The European spruce bark beetle <Emphasis Type="Italic">Ips typographus</Emphasis> in a national park: from pest to keystone species

The influence of natural disturbance on biodiversity is poorly known in the intensively cultivated landscape of Europe. As an example of insect disturbance we studied effects of gaps generated by outbreaks of the spruce bark beetle (Ips typographus) on biodiversity in the area of the National Park “Bavarian Forest” and compared them with openings (e.g. meadows) created by humans in these forests. Insects were sampled using flight interception traps across twelve ecotones between edges of closed forest, six bark beetle gaps and six meadows. The diversity and species density of true bugs and of bees/wasps increased significantly from the closed stand to the edge, and continued to increase inside the openings at interior and exterior edges. Species density in saproxylic beetles also increased significantly from closed forest to opening, but only across ecotones including bark beetle gaps. Similarly, the number of critically endangered saproxylic beetles increased significantly in bark beetle gaps. Using indicator species analysis a total of 60 species were identified as possessing a statistically significant value indicating preference for one of the habitat types along the ecotones: 29 of them preferred gaps, 24 preferred meadows, three were characteristic for edges of meadows, three for edges of bark beetle gaps, but only one was typical of closed forest. Most of our results support the thesis that I. typographus fulfils the majority of criteria for a keystone species, particularly that of maintenance of biodiversity in forests. Our results emphasize the value for the study and conservation of insect diversity of the policy of non-interference with natural processes pursued in some protected areas. As a recommendation to forest management for increasing insect diversity even in commercial forest, we suggest that logging in recent gaps in medium aged mixed montane stands should aim at retention of a part of the dead wood. Planting should be avoided, to lengthen the important phase of sunlit conditions.     

Population dynamics in changing environments: the case of an eruptive forest pest species

In recent decades we have seen rapid and co‐occurring changes in landscape structure, species distributions and even climate as consequences of human activity. Such changes affect the dynamics of the interaction between major forest pest species, such as bark beetles (Coleoptera: Curculionidae, Scolytinae), and their host trees. Normally breeding mostly in broken or severely stressed spruce; at high population densities some bark beetle species can colonise and kill healthy trees on scales ranging from single trees in a stand to multi‐annual landscape‐wide outbreaks. In Eurasia, the largest outbreaks are caused by the spruce bark beetle, Ips typographus (Linnaeus), which is common and shares a wide distribution with its main host, Norway spruce (Picea abies Karst.). A large literature is now available, from which this review aims to synthesize research relevant for the population dynamics of I. typographus and co‐occurring species under changing conditions. We find that spruce bark beetle population dynamics tend to be metastable, but that mixed‐species and age‐heterogeneous forests with good site‐matching tend to be less susceptible to large‐scale outbreaks. While large accumulations of logs should be removed and/or debarked before the next swarming period, intensive removal of all coarse dead wood may be counterproductive, as it reduces the diversity of predators that in some areas may play a role in keeping I. typographus populations below the outbreak threshold, and sanitary logging frequently causes edge effects and root damage, reducing the resistance of remaining trees. It is very hard to predict the outcome of interspecific interactions due to invading beetle species or I. typographus establishing outside its current range, as they can be of varying sign and strength and may fluctuate depending on environmental factors and population phase. Most research indicates that beetle outbreaks will increase in frequency and magnitude as temperature, wind speed and precipitation variability increases, and that mitigating forestry practices should be adopted as soon as possible considering the time lags involved.     

