Stand structure and regeneration of <Emphasis Type="Italic">Populus euphratica</Emphasis> forest in the lower reaches of the Heihe River,NW China

Populus euphratica Oliv. is a main tree species that forms natural riparian forests in arid and semi-arid areas from Morocco to the Ordos Plateau. This study is designed to clarify the forest structure and dynamics of P. euphratica and to elucidate the ecological mechanisms sustaining riparian forests under unreliable environmental conditions. This study was conducted in a P. euphratica forest of the Ejina Oasis in Inner Mongolia, China, which is a hyperarid area. According to their tree size distribution, P. euphratica forests can be grouped into juvenile, mature, and overmatured stages. Almost all large P. euphratica showed dieback. The regeneration density on the forest floor shows a relation with the degree of height decrease due to dieback damage, as evaluated using the ratio of actual height to the maximum height estimated from the D–H relation. Therefore, after the mature stage, individual trees continue to grow while controlling their canopy size to adjust to changing environmental conditions in the overmatured stage. Our results suggest that P. euphratica growing under large fluctuations in groundwater levels exhibit a sophisticated regeneration system with canopy degradation.     

Soil carbon dynamics in regrowing forest of eastern Amazonia

The future flora of Amazonia will include significant areas of secondary forest as degraded pastures are abandoned and secondary succession proceeds. The rate at which secondary forests regain carbon (C) stocks and re-establish biogeochemical cycles that resemble those of primary forests will influence the biogeochemistry of the region. Most studies have focused on the effects of deforestation on biogeochemical cycles. In this study, we present data on the recuperation of carbon stocks and carbon fluxes within a secondary forest of the eastern Amazon, and we compare these measurements to those for primary forest, degraded pasture, and productive pasture. Along a transect from a 23-y-old degraded pasture, through a 7-y-old secondary forest, through a 16-year-old secondary forest, and to a primary forest, the δ¹³C values of soil organic matter (SOM) in the top 10 cm of soil were – 21.0, – 26.5, – 27.4, and – 27.9‰, respectively, indicating that the isotopic signature of SOM from C3 forest plants was rapidly re-established. The degraded pasture also had significant inputs of C from C3 plants. Radiocarbon data indicated that most of the C in the top 10 cm of soil had been fixed by plants during the last 30 years. Differences in soil C inventory among land use types were small compared to uncertainties in their measurement. Root inputs were nearly identical in primary and secondary forests, and litterfall in the secondary forest was 88% of the litterfall rate of the primary forest. In contrast, the secondary forest had only 17% of the above ground biomass. Because of rapid cycling rates of soil C and rapid recovery of C fluxes to and from the soil, the below ground C cycle in this secondary forest was nearly identical with those of the unaltered primary forest.     

Roles of riparian and secondary forests in maintaining the near-threatened butterfly, Sasakia charonda (Lepidoptera, Nymphalidae), populations in Japan

To clarify the habitat requirements of the near-threatened butterfly, Sasakia charonda (Lepidoptera, Nymphalidae), we studied the distribution pattern of its host trees, Celtis sinensis and Celtis jessoensis, and the utilization patterns of various vegetation types by this butterfly in the Oofukasawa River basin in Hokuto City, Yamanashi Prefecture, central Japan. Two species of host trees, C. sinensis and C. jessoensis (height = 2 m or more) were found in riparian forests on sandbanks (hereinafter, riparian forest), in forest regenerated after landslides on valley walls (landslide tracks), in secondary deciduous forests consisting mainly of Quercus acutissima or Quercus serrata and in forests established at abandoned paddy fields and their periphery, where weeds and shrubs used to be mown frequently to avoid shade on the paddies before their abandonment. This suggests that they are pioneer species, and their distribution and regeneration depend on natural and/or human disturbances. Host trees above 2 m were preferred by larvae, and there were very few such trees in secondary forests. More overwintering larvae occurred in riparian forests than at other sites. The number of S. charonda adults was highest at the edge of riparian forests, and we observed a variety of behaviors such as puddling, chasing and mating there. Although the number of adult butterflies was smaller inside and at the edge of secondary forests than in riparian forests, puddling by males and roosting on the trunk of Q. acutissima or Q. serrata by females were observed more frequently there than in riparian forests. Thus, we conclude that landscapes including both riparian forests with natural disturbance and secondary forests with Quercus trees are necessary to maintain host Celtis trees and S. charonda populations.     

