Synchrony in small mammal populations of montane forest patches in southern India

1. Small mammal populations were studied in montane evergreen forests in the Nilgiris, southern India, using live-trapping from January 1994 to September 1996. Two sites were selected, one with a single large forest patch and adjoining plantations, and the other with several small forest patches separated by grassland.
2. Nine species were recorded, of which eight were trapped in the forest patches, two in the grasslands and five in plantations. Rattus rattus was the most abundant species (2–36 individuals ha⁻¹) followed by Suncus montanus (0–11 individuals ha⁻¹). Densities of seven other species recorded were very low.
3. Synchrony in various population variables – density, biomass, mean weight, density of adults and adult females, and proportion of adults, adult females and sub-adults – was examined for Rattus rattus populations in the forest patches and plantations. Density and biomass were studied in seven other species recorded at these sites. Spearman's rank order correlation was used as a measure of synchrony between the population variables.
4. Within-site synchrony was higher than between-site synchrony in population characters. Synchrony was also higher between plots within the unfragmented site than they were between plots in the fragmented site. Relatively high synchrony in proportions of adults, adult females and juveniles in the forest patches implied that breeding is probably influenced by climate and food availability, which are seasonal in this habitat.
5. Given the small patch sizes (≈1–10 ha) and low population sizes, asynchrony is likely to be an outcome of demographic and environmental stochasticity, and low dispersal rates may impede establishment of synchrony.     

Composition,structure and diversity of cove forest stands in the Great Smoky Mountains: a patch dynamics perspective

Abstract. Cove forests of the Great Smoky Mountains are North American examples of old-growth temperate forest. Ecological attributes of seven stands were studied using one 0.6 - 1.0 ha plot per stand. Stand basal area (39 - 55 m²/ha) and biomass (326 - 471 Mg/ha) were high for temperate deciduous forest. Density ranged from 577 to 1075 stems/ha. All stands had a mixture of deciduous canopy species. Only rarely did a single species comprise more than half of the stand by density, basal area or biomass. Shade-intolerant species were present at low levels (1 - 5 % of total stand density). A wide range of stem diameters was characteristic of most species. However, some species lacked small stems, indicating discontinuous regeneration. Stands tended to have 10 - 20 tree species per ha and at least five species had biomass levels > 10 Mg/ha, indicating high evenness. Canopy gaps covered 10 % of the total area (2 - 21 % by stand). Gaps and conspecific patches of canopy trees > 0.05 ha in size were infrequent. Spatial analyses revealed a variety of patterns among species at inter-tree distances of 1 to 25 m. When all species were combined, juveniles showed aggregation, and adults were often hyperdispersed. Analyses for individual species confirmed that the mosaic of canopy species is influenced by non-random spatial processes. Adults of several species were aggregated at distances > 10 m. Juveniles of all major species exhibited aggregation. Several species exhibited regeneration near conspecific adults. This pattern suggested limited mobility for such species within the shifting mosaic. A diverse patchwork resulted despite the fact that many species did not exhibit segregation of adults and juveniles. Further understanding of patch dynamics and the potential for compositional steady state in cove forests requires long-term study with spatial data.     

Patch size of forest openings and arthropod populations

Summary Five sizes of canopy openings (0.016 ha to 10 ha) were established in the Southern Appalachian Mountains in early 1982 to examine the initial patterns of plant and arthropod establishment across a size range of forest disturbances. Vegetation standing crop after the first growing season was considerably higher in large than small openings in apparent response to greater resource release (e.g., sunlight) in larger openings. Woody stump and root sprouts were the dominant mode of revegetation in each patch size. Forest dominants such as Quercus rubra, Q. prinus and Carya spp. were less important as sprouters in openings than several minor forest components (e.g., Robinia pseudo-acacia, Acer rubrum, Halesia carolina and Cornus florida). Arthropod abundance and community composition varied across the size range of forest openings. Arthropods from the surrounding forest readily utilized the smallest canopy openings (0.016 ha). All feeding guilds were well represented in these small openings and herbivore biomass and load (mg of herbivores/g of foliage) were much higher than in larger patches. In contrast, arthropod abundance and species richness were significantly lower in mid-size than smaller patches. The relatively sparse cover and high sunlight in mid-size openings may have promoted surface heat buildups or soil surface/litter moisture deficits which restricted arthropod entry from the surrounding forest. Arthropod abundance and species richness were higher in large than mid-size patches. The greater vegetation cover in larger openings may have minimized the deleterious effects on arthropod populations. However, the absence of population increases among these arthropod species maintained herbivore loads at very low levels in large patches. Our results suggest that arthropod abundance and diversity in sprout-dominated forest openings are highly dependent on the extent of environmental differences between patch and surrounding forest.     