Differences in the fruit maturation stages at which oviposition occurs among insect seed predators feeding on the fruits of five dipterocarp tree species

The seeds of dipterocarp trees are the main food resources for many species of weevils, bark beetles and small moths; however, for most seed‐eating insects on dipterocarp tropical trees, seed utilization patterns remain poorly investigated. This study aimed to determine the fruit maturation stages at which eggs are laid by different insect seed predators feeding on the seeds or fruits of the following five dipterocarp species: Dipterocarpus globosus, Dryobalanops aromatica, Shorea beccariana, S. acuta and S. curtisii, which reproduced during the same period. We investigated the occurrence frequencies of the insect seed predators at various growth stages by collecting both unfallen and fallen fruit on several occasions during the period of seed/fruit maturation in a tropical rainforest in Borneo from September to December 2013. Weevils and bark beetles were the dominant insect seed predators of the five tree species. One or two weevil species of Alcidodes, Damnux and/or Nanophyes preyed on the seeds of each of the five tree species, and one bark beetle species, Coccotrypes gedeanus, preyed on the seeds of all five tree species. Many larvae, pupae and adults of each weevil species were found in pre‐dispersal (unfallen) fruit, whereas bark beetles at various growth stages were found in post‐dispersal (fallen) fruit. These results suggested that, among the dominant insect seed predators of the five dipterocarp species, weevil species oviposit on pre‐dispersal fruit and begin their larval growth before seed dispersal, whereas the oviposition and larval development of bark beetle species occurs in post‐dispersal fruit.     

Habitat Islands,Forest Edge and Spring-active Invertebrate Assemblages

Forest management results in forest patches of varying sizes within a clearcut matrix. The result is a large amount of edge habitat and many small patches across the landscape. Here we describe the spring-active epigeal spider and carabid fauna found at the forest-clearcut edge of spruce forest in northern Ontario, Canada. We include two types of edge: the forest-clearcut interface and the small habitat patches formed by forest residuals within the clearcut. Spring-active forest spiders and carabids appear little affected by adjacent clearcutting activity, and some forest species, such as Agyneta olivacea (Emetron), Diplocentria bidentata (Emetron) and Microneta viaria (Blackwall), are more prevalent at the forested edge. Common and abundant spider species were equally recorded in forest interior and forest edge. Generally, no invasion of open-habitat species was observed within the forest, although smaller forest patches may be at higher risk.     

Distribution of ground-dwelling beetles （Coleoptera） across a forest-clearcut ecotone in Wolong Natural Reserve, southwestern China

Abstract The influence of edge effect on ground-dwelling beetles (Coleoptera) across a forest-clearcut ecotone was studied in Wolong Natural Reserve, southwestern China. During the field research, a total of 30 739 beetles were collected with pitfall traps along transects, which extending 100 m from the edge into the forest interior and 100 m into the clearcut. Of the collection, Carabidae comprised 92%, Staphylinidae 3%, Curculionidae 2%, and Tenebrionidae 2%, and these four families can be considered as abundant groups. Family richness, Shannon diversity and equitability display a significant decrease from forest interior, edge to clearcut. Based on the family composition and abundance, ground-dwelling beetles of the forest interior can be separated from those in the clearcut by Principal coordinate analysis ordination, and beetle assemblages in the forest edge were more similar to forest assemblages than to those found in the clearcut by cluster analysis. Seasonal dynamics of family richness showed a monotone peak in the middle season, with a highest value in the forest interior and a lowest value in the clearcut. Family abundance showed two peaks in the middle season, always with more individuals in the clearcut than in the forest interior or in the edge. Multiple linear regression analyses showed that the cover of shrub and litter were the two most important factors in determining family richness, Shannon diversity, equitability and abundance. Our results show that the forest edge and clearcut have obviously different composition and diversity of ground-dwelling beetles from forest interior at the family level. However, more edges have been formed due to increasing forest fragmentation (clearcutting or logging), so it is necessary to preserve large and intact forest to protect the diversity of ground-dwelling beetles in Wolong Natural Reserve.     