Recovery of species diversity and composition of braconid parasitic wasps after reforestation of degraded grasslands in lowland East Kalimantan

Human-induced disturbances and wildfires can transform areas of tropical rainforest into Imperata-dominated grassland, but it may be possible that recovery of biodiversity is facilitated by reforestation with fast-growing trees. We compared the assemblages of braconid wasps as parasitoids of taxonomically diverse groups of insects among Imperata grasslands, young and mature plantations of Acacia mangium, young secondary forests after wildfires, and old secondary forests in the lowland of East Kalimantan. The abundance and species richness of braconids, which had declined in Imperata grasslands, somewhat increased in Acacia plantations, and also the species composition of braconids in Acacia plantations was transitional between Imperata grasslands and old secondary forests. Parasitoids of detritivores and wood borers increased markedly after plantation, while those of herbivores showed a distinct turnover of species all over the range from grasslands to old secondary forests. The plantation of A. mangium had most likely facilitated the recovery of the diversity of host forest and their parasitoids, but the recovery was just at the rudimentary stage even in mature plantations. Monitoring of parasitic wasps would be useful to test the continuous recovery of forest biodiversity in plantation stands.     

Modelling riparian forest distribution and composition to entire river networks

Aim: Developing a methodology to map the distribution of riparian forests to entire river networks and determining the main environmental factors controlling their spatial patterns. Location: Cantabrian region, northern Spain. Methods: We mapped the riparian forests at a physiognomic and phytosociological level by delimiting riparian zones and generating vegetation distribution models based on remote sensing data (Landsat 8 OLI and LiDAR PNOA). We built virtual watersheds to define a spatial framework where the catchment environmental information can be specified for each river reach, in combination with the vegetation map. In order to determine the drivers that play a significant role in the observed spatial patterns in riparian forests, based on our data sets we modelled interactions between environmental information and riparian vegetation by using the Random Forest algorithm. Results: The modelling results obtained reliably reproduced the variation of riparian forest structure and composition across Cantabrian watersheds. The produced maps were highly accurate, with a more than 70% overall accuracy for forest occurrence. A clear differentiation between Eurosiberian (habitats 91E0 and 9160) and Mediterranean (92E0) riparian forests was shown on both sides of the mountain range. Topography and land use were the main drivers defining the distribution of riparian forest as a physiognomic unit. In turn, altitude, climate and percentage of pasture were the most relevant factors determining their composition (phytosociological approach). Conclusions: Our study confirms that anthropic control ultimately defines the distribution of vegetation in the riparian area at a regional to local scale. Human disturbances constrain the extension of forest patches across their potential distribution defined by topoclimatic boundaries, which establish a clear limit between Mediterranean and Eurosiberian biogeographical regions.     

Different Population Dynamics of Microcebus murinus in Primary and Secondary Deciduous Dry Forests of Madagascar

The goal of this study was to identify causes for lower population densities of mouse lemurs (Microcebus murinus) in secondary than in primary dry deciduous forests of western Madagascar. Variations in the abundance of Microcebus murinus are linked to the capacity to enter energy-saving torpor during the dry season. Under natural conditions in primary forest, Microcebus murinus can maintain daily torpor (and possibly hibernation) as long as body temperatures remain below 28°C. Females are more likely to hibernate than males, resulting in skewed sex ratios of captured Microcebus murinus in the primary forest. In the secondary forest the sex ratio of subjects captured during the dry season is more balanced than in primary forest, indicating that fewer females go into torpor in secondary than in primary forest. Secondary forests have fewer large standing or fallen trees that might provide holes as shelter for Microcebus murinus. Ambient temperatures are higher in secondary than in primary forests and higher outside than inside tree holes. These high ambient temperatures might hinder the ability of Microcebus murinus to maintain torpor for prolonged periods in secondary forests. Mouse lemurs from secondary forest have lower body mass than in primary forest. The year-to-year recapture rate is zero in secondary forest and thus significantly lower than in primary forest. This indicates that survival rates are lower in secondary than in primary forests. Thus, secondary forests may be of limited value as buffer zones or even corridors for mouse lemurs.     

