during the next. In summary, negative intraspecific interactions appear to cause spatial segregation of Ambrosia shrubs, while a combination of positive and negative interactions apparently contribute to the directional association between Ambrosia and Acamptopappus. Thus plant interactions in this desert appear to shape community structure in at least two dimensions by influencing the distances and in which directions to their neighbors plants can grow and survive.     

Zonation of shrubs in western Atlantic salt marshes

We explored the generality of the processes mediating shrub zonation in western Atlantic salt marshes by comparing the results of our experiments in Georgia, USA with previous studies from Rhode Island, USA. The shrub Borrichia frutescens dominates the terrestrial border of many Georgia salt marshes. Within the shrub zone, physical stress increased at lower elevations, shrubs at lower elevations were stunted, and experimentally reducing physical stress reduced shrub stunting. Below the shrub zone, physical stress increased further, and the grass Spartina alterniflora dominated. Transplant and neighbor-removal experiments indicated that the lower border of the shrub zone was set more by physical stress than by competition, but that the upper border of the grass zone was set primarily by competition with shrubs. Laboratory experiments indicated that S. alterniflora seedlings survived best and shrub seedlings worst in the flooded, salty treatment that mimicked low-marsh conditions. These processes are similar to those maintaining zonation patterns between the shrub Iva frutescens and the rush Juncus gerardi in Rhode Island salt marshes. However, markedly different processes appear to occur further to the north, where woody shrubs are absent from coastal marshes, and further to the south, where woody plants (mangroves) dominate coastal wetlands.     

Photosynthesis, photoinhibition, and nitrogen use efficiency in native and invasive tree ferns in Hawaii

Photosynthetic gas exchange, chlorophyll fluorescence, nitrogen use efficiency, and related leaf traits of native Hawaiian tree ferns in the genus Cibotium were compared with those of the invasive Australian tree fern Sphaeropteris cooperi in an attempt to explain the higher growth rates of S. cooperi in Hawaii. Comparisons were made between mature sporophytes growing in the sun (gap or forest edge) and in shady understories at four sites at three different elevations. The invasive tree fern had 12-13 cm greater height increase per year and approximately 5 times larger total leaf surface area per plant compared to the native tree ferns. The maximum rates of photosynthesis of S. cooperi in the sun and shade were significantly higher than those of the native Cibotium spp (for example, 11.2 and 7.1 µmol m^-2 s^-1, and 5.8 and 3.6 µmol m^-2 s^-1 respectively for the invasive and natives at low elevation). The instantaneous photosynthetic nitrogen use efficiency of the invasive tree fern was significantly higher than that of the native tree ferns, but when integrated over the life span of the frond the differences were not significant. The fronds of the invasive tree fern species had a significantly shorter life span than the native tree ferns (approximately 6 months and 12 months, respectively), and significantly higher nitrogen content per unit leaf mass. The native tree ferns growing in both sun and shade exhibited greater photoinhibition than the invasive tree fern after being experimentally subjected to high light levels. The native tree ferns recovered only 78% of their dark-acclimated quantum yield (F_v/F_m), while the invasive tree fern recovered 90% and 86% of its dark-acclimated F_v/F_m when growing in sun and shade, respectively. Overall, the invasive tree fern appears to be more efficient at capturing and utilizing light than the native Cibotium species, particularly in high-light environments such as those associated with high levels of disturbance.     

Positive correlation between levels of retained zeaxanthin + antheraxanthin and degree of photoinhibition in shade leaves of Schefflera arboricola (Hayata) Merrill

Attached intact leaves of Schefflera arboricola grown at three different photon flux densities (PFDs) were subjected to 24-h exposures to a high PFD and subsequent recovery at a low PFD. While sun leaves showed virtually no sustained effects on photosystem II (PSII), shade-grown leaves exhibited pronounced photoinhibition of PSII that required several days at low PFD to recover. Upon transfer to high PFD, levels of nonphotochemical quenching in PSII as well as levels of zeaxanthin were initially low in shade leaves but continued to increase gradually during the 24-h exposure. The xanthophyll cycle pool size rose gradually during and also subsequent to the photoinhibitory treatment in shade leaves. Upon return to low PFD, a marked and extremely long-lasting retention of zeaxanthin and antheraxanthin was observed in shade but not sun leaves. During recovery, changes in the conversion state of the xanthophyll cycle therefore closely mirrored the slow increases in PSII efficiency. This novel report of a close association between zeaxanthin retention and lasting PSII depressions in these shade leaves clearly suggests a role for zeaxanthin in photoinhibition of shade leaves. In addition, there was a decrease in β-carotene levels, some decrease in chlorophyll, but no change in lutein and neoxanthin (all per leaf area) in the shade leaves during and subsequent to the photoinhibitory treatment. These data may be consistent with a degradation of a portion of core complexes but not of peripheral light-harvesting complexes. A possible conversion of β-carotene to form additional zeaxanthin is discussed. Received: 24 October 1997 / Accepted: 12 November 1997     