Gap size and succession in cutover southern Appalachian forests: an 18 year study of vegetation dynamics

We used clearcut logging in establishing four replicated sizes of canopy openings (0.016, 0.08, 0.4, and 2.0 ha) in a southern Appalachian hardwood forest in 1981 to examine the long-term effects of disturbance size on plant community structure, biomass accumulation, aboveground net primary productivity (NPP), and mode of recovery. The reestablishment of NPP and biomass following logging was 6–7-fold greater in large than small openings by 17 years. Total biomass in the 2.0 ha openings (127.3 Mg ha⁻¹) recovered 59.5% as NPP (19.7 Mg ha⁻¹ yr⁻¹) reached 225% of precut forest levels. Biomass accumulation was 2.6–3.6-fold greater in interior than edge locations of all but the 0.016 ha gaps. The absence of significant patch size or edge vs. interior differences in tree densities suggests that growth rates of individual trees were enhanced in more insolated microenvironments. Sprouting (86–95% of tree NPP) was much more important than advance regeneration (4–10%) or seedling germination (<2%) during early recovery in all opening sizes. Canopy dominant Quercus and Carya trees exhibited limited sprouting following disturbance. Instead, shade-intolerant Robinia pseudoacacia and Liriodendron tulipifera were major sprouters that used N-fixation (Robinia) and rapid growth (Liriodendron) in attaining 7.4 and 5.9 fold greater biomass accumulation, respectively in 2.0 ha than 0.016 ha opening sizes. Seedling germination and understory production were extensive in all openings following logging, but declined rapidly as the young tree canopy began closing by 4–6 years. The relative importance of shade-intolerant tree biomass approximately doubled over 17 years as shade-tolerant tree seedlings, herbs, and shrubs gradually regained importance under the emerging canopy. Sprouting caused the persistence of a tree species composition in all openings that remained relatively similar to the precut forest. Large disturbances on mountain slopes of the southern Appalachians generally promote sprouting and rapid recovery, whereas small disturbances in low-elevation cove forests lead to a gradual recovery through seedling germination and/or advance regeneration. Continued logging in the southern Appalachians will increase the relative size and frequency of large disturbances, further the importance of sprouting of shade-intolerant species, and lead to more even-aged forest stands throughout the region.     

Population density and area: the role of between- and within-patch processes

The relationship between population density and the size of host plant patches was investigated for the red milkweed beetle Tetraopestetraophthalmus inhabiting unmanipulated patches of Asclepias syriaca. The resource concentration hypothesis proposes that density-area patterns, specifically that of increasing herbivore density with patch size, are primarily a function of movement between host plant patches. This research investigated the degree to which movement accounted for density-area patterns. Poisson regression analysis of beetle abundance versus milkweed patch size revealed that beetle density tended to increase with patch size. The pattern of density and patch size resulted from local reproduction and residence time. The density of emerging beetles tended to increase with patch size while emigration rates were unrelated to patch size. Immigration rates were constant with patch size for male beetles, and decreased with patch size for female beetles. Net flux of beetles (immigration – emigration) did not vary with patch size for male beetles and decreased with patch size for female beetles. Comparisons are made between this system and previously studied systems where movement plays a significant role in forming density area patterns. Additionally, several hypotheses are presented which may account for greater in situ recruitment and residence time in large patches. Received: 23 February 1996 / Accepted: 8 January 1997     

Importance of habitat patch size for occupancy and density of aspen-associated saproxylic beetles

In Fennoscandian boreal forests, aspen (Populus tremula) is one of the most important tree species for biodiversity. In this study we explore how occupancy and density of beetles associated with dead aspen are related to habitat patch size and connectedness in a 45,000 ha boreal managed forest landscape in central Sweden. Patch size was estimated as amount of breeding substrate and connectedness as crown cover of living aspen in the surrounding landscape. The beetles were sampled by sieving of bark or by inspection of species-characteristic galleries in 56 patches with dead aspen. Six of nine aspen-associated species (Xylotrechus rusticus, Ptilinus fuscus, Mycetophagus fulvicollis, Cyphaea curtula, Homalota plana and Endomychus coccineus) showed a positive significant relationship between habitat patch size and occupancy. For all these species, except C. curtula, there was also a significant positive relationship between patch size and density. Connectedness was not retained as a significant variable in the analyses. Species not defined as aspen-associated constituted a significantly larger proportion of the total density of individuals of saproxylic beetles in smaller habitat patches than in larger patches. Richness of aspen-associated species was positively related to habitat patch size. Efforts in the managed forest should be directed towards preserving and creating larger patches of living and dead aspen trees and increasing the amount of aspen at the landscape level.     