Effects of forest-floor avalanche disturbance on the structure and dynamics of a subalpine forest near the forest limit on Mt. Fuji

In subalpine forests near the forest limit on Mt. Fuji in central Japan, slush avalanches occasionally destroy forest-floor vegetation through an influx of volcanic gravel from bare upper sites. The vegetation structure of Larix kaempferi–Abies veitchii forests near and distant from avalanche paths was investigated to determine the effects of forest-floor disturbance on successional processes. The Larix population in a forest near an avalanche path, where there had been signs of forest-floor disturbance by avalanches, had a discontinuous age structure with three age groups, indicating that Larix seedlings established under the Larix canopy after the forest-floor disturbance. In contrast, the Larix population in a forest distant from avalanche paths, where there had been no forest-floor disturbance, had a continuous age structure, with no plants younger than the 90-year-old trees, indicating that this population had established on bare ground over a long period. These data suggest that the primary requirement for the regeneration of L. kaempferi on the forest floor is a mineral substrate. Conversely, A. veitchii had a continuous age structure in both forests. However, forest-floor disturbances by avalanche(s) may exclude A. veitchii from the forest because A. veitchii is very sensitive to scoria deposition. In conclusion, forest-floor disturbance by avalanche(s) provides L. kaempferi with an opportunity to establish on the forest floor, resulting in the maintenance of Larix forests alongside avalanche paths in the upper subalpine area on Mt. Fuji.     

Parasite loss and introduced species: a comparison of the parasites of the Puerto Rican tree frog, (<Emphasis Type="Italic">Eleutherodactylus coqui</Emphasis>), in its native and introduced ranges

The Puerto Rican frog, Eleutherodactylus coqui has invaded Hawaii and reached densities far exceeding those in their native range. One possible explanation for the success of E. coqui in its introduced range is that it lost its co-evolved parasites in the process of the invasion. We compared the parasites of E. coqui in its native versus introduced range. We collected parasite data on 160 individual coqui frogs collected during January-April 2006 from eight populations in Puerto Rico and Hawaii. Puerto Rican coqui frogs had higher species richness of parasites than Hawaiian coqui frogs. Parasite prevalence and intensity were significantly higher in Hawaii, however this was likely a product of the life history of the dominant parasite and its minimal harm to the host. This suggests that the scarcity of parasites may be a factor contributing to the success of Eleutherodactylus coqui in Hawaii.     

Influence of Forest Types and Effects of Forestry Activities on Species Richness and Composition of Chrysomelidae in the Central Mountainous Region of Japan

Species richness and composition of the Chrysomelidae (Coleoptera) were studied in larch (Larix kaempheri [Lamb.] Carrière) plantations, secondary forests, and primary forests. In addition, the effects of forest management practices, such as thinning and long rotation, were examined in the larch plantation. The species richness of Chrysomelidae was higher in the larch plantation than in the secondary forest or in the primary forest. Among the larch plantations, the species richness in old-aged plantations was higher than that in middle-aged plantations. The composition of the beetle assemblages in the larch plantation differed from that in the secondary forest or in the primary forest. Exosoma akkoae (Chujo), Batophila acutangula Heikertinger, and Calomicrus nobyi Chujo were caught with a bias toward the larch plantation. Longitarsus succineus (Foudras) and Sphaeroderma tarsatum Baly were caught more in the secondary forest and the primary forest, respectively. More B. acutangula and S. tarsatum were caught in stands where their host plants occurred at higher rates. Species richness of understory plants was an important factor for chrysomlid species richness, and frequency of host occurrence affected the number of individuals of leaf beetles examined. It seems that forest types and forest management practices affect host plants as well as Chrysomelidae, and that these effects on the host plants also influence chrysomelid assemblages.     

Effects of past logging and grazing on understory plant communities in a montane Colorado forest

Throughout Pinus ponderosa–Pseudotsuga menziesii forests of the southern Colorado Front Range, USA, intense logging and domestic grazing began at the time of Euro-American settlement in the late 1800s and continued until the early 1900s. We investigated the long-term impacts of these settlement-era activities on understory plant communities by comparing understory composition at a historically logged and grazed site to that of an environmentally similar site which was protected from past use. We found that species richness and cover within functional groups rarely differed between sites in either upland or riparian areas. Multivariate analyses revealed little difference in species composition between sites on uplands, though compositional differences were apparent in riparian zones. Our findings suggest that settlement-era logging and grazing have had only minor long-term impacts on understories of upland Front Range P. ponderosa–P. menziesii forests, though they have had a greater long-term influence on riparian understories, where these activities were likely the most intense. This article was written and prepared by US Government employees on official time, and therefore it is in the public domain and not subject to copyright.     