Attraction to monoterpenes and beetle‐produced compounds by syntopic Ips and Dendroctonus bark beetles and their predators

• 1 Bark beetles are significant mortality agents of conifers. Four beetle species, the pine engraver Ips pini, the six‐spined pine engraver Ips calligraphus sub. ponderosae, the southern pine beetle Dendroctonus frontalis, and the western pine beetle Dendroctonus brevicomis, cohabitate pines in Arizona.
• 2 A pheromone trapping study in ponderosa forests of Arizona determined the attraction of beetles to conspecific and heterospecific pheromone components in the presence and absence of host volatiles, and tested whether predators differ in their attraction to combinations of pheromone components and tree monoterpenes.
• 3 All four bark beetle species differed in their responses to heterospecific lures and monoterpenes. Ips calligraphus was the only species that increased in trap catches when heterospecific lures were added. Heterospecific lures did not inhibit the attraction of either Dendroctonus or Ips species. The replacement of myrcene with α‐pinene increased the attraction of Dendroctonus, whereas the addition of α‐pinene had mixed results for Ips. The prominent predators Temnochila chlorodia and Enoclerus lecontei were more attracted to the I. pini lure than the D. brevicomis lure, and the combination of the two lures with α‐pinene was most attractive to both predator species.
• 4 Cross attraction and limited inhibition of bark beetles to heterospecific pheromones suggest that some of these species might use heterospecific compounds to increase successful location and colonization of trees. Predator responses to treatments suggest that tree volatiles are used to locate potential prey and predators are more responsive to Ips than to Dendroctonus pheromone components in Arizona.

     

边缘效应对卧龙自然保护区森林-草地群落交错带地表甲虫多样性的影响

本文研究了卧龙国家自然保护区地表甲虫群落在天然落叶阔叶林森林边缘与森林内部和周围草地间多样性差异,在科级水平上探讨边缘效应对地表甲虫群落的影响。调查共设5个重复样带（间距大于500 m）;每个样带以距离梯度（25 m）的方式设置样点,分别由边缘深入到森林内部和草地中央100 m,共设45个样点,通过巴氏罐诱法调查地表甲虫群落组成和季节变化。本研究采集甲虫标本4 736 号,隶属于28个科,步甲科、隐翅虫科和叩甲科分别占总数的49.5%、23.5%和13.0%,共同构成本研究地区地表甲虫的优势类群。甲虫的个体数量从森林内部、边缘到周围草地依次降低,而科多样性和均匀度则依次增高,都达到了显著差异。主坐标分析（PCoA）排序表明,森林内部和周围草地间的地表甲虫群落组成差异较大;而森林边缘的群落组成与两者都有较高程度的相似性,反映了森林边缘的地表甲虫群落已经与森林内部的群落组成发生明显分化,除了森林物种成分外,还包含了若干从周围草地环境扩散来的物种成分。从季节动态看,森林边缘和森林内部在丰富度和个体数量的季节变化曲线方面相似性更高;捕食类和腐食类地表甲虫的季节变化在森林内部和边缘相似性更高,而植食类则在森林边缘和草地的相似性更高。多元回归分析表明,枯落物覆盖率是影响地表甲虫科多样性和均匀度的决定因素,枯落物厚度和林冠层覆盖率是决定甲虫个体数量分布的重要因素。以上结果表明,在科级水平上,地表甲虫群落在天然落叶阔叶林边缘已经与森林内部生境发生分化,而且正在加剧的森林片断化进程将会产生更多的森林边缘,因此,保留大面积的天然落叶阔叶林免受破坏和干扰是保护地表甲虫多样性的必要措施。     

Ophiostomatoid fungi isolated from Japanese red pine and their relationships with bark beetles

We isolated ophiostomatoid fungi from bark beetles infesting Pinus densiflora and their galleries at 24 sites in Japan. Twenty-one ophiostomatoid fungi, including species of Ophiostoma, Grosmannia, Ceratocystiopsis, Leptographium, and Pesotum, were identified. Among these, 11 species were either newly recorded in Japan or were previously undescribed species. Some of these fungal species were isolated from several bark beetles, but other species were isolated from only a particular beetle species. Thus, it is suggested that some ophiostomatoid fungi have specific relationships with particular beetle species. In addition, fungus-beetle biplots from redundancy analysis (RDA) summarizing the effects of beetle ecological characteristics suggested that the association patterns between bark beetles and the associated fungi seemed to be related to the niches occupied by the beetles.     