Evidence of a Minimum Patch Size Threshold of Reproductive Success in an Endangered Songbird

ABSTRACT That area-sensitive songbirds breed only in relatively large patches suggests that there may be a minimum patch size threshold in which they will breed, even when controlling for the total amount of habitat in the landscape. We searched for minimum patch size thresholds of presence, territory establishment by males, pairing success, and reproductive success for 2 migratory songbirds that differed in sensitivity to patch size: golden-cheeked warblers (Dendroica chrysoparia) and white-eyed vireos (Vireo griseus). We assessed 2 potential limiting factors: brown-headed cowbird (Molothrus ater) parasitism and arthropod biomass (food resource). We determined whether either factor was related to patch size and compared measurements of each above and below the observed thresholds. We monitored 24 golden-cheeked warbler and 47 white-eyed vireo territories in 12 patches. We found evidence of a minimum patch size threshold (between 15.0 ha and 20.1 ha) of reproductive success for golden-cheeked warblers, but not for white-eyed vireos. We found no minimum patch size thresholds for presence, territory establishment by males, or pair formation for either species. Conservation practices based on thresholds of presence, territory establishment, or pair formation might not address issues of reproduction for golden-cheeked warblers. We failed to find evidence that cowbird parasitism or arthropod biomass were limiting factors. The ability to identify patch size thresholds of reproductive success for target species could be useful in conservation and management for setting goals for retention and restoration of target species’ habitat patch size.     

Forest Fragmentation Modifies Habitat Quality for Alouatta palliata

Fragmentation reduces habitat area, increases the number of habitat patches, decreases their size, and increases patch isolation. For arboreal mammals such as howlers (Alouatta palliata), canopy modifications from fragmentation processes could also negatively affect habitat quality. We analyzed changes in the composition and plant structure of 15 fragments (1–76 ha) and compared them with vegetation from a continuous tropical rain forest reserve (700 ha) in Los Tuxtlas, Mexico. At each site, we sampled 1000 m² of all trees, shrubs, and lianas with a diameter at breast height (DBH) ≥10 cm. We obtained estimates of species richness, density, and basal area for different ecological groups, DBH ranges, and top food resources for howlers. We used a stepwise multiple regression analysis to determine relationships between fragment characteristics (size, shape index, and isolation) and plant variables. Compared to continuous forest, fragments have altered composition and plant structure, with large trees absent from the canopy. The basal area of top food resources is higher in continuous forest. Fragment size is the best explanation for the differences in composition and plant structure. The largest fragments had greater basal area of top food resources and more large primary trees in the canopy. Overall, our results suggest that fragmentation altered the habitat quality for howlers.     

Regeneration in subalpine coniferous forests

The regeneration process of a subalpine coniferous forest, a mixed forest ofTsuga diversifolia (dominant species),Abies veitchii, Abies mariessi, andPicea jezoensis var.hondoensis, was studied on the basis of annual ring data. The age class distribution was discontinuous and four age groups occurred in the study plot (30m×30m). The canopy layer was a mosaic of patches (83.8–133.7 m² patch area), which had different mean ages. The recruitment of canopy trees was carried out only by advance regeneration in the plot. The diameter growth ofAbies andPicea exceeded diameter growth ofTsuga in the gap.Abies lived for 200–300 years and their trunks were susceptible to heart rot.Picea lived for 300–400 years andTsuga for more than 400 years. The regeneration process derived from the analysis of the plot consisted of three phases leading to the development of a even-aged patch; (1) the establishment of saplings before a gap opening, (2) the opening of a gap in the canopy and repair of the canopy by advance regenerated saplings dominated by rapid growth species,Abies andPicea, and (3) the dying off of canopy trees as each species reached the end of its life-span, resulting in pure patches of long-livedTsuga.     

Scales of aboveground and below-ground competition in a semi-arid woodland detected from spatial pattern

Abstract. Semi-arid woodlands are two-phase mosaics of canopy and inter-canopy patches. We hypothesized that both aboveground competition (within canopy patches), and below-ground competition (between canopy patches), would be important structuring processes in these communities. We investigated the spatial pattern of trees in a Pinus edulis-Juniperus monosperma woodland in New Mexico using Ripley's K-function. We found strong aggregation of trees at scales of 2 to 4 m, which indicates the scale of canopy patches. Canopy patches were composed of individuals of both species. Crown centers of both species were always less aggregated than stem centers at scales less than canopy patch size, indicating morphological plasticity of competing crowns. In the smallest size classes of both species, aggregation was most intense, and occurred over a larger range of scales; aggregation decreased with increasing size as is consistent with density-dependent mortality from intraspecific competition. Within canopy patches, younger trees were associated with older trees of the other species. At scales larger than canopy patches, younger trees showed repulsion from older conspecifics, indicating below-ground competition. Hence, intraspecific competition was stronger than interspecific competition, probably because the species differ in rooting depth. Woodland dynamics depend on the scale and composition of canopy patches, aggregated seed deposition and facilitation, above- and below-ground competition, and temporal changes in the spatial scale of interactions. This woodland is intermediate in a grassland-forest continuum (a gradient of increasing woody canopy cover) and hence we expected, and were able to detect, the effects of both above- and below-ground competition.     