Neighborhood interactions influencing tree population dynamics in nonpyrogenous boreal forest in northern Finland

Open-canopy moss-rich woodlands dominated by Picea abies and Betula pubescens in northern Finland may undergo cyclic development with reciprocal replacement of the tree species due to the positive feedbacks on soil conditions. Although relations to the abiotic environment are well understood, intra- and interspecific interactions during development of sparse boreal forests have received less attention. We studied tree regeneration, growth and survival with respect to size and density of neighboring trees in four stands representing roughly four stages of the Picea–Betula forest cycle. We conducted spatial analysis (Ripley’s K-function) of mapped locations of live and dead stems to reconstruct the distribution of stems prior to mortality, and to infer possible causes of tree population decline. The prevalence of standing dead stems enabled us to test if mortality was associated with density and size of neighboring trees. Logistic regression was used to test for relationships between tree survival and local crowding indices. We also examined spatial autocorrelations of individual size characteristics to determine the mode and spatial extent of tree interactions. Picea abies had reduced recruitment in open areas occupied by mosses and dwarf-shrubs, and preferentially regenerated near B. pubescens, whereas B. pubescens formed small clumps (and occasionally these consisted entirely of stems from a single tree) that showed local repulsion from large P. abies trees. Size of neighboring trees was the primary determinant of individual growth and survival, whereas neighborhood density per se had only a weak effect. Picea abies had negatively correlated sizes among close neighbors (0–4 m radius) indicating that dominant trees suppress their smaller neighbors. Negative autocorrelations prevailed at the transition stages where the patches of smaller trees were concentrated around evenly spaced large trees. Tree sizes became spatially independent at the mature phase. We hypothesize that both low light and soil nutrient availability causes the P. abies population decline. Dominant trees in this high latitude forest have large light depletion zones and shallow root system to promote strong above- and below-ground competition with younger trees. Higher mortality rates within canopy patches were not compensated for by recruitment in gaps, causing P. abies population to decline steadily.     

Depth distribution and composition of seed banks under different tree layers in a managed temperate forest ecosystem

In the present work we examined the composition and distribution across three soil layers of the buried soil seed bank under three different overstory types (Fagus sylvatica, Quercus robur, Pinus sylvestris) and in logging areas in a 4383-ha forest in central Belgium. The objectives were: (1) to investigate whether species composition and species richness of soil seed banks are affected by different forest stands; (2) to examine how abundant are habitat-specific forest species in seed banks under different planted tree layers. The study was carried out in stands which are replicated, managed in the same way (even-aged high forest), and growing on the same soil type with the same land-use history. In the investigated area, the seed bank did show significant differences under oak, beech, pine and in logging areas, respectively in terms of size, composition and depth occurrence. All species and layers taken together, the seed bank size ranked as follows: oakwood > beechwood > logging area > pinewood. The same pattern was found for forest species. Seed numbers of Betula pendula, Calluna vulgaris, Dryopteris dilatata and Rubus fruticosus were significantly higher under the beech canopy. Carex remota, Impatiens parviflora and Lotus sp. showed a significantly denser seed bank in logging areas, while Digitalis purpurea seeds were significantly more abundant in soils under the oak canopy. The fact that the seed bank of an originally homogeneous forest varies under different planted stands highlights that a long period of canopy conversion can affect the composition and depth of buried seeds.     

Recovery of mycorrhizas of a fungus, <Emphasis Type="Italic">Cenococcum geophilum</Emphasis>, after urea treatment in warm temperate forests in Japan

We studied recovery of the ectomycorrhizal species Cenococcum geophilum (Cg) after urea treatment in two types of vegetation. The recovery was as quick as 6 months after the treatment and overlapped the fruiting period of ectomycorrhizal ammonia fungi. Ectomycorrhizas investigated as Cg belonged to a single species irrespective of treatment and vegetation type. Cg ectomycorrhizas were significantly more abundant in density in nontreated soil than in treated soil. Between vegetation types, Cg ectomycorrhizas were significantly more in density in broad-leaved deciduous forest than in broad-leaved evergreen forest. Moreover, Cg was dominant in the nontreated soils of the former type of forest.     