Fungi Vectored by the Bark Beetle Ips typographus Following Hibernation Under the Bark of Standing Trees and in the Forest Litter

The bark beetle Ips typographus has different hibernation environments, under the bark of standing trees or in the forest litter, which is likely to affect the beetle-associated fungal flora. We isolated fungi from beetles, standing I. typographus-attacked trees, and forest litter below the attacked trees. Fungal identification was done using cultural and molecular methods. The results of the two methods in detecting fungal species were compared. Fungal communities associated with I. typographus differed considerably depending on the hibernation environment. In addition to seven taxa of known ophiostomoid I. typographus-associated fungi, we detected 18 ascomycetes and anamorphic fungi, five wood-decaying basidomycetes, 11 yeasts, and four zygomycetes. Of those, 14 fungal taxa were detected exclusively from beetles that hibernated under bark, and six taxa were detected exclusively from beetles hibernating in forest litter. The spruce pathogen, Ceratocystis polonica, was detected occasionally in bark, while another spruce pathogen, Grosmannia europhioides, was detected more often from beetles hibernating under the bark as compared to litter. The identification method had a significant impact on which taxa were detected. Rapidly growing fungal taxa, e.g. Penicillium, Trichoderma, and Ophiostoma, dominated pure culture isolations; while yeasts dominated the communities detected using molecular methods. The study also demonstrated low frequencies of tree pathogenic fungi carried by I. typographus during its outbreaks and that the beetle does not require them to successfully attack and kill trees.     

Selective manipulation of predators using pheromones: responses to frontalin and ipsdienol pheromone components of bark beetles in the Great Lakes region

Abstract 1 One proposed approach to improving biological control of bark beetles (Coleoptera: Scolytidae; alt. Curculionidae: Scolytinae) is to manipulate predator movement using semiochemicals. However, selective manipulation is impeded by attraction of both predators and pests to bark beetle pheromones. 2 The primary bark beetle affecting pine plantations in Wisconsin, U.S.A., is the pine engraver, Ips pini (Say). Other herbivores include Ips grandicollis (Eichhoff) and Dryophthorus americanus Bedel (Curculionidae). The predominant predators are the beetles Thanasimus dubius (Cleridae) and Platysoma cylindrica (Histeridae). 3 We conducted field assays using two enantiomeric ratios of ipsdienol, and frontalin plus α‐pinene. Ipsdienol is the principal pheromone component of I. pini, and frontalin is produced by a number of Dendroctonus species. α‐Pinene is a host monoterpene commonly incorporated into commercial frontalin lures. 4 Thanasimus dubius was attracted to frontalin plus α‐pinene, and also to racemic ipsdienol. By contrast, I. pini was attracted to racemic ipsdienol, but showed no attraction to frontalin plus α‐pinene. Platysoma cylindrica was attracted to 97%‐(–)‐ipsdienol and, to a lesser extent, racemic ipsdienol, but not to frontalin plus α‐pinene. Ips grandicollis was attracted to frontalin plus α‐pinene but not to ipsdienol. Dryophthorus americanus was attracted to both ipsdienol and frontalin plus α‐pinene. 5 This ability to selectively attract the predator T. dubius without attracting the principal bark beetle in the system, I. pini, provides new opportunities for research into augmentative biological control and basic population dynamics. Moreover, the attraction of T. dubius, but not P. cylindrica, to frontalin plus α‐pinene creates opportunities for selective manipulation of just one predator. 6 Patterns of attraction by predators and bark beetles to these compounds appear to reflect various degrees of geographical and host tree overlap with several pheromone‐producing species.     

Dung beetles in continuous forest, forest fragments and in an agricultural mosaic habitat island at Los Tuxtlas, Mexico