Influence of forest type and tree species on canopy ants (Hymenoptera: Formicidae) in Budongo Forest, Uganda

The arboreal ant fauna was investigated in Budongo Forest, a seasonal rain forest in Uganda, using the insecticidal fogging technique. Ants were collected from 61 trees, between 7 and 33 m in height, belonging to four tree species. Trees were growing in adjacent plots of forests characterized by different use and structure: an old primary forest, a primary swamp forest along a small river, and a secondary forest where selective logging was carried out for 30 years. A total number of 37,065 ants, belonging to 161 species in 30 genera were collected. Considering the high number of species found only once, the completeness of the canopy ant fauna was relatively high and of relatively similar magnitude as samples from the Neotropics or the Oriental region. Up to 37 ant species on a single tree, with an average of 18.2 species per tree, were found. Forty-four ant species (28.1%) were found only once, less than ten individuals were found for each of 88 species (54.7%), but 64.0% of all individuals belonged to one of five species. Considering the high numerical dominance of a few ant species like a Pheidole sp., Tetramorium aculeatum (Mayr) and a Crematogaster sp., there is some evidence for an ant mosaic in the lower canopy of the Budongo Forest. Individual numbers of ants were strongly correlated with nests in the fogged tree, though the ants were not homogeneously distributed in the tree crowns. Diversity measures that strongly depend on individual numbers such as the Morisita-Horn index or rarefaction methods were calculated, but results were not concordant with those of incidence-based estimates such as jack-knife calculations. Differences in ant species richness and faunal composition between tree species were low, but more significant between forest types. The ant fauna in the secondary forest was less diverse with 12.6% fewer species compared to the primary forest sites. The average number of ant species per tree was significantly lower in the secondary forest (<20% of the species; F=8.03, df=59, P<0.01) than in the undisturbed forest types. Cataulacus, Leptothorax, Tetraponera, and Polyrhachis, which are typical canopy-dwelling ant genera, had a significantly higher diversity and frequency in the two primary forest types (F=4.17, df=53, P<0.05). Secondary forest trees are often younger, lacking dead branches and epiphytes which are important requisites for ant colonization on trees.     

ACCLIMATION OF EMILIANIA HUXLEYI (PRYMNESIOPHYCEAE) TO PHOTON FLUX DENSITY1

Growth rate, pigment composition, and noninvasive chl a fluorescence parameters were assessed for a noncalcifying strain of the prymnesiophyte Emiliania huxleyi Lohman grown at 50, 100, 200, and 800 μmol photons·m^?2·s^?1. Emiliania huxleyi grown at high photon flux density (PFD) was characterized by increased specific growth rates, 0.82 d^?1 for high PFD grown cells compared with 0.38 d^?1 for low PFD grown cells, and higher in vivo chl a specific attenuation coefficients that were most likely due to a decreased pigment package, consistent with the observed decrease in cellular photosynthetic pigment content. High PFD growth conditions also induced a 2.5‐fold increase in the pool of the xanthophyll cycle pigments diadinoxanthin and diatoxanthin responsible for dissipation of excess energy. Dark‐adapted maximal photochemical efficiency (F_v/F_m) remained constant at around 0.58 for cells grown over the range of PFDs, and therefore the observed decline, from 0.57 to 0.33, in the PSII maximum efficiency in the light‐adapted state, (F_v′/F_m′), with increasing growth PFD was due to increased dissipation of excess energy, most likely via the xanthophyll cycle and not due to photoinhibition. The PSII operating efficiency (F_q′/F_m′) decreased from 0.48 to 0.21 with increasing growth PFD due to both saturation of photochemistry and an increase in nonphotochemical quenching. The changes in the physiological parameters with growth PFD enable E. huxleyi to maximize rates of photosynthesis under subsaturating conditions and protect the photosynthetic apparatus from excess energy while supporting higher saturating rates of photosynthesis under saturating PFDs.