Influence of urbanization on riparian forest diversity and structure in the Georgia Piedmont, US

Riparian forests are increasingly threatened by urban expansion and land use change worldwide. This study examined the relationships between landscape characteristics and woody plant diversity, structure, and composition of small order riparian corridors along an urban-rural land use gradient in the Georgia Piedmont, US. Riparian plant diversity, structure, and composition were related to landscape metrics and land use. Species richness was negatively associated with impervious surfaces and landscape diversity, and positively associated with forest cover and largest forest patch index. Shannon species diversity was strongly related to the biomass of non-native species, especially for the regeneration layer. Urban sites were characterized by high richness of non-native and pioneer species. Developing sites were dominated by the non-native shrub, Ligustrum sinense Lour., and several native overstory trees, mainly Acer negundo L. While agricultural and managed forest sites were composed of ubiquitous species, unmanaged forest sites had a structurally distinct midstory indicative of reduced disturbance. Urban and agricultural land uses showed decreased native stem densities and signs of overstory tree regeneration failure. Results from this study highlight the impact of the surrounding landscape matrix upon riparian forest plant diversity and structure.     

Comparing forest structure and biodiversity on private and public land: secondary tropical dry forests in Costa Rica

Secondary forests constitute a substantial proportion of tropical forestlands. These forests occur on both public and private lands and different underlying environmental variables and management regimes may affect post‐abandonment successional processes and resultant forest structure and biodiversity. We examined whether differences in ownership led to differences in forest structure, tree diversity, and tree species composition across a gradient of soil fertility and forest age. We collected soil samples and surveyed all trees in 82 public and 66 private 0.1‐ha forest plots arrayed across forest age and soil gradients in Guanacaste, Costa Rica. We found that soil fertility appeared to drive the spatial structure of public vs. private ownership; public conservation lands appeared to be non‐randomly located on areas of lower soil fertility. On private lands, areas of crops/pasture appeared to be non‐randomly located on higher soil fertility areas while forests occupied areas of lower soil fertility. We found that forest structure and tree species diversity did not differ significantly between public and private ownership. However, public and private forests differed in tree species composition: 11 percent were more prevalent in public forest and 7 percent were more prevalent in private forest. Swietenia macrophylla, Cedrela odorata, and Astronium graveolens were more prevalent in public forests likely because public forests provide stronger protection for these highly prized timber species. Guazuma ulmifolia was the most abundant tree in private forests likely because this species is widely consumed and dispersed by cattle. Furthermore, some compositional differences appear to result from soil fertility differences due to non‐random placement of public and private land holdings with respect to soil fertility. Land ownership creates a distinctive species composition signature that is likely the result of differences in soil fertility and management between the ownership types. Both biophysical and social variables should be considered to advance understanding of tropical secondary forest structure and biodiversity.     

Assessing the extent and diversity of riparian ecosystems in Sonora, Mexico

Conservation of forested riparian ecosystems is of international concern. Relatively little is known of the structure, composition, diversity, and extent of riparian ecosystems in Mexico. We used high- and low-resolution satellite imagery from 2000 to 2006, and ground-based sampling in 2006, to assess the spatial pattern, extent, and woody plant composition of riparian forests across a range of spatial scales for the state of Sonora, Mexico. For all 3rd and higher order streams, river bottomlands with riparian forests occupied a total area of 2,301 km². Where forested bottomlands remained, on average, 34% of the area had been converted to agriculture while 39% remained forested. We estimated that the total area of riparian forest along the principal streams was 897 km². Including fencerow trees, the total forested riparian area was 944 km², or 0.5% of the total land area of Sonora. Ground-based sampling of woody riparian vegetation consisted of 92, 50 m radius circular plots. About 79 woody plant species were noted. The most important tree species, based on cover and frequency, were willow species Salix spp. (primarily S. goodingii and S. bonplandiana), mesquite species Prosopis spp. (primarily P. velutina), and Fremont cottonwood Populus fremontii. Woody riparian taxa at the reach scale showed a trend of increasing diversity from north to south within Sonora. Species richness was greatest in the willow-bald cypress Taxodium distichum var. mexicanum—Mexican cottonwood P. mexicana subsp. dimorphia ecosystem. The non-native tamarisk Tamarix spp. was rare, occurring at just three study reaches. Relatively natural stream flow patterns and fluvial disturbance regimes likely limit its establishment and spread.     