With the aim of determining what kind of landscape mosaics might sustainmaximum diversity and minimum species loss, dung beetles were sampled withbaited pitfall traps to compare species richness and species composition in atract of continuous forest, forest fragments and a habitat island consisting ofa mosaic of forest and arboreal crops in Los Tuxtlas, southern Mexico. Wecaptured 7332 dung beetles representing 33 species. Similar numbers of specieswere captured in the three habitats. However, 56% of individuals were capturedin the continuous forest, 29% in the mosaic habitat and 15% in the forestfragments. Eight species (Canthon femoralis,Copris laeviceps, Canthidium centrale,Onthophagus batesi, Deltochilumpseudoparile, O. rhinolophus, Canthonviridis vazquezae and Dichotomius satanus) accounted for 90% of thecaptures, but their relative dominance varied among habitats. A clear trend wasevident in the number of dung beetles captured in the dung processing guilds(rollers/tunnelers) as well as in the diurnal and nocturnal guilds, withcaptures decreasing from the continuous forest to the mosaic habitat to theforest fragments. A similar trend was detected in detection rates for medium andsmall size dung producing mammals. Species richness of forest fragments andmosaic habitat did not differ from that found in the continuous forest, butthese habitats differ significantly in species richness from isolated shaded andunshaded plantations, linear strips of vegetation, the forest–pasture edge andpastures according to rarefaction analysis. The co-occurrence of the continuousforest, the mosaic habitat and the cluster of forest fragments in closeproximity seems to be preserving a diverse assemblage of dung beetle species inthe local landscape.     

Forest complexity drives dung beetle assemblages along an edge-interior gradient in the southwest Amazon rainforest

1. The habitat heterogeneity hypothesis predicts that heterogeneous habitats may provide more niches and diverse ways of exploiting environmental resources, thereby allowing more species to coexist, persist and diversify. 2. We aimed to investigate how an edge-interior gradient related to forest complexity influences species composition, abundance and richness of dung beetles in the western Amazon rainforest. We expected dung beetle abundance and richness to increase along the forest edge-interior gradient, in accordance with the habitat heterogeneity hypothesis. We also expected strong changes in species composition driven by species turnover in the forest interior and nestedness along the forest edges. We sampled dung beetles using baited pitfall traps across an edge-interior gradient. We also assessed the variation in forest features along the edge-interior gradient to identify changes in forest complexity. 3. Both species richness and abundance of dung beetles increased along the forest edge-interior, following the gradient of forest complexity. The Sorensen dissimilarity of dung beetle assemblages was higher among sampling units placed near the forest edge, although neither turnover, nor nestedness was different between the extremes of the forest edge-interior gradient. There was a clear compositional change along the edge-interior gradient mostly driven by species turnover. Individual indicator value analysis revealed that species were strongly associated with the forest interior conditions. 4. The simplification of the Amazon rainforest near clearings causes compositional changes in dung beetle assemblages. These changes are characterised by species-poor and low-abundance assemblages and may impair dung beetle ecological functions and therefore forest recovery.     

Defence syndromes in lodgepole – whitebark pine ecosystems relate to degree of historical exposure to mountain pine beetles

Warming climate is allowing tree‐killing bark beetles to expand their ranges and access naïve and semi‐naïve conifers. Conifers respond to attack using complex mixtures of chemical defences that can impede beetle success, but beetles exploit some compounds for host location and communication. Outcomes of changing relationships will depend on concentrations and compositions of multiple host compounds, which are largely unknown. We analysed constitutive and induced chemistries of Dendroctonus ponderosae's primary historical host, Pinus contorta, and Pinus albicaulis, a high‐elevation species whose encounters with this beetle are transitioning from intermittent to continuous. We quantified multiple classes of terpenes, phenolics, carbohydrates and minerals. Pinus contorta had higher constitutive allocation to, and generally stronger inducibility of, compounds that resist these beetle–fungal complexes. Pinus albicaulis contained higher proportions of specific monoterpenes that enhance pheromone communication, and lower induction of pheromone inhibitors. Induced P. contorta increased insecticidal and fungicidal compounds simultaneously, whereas P. albicaulis responses against these agents were inverse. Induced terpene accumulation was accompanied by decreased non‐structural carbohydrates, primarily sugars, in P. contorta, but not P. albicaulis, which contained primarily starches. These results show some host species with continuous exposure to bark beetles have more thoroughly integrated defence syndromes than less‐continuously exposed host species.     