新疆叶尔羌河流域胡杨林时空格局特征

叶尔羌河流域胡杨林对保障我国新疆南疆地区生态安全有着重要的功能,到目前对该区胡杨林时空格局特征规律的研究极少。基于1995—2015年Landsat TM/OLI长时间序列遥感监测数据,通过土地利用程度变化指数、转移矩阵以及景观指数对叶尔羌河流域胡杨林时空格局变化进行动态监测与分析。结果表明:(1)胡杨林面积从1995年的1916.15 km~2减少到2015年的1652.25 km~2,面积减少了263.90 km~2。永安坝至夏河林场段,耕地向河道扩张,胡杨林转为耕地;而夏河林场至三河口段,水资源匮乏,胡杨林被迫向河道收缩,转为未利用地。(2)胡杨林动态度和空间动态度降低,而胡杨林变化状态指数、斑块密度、最大斑块指数和散布与并列指数增加,斑块结合度指数高,胡杨林空间分布逐渐集中、破碎度降低、连通性增加。流域景观要素连接性逐渐减弱,破碎化程度较高,优势度斑块类型比例逐渐下降,景观类型向复杂化方向发展。(3)1995—2005年胡杨林受气候变化和人类活动的干扰,胡杨林面积减少较快,景观格局变化剧烈,而2005—2015年胡杨林保护措施的实施、区域产业结构的调整以及生态安全输水区域的扩大等,胡杨林面积减少速度减缓,但形势仍较为严峻。在叶尔羌河流域合理利用水资源,统筹协调生活、生产和生态用水、强化水资源管理力度,以及调整产业结构、控制人口增长速度等是该区胡杨林生态系统恢复的有效手段和途径。研究可以为叶尔羌河流域生态系统恢复以及胡杨林保护提供参考。     

Vegetation dynamics in the Bangladesh Sundarbans mangroves: a review of forest inventories

The Sundarbans, the largest contiguous mangrove forest in the world, covers 6,017 km² of the coastal zone of Bangladesh. Heritiera fomes Buch.-Ham., Excoecaria agallocha L. and Ceriops decandra (Griff.) Ding Hou jointly cover 95% of the forest area. In this study, the results of four forest inventories have been analyzed to understand observed vegetation dynamics of the Sundarbans from 1926 to 1997. The diversity of forest types has been gradually reduced over time, but H. fomes and E. agallocha have maintained their dominance over large portions of the forest. H. fomes is spread over 67% of the vegetated area of the forest, concentrated mostly in the northeastern area, and constitutes 75% of the density of trees with >15 cm dbh. The distribution and stem density of H. fomes show negative relationships with that of E. agallocha and C. decandra. In terms of coverage, E. agallocha is the most common species, spread over 74% of the vegetated area of the forest, and constitutes 39% of the density of trees with >2.5 cm dbh. On a longer timeframe (1926–1997), the dominance (coverage and density of larger diameter trees) of H. fomes as well as that of E. agallocha is declining. Even on parameters such as density of trees with >15 cm dbh, the dominance of E. agallocha is declining at a much greater rate than H. fomes. This observation contradicts the successional schemes proposed by different authors. This might indicate that theorizing successional schemes based on short-term observations on vegetation dynamics is not sufficient. The effect of human interference, changes in hydroedaphic condition and species interaction should be taken into consideration during explaining observed vegetation dynamics. Moreover, the need to understand vegetation trajectories at the micro-scale should be emphasized.     

Distribution and size of capercaillie leks in relation to old forest fragmentation

Summary Distribution and size of 38 capercaillie Tetrao urogallus leks were related to amount and configuration of old forest patches in two south-east Norwegian coniferous forests. The smallest occupied patch was 48 ha containing a solitary displaying cock. All patches larger than 1 km² contained leks. Number of cocks per lek increased with increasing patch size. Number of leks per patch increased in a step-wise manner with one lek added for each 2.5–3 km² increase in patch size. In large patches there was one lek per 3–5 km² old forest, and density of lekking cocks was 2–2.5 per km². In small patches density of cocks varied considerably. Density of cocks was not related to patch isolation or patch shape. However, among leks surrounded by 50–60% old forest within a 1 km radius, number of cocks increased with increasing old forest fine-graininess. We argue that when old forests cover more than 50%, a fine-grained mosaic may support higher densities of lekking cocks than a coarse-grained mosaic. Conversely, when old forests cover less than 50%, a fine-grained mosaic is unfavourable, because each old forest patch becomes too small and isolated. Finally, we present a predictive model of how old forest fragmentation influences density of leks, number of cocks per lek, and total density of cocks.