Investigating the abundance and flight period of bark beetles (Coleoptera: Curculionidae: Scolytinae) over elevational gradients in Sitka spruce forests

1. A warming climate, as predicted under current climate change projections, is likely to influence the population dynamics of many forest insect species. Numerous bark beetle species in both Europe and North America have already responded to a warming climate by significantly expanding their geographical ranges.
2. The aim of the current study was to investigate how populations of bark beetles within stands of Sitka spruce, a widely planted non-native commercial plantation tree species in the U.K., were likely to respond to a warming climate. Experimental plots were established in stands of Sitka spruce over elevational gradients in two commercial forest plantations, and the abundance and emergence times of key bark beetle species were assessed over a 3-year period using flight interception traps. The air temperature difference between the lowest and highest experimental plot in each forest was consistently >1°C throughout the 3-year period.
3. In general, the abundance of the most dominant bark beetle species (e.g. Trypodendron, Dryocoetes, Hylastes spp.) was higher, and emergence times tended to be earlier in the year at the lower elevation plots, where temperatures were higher, although not all bark beetle species responded in the same manner.
4. The results of the study indicated that, under the projected future climate warming scenarios, monoculture Sitka spruce stands at low elevations may potentially be more vulnerable to significant outbreak events from existing or invasive bark beetle species. Hence, consideration of establishing more resilient forests of Sitka spruce by diversifying the species composition and structure of Sitka spruce stands is discussed.

     

Synchronous rise and fall of bark beetle and parasitoid populations in windthrow areas

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Oviposition behavior and reproductive success of the cerambycid Acanthocinus aedilis in the presence and absence of the bark beetle Tomicus piniperda

The influence of Tomicus piniperda (L.)(Col.:Scolytidae) attacks on Acanthocinus aedilis (L.)(Col.:Cerambycidae) (1) oviposition behavior, (2) breeding material preference and (3) reproductive success was investigated in caged pine bolts with and without egg galleries of T. piniperda. In addition, A. aedilis oviposition behavior was studied in the field. In the presence of T. piniperda attacks most A. aedilis eggs were laid through the entrance holes of the bark beetle egg galleries. In the absence of bark beetle attacks most A. aedilis eggs were laid at places where the outer bark had been damaged and the phloem was exposed. In cages, A. aedilis preferred to oviposit in bolts with bark beetle attacks. Thus, A. aedilis oviposited in all 17 of bark beetle attacked bolts but in only four of nine bolts without T. piniperda attacks. Neither the number of A. aedilis offspring nor the body size of emerging adults differed significantly between bolts attacked by bark beetles and unattacked bolts. Nor was there any significant relationship between the density of bark beetle egg galleries and the production of A. aedilis offspring.     

Horizontal and vertical distribution of saproxylic beetles (Col., Buprestidae, Cerambycidae, Scolytinae) across sections of forest edges

Abstract:  We evaluated the preferred home ranges of three saproxylic beetle taxa along transects from the open field into the forest interior, and from the forest floor up to the canopy. By means of trap sets on metal scaffolds, vertical and horizontal strata were sampled across two types of forest edges: soft-edge ecotones with a gradual transition from the field into the forest and hard edges with an abrupt transition. The forest edges consisted of different strata such as herbaceous fringe, shrub belt, unmanaged forest and managed forest. The thermophilic buprestids were mainly caught in the open land (herbaceous fringe and agricultural land) and in the upper forest mantle. In general, the cerambycids were most abundant in the open land and the lower forest mantle, but a few species favoured the forest interior. The bark beetles (Scolytinae) were equally distributed in all habitats. These distribution patterns of the taxa were observed in terms of both species numbers and abundances. Each species with at least five collected specimens was assigned to one of the three habitat types: open land, forest mantle and forest interior. Of 74 ranked species, only 16% were prevalent in the forest interior and are thus considered to be true forest species. The other 84% of the species were attributed to open land or the forest mantle and are, therefore, forest edge species. Soft forest edges generally supported a higher species richness than hard edges, particularly as regards Cerambycidae and Scolytinae. In terms of Shannon diversity, soft edges tended to be more diverse in buprestids and cerambycids. Overall, the forest interior showed the least species richness and diversity. Therefore, for the conservation of saproxylic beetles, not only the amount and quality of dead wood is important, but also the presence and design of forest boundary structures.     

Are bark beetle outbreaks less synchronous than forest Lepidoptera outbreaks?

Comparisons of intraspecific spatial synchrony across multiple epidemic insect species can be useful for generating hypotheses about major determinants of population patterns at larger scales. The present study compares patterns of spatial synchrony in outbreaks of six epidemic bark beetle species in North America and Europe. Spatial synchrony among populations of the Eurasian spruce bark beetle Ips typographus was significantly higher than for the other bark beetle species. The spatial synchrony observed in epidemic bark beetles was also compared with previously published patterns of synchrony in outbreaks of defoliating forest Lepidoptera, revealing a marked difference between these two major insect groups. The bark beetles exhibited a generally lower degree of spatial synchrony than the Lepidoptera, possibly because bark beetles are synchronized by different weather variables that are acting on a smaller scale than those affecting the Lepidoptera, or because inherent differences in their dynamics leads to more cyclic oscillations and more synchronous spatial dynamics in the Lepidoptera.     

Influence of Forest Type and Tree Species on Canopy‐Dwelling Beetles in Budongo Forest,Uganda1

Beetles were collected on 64 trees of four species (Cynometra alexandri C. H. Wright, Rinorea beniemis (Welwitsch ex Olivier) Kuntze, Teclea nobilis Delile, and Trichilia rubescens Olivier) in Budongo Forest, Uganda, using an insecticidal fogging technique. Selected tree species were abundant, taxonomically not closely related, and different in the shape of leaves, growth form, and size, with heights between 7 and 35 m. Trees were fogged in an old primary forest stand, in an area of secondary forest where selective logging was performed, and in a swamp forest. Eight conspecific trees per forest type were fogged. A total of 29,736 beetles were collected from all trees that could be assigned to 1433 (morpho)‐species; 41.6 percent were singletons and 89.6 percent of species were found with less than ten individuals. Abundant beetle taxa included Latridiidae (N= 4093), Chrysomelidae (3952), Staphylinidae (2931), Apioninae (2621), and Curculionidae (2457). Most species‐rich groups were Staphylinidae (N= 196 spp.), Curculionidae (189), and Chrysomelidae (148). Abundance increased in the order: primary < secondary < swamp forest. Due to the relatively high dominance of some species in the secondary forest, species richness increased in the order: secondary < primary < swamp forest. Beta diversity measures and factor analysis showed distinct differences among forest types but higher similarity of beetle communities on different tree species within one forest type. The taxonomic distribution of beetles in the secondary forest was more heterogeneous than in the primary forest. Analyses of the data revealed low host specificity even for phytophagous beetles, underlining the importance of habitat structure and chance effects on the spatial distribution of beetles in the canopy of Budongo Forest.     

Tree Ferns in the Interior and at the Edge of a Mexican Cloud Forest Remnant: Spore Germination and Sporophyte Survival and Establishment1

The three most common tree fern species in a Mexican montane cloud forest fragment (Alsophila firma, Lophosoria quadripinnata and Sphaeropteris horrida) were selected for laboratory and transplant experiments. The objectives were: (1) to determine the percentage of spore germination and gametophytes producing sporophytes, and (2) to compare early establishment of sporophytes at the edge and in the interior of the forest. Percent spore germination varied between 16 and 86 percent, and the number of gametophytes that produced sporophytes was high (>50%). Survival and growth of sporophytes differed between species and habitats. Survival was greater at the edge than in the forest interior for Lophosoria, but it was similar for Alsophila and Sphaeropteris. Number and length of fronds were higher at the forest edge for individuals of Alsophila and Lophosoria, but not for Sphaeropteris. RGR was higher at the forest edge than in the forest interior for sporophytes of the three species. This study suggests that the forest edge is an appropriate habitat for establishment of Alsophila and Lophosoria, but Sphaeropteris is apparently a forest interior species.