Evaluating Residual Tree Patches as Stepping Stones and Short-Term Refugia for Red-Legged Frogs

Abstract: Temperate pond-breeding amphibians are vulnerable to forest fragmentation because they must access upland terrestrial sites during the nonbreeding season but are prone to desiccation in hot, dry environments without canopy cover. Harvesting techniques that retain live trees in the cut block are advocated for sustaining forest biodiversity, but the effects of these practices on amphibians are unknown. We studied red-legged frogs (Rana aurora) in movement trials to assess: 1) how short-term use of residual trees was affected by tree patch size, streams, and neighborhood features; 2) whether residual tree patches were used as stepping stones in negotiating cut blocks; 3) the effects of patch size and patch proximity in altering movement paths; and 4) the effects of retention level and patch size on interpatch distance. Residual tree patches were potentially valuable short-term refugia but their value was size dependent. Virtually all frogs released at the base of single trees or inside small tree clusters left within 72 hours, but the proportion leaving decreased curvilinearly with increasing patch size. Frogs were less likely to leave tree patches with a running stream or where neighborhood stream density was high. Residual tree patches did not systematically alter movement paths. Frogs intercepted residual tree patches mostly at random and had to be within 5-20 m of a tree patch before moving to it in greater proportions than expected by chance. However, amphibian movements were biased toward large (0.8 ha) patches and away from small (0.3 ha) patches 50 m away. Our results indicated that residual trees should not be retained singly but should be aggregated in groups between 0.8 ha and 1.5 ha, preferably in stream locations.     

Effects of patch size and basal area on avian taxonomic and functional diversity in pine forests: Implication for the influence of habitat quality on the species–area relationship

Relationships between avian diversity and habitat area are assumed to be positive; however, often little attention has given to how these relationships can be influenced by the habitat structure or quality. In addition, other components of biodiversity, such as functional diversity, are often overlooked in assessing habitat patch value. In the Sandhills Ecoregion of Georgia, USA, we investigated the relationship between avian species richness and functional diversity, forest basal area, and patch size in pine forests using basal area as a surrogate for overstory structure which in turn impacts vegetation structure and determines habitat quality within a patch. We conducted bird surveys in planted mature pine stands, during breeding season of 2011. We used three classes of stand basal area (BA): OS, overstocked (BA ≥ 23 m²/ha); FS, fully/densely stocked (13.8 m²/ha ≤ BA < 23 m²/ha); and MS, moderately stocked (2.3 m²/ha ≤ BA < 13.8 m²/ha). MS patches showed more structural diversity due to higher herbaceous vegetation cover than other two pine stocking classes of patches. Total species richness and functional richness increased with the size of MS patches, whereas functional divergence decreased with the size of OS patches (p < 0.05). Functional richness tended to be lower than expected as the size of OS patches increased. Greater richness of pine–grassland species was also found at MS patches. Percent cover of MS patches within a landscape influenced positively the richness of pine–grassland species (p < 0.05). Our results suggest that (a) avian species–habitat area relationship can be affected by habitat quality (structural diversity) and varies depending on diversity indices considered, and (b) it is important to maintain moderate or low levels of pine basal area and to preserve large‐sized patches of the level of basal area to enhance both taxonomic and functional diversity in managed pine forests.     

Competition, coexistence, and adaptation amongst rodent invaders to Pacific and New Zealand islands

Variation in skull size was investigated for three species of rats (kiore –Rattus exulans Peale; ship rat –R. rattus L.; Norway rat –R. norvegicus Berkenhout) which were introduced by humans to various islands in New Zealand and other Pacific islands. Data from seventy-one islands and 882 specimens are examined for evidence of the effects of latitude, island size and interspecific competition among rats and the house mouse (Mus musculus L.) on skull size, using multiple regressions. For R. exulans, skull size increases with latitude as predicted by Bergmann's rule, but no such effect occurs for the other two rats. There was a positive relationship between island size and the number of species inhabiting it, and some species combinations were more likely to occur than others. For example, R. exulans and R. norvegicus were more likely to occur together, while R. rattus and R. exulans were rarely sympatric. R. exulans and R. rattus skull size was negatively correlated with the number of other rodents on the same island. R. exulans skull size increased on smaller islands in some island groups, perhaps because increased density and consequent increased intraspecific competition on smaller islands favours increased body size. This effect is more pronounced in tropical islands (Solomon islands), than in subtropical ones (Hawaiian islands) and less so in temperate New Zealand. Collectively the data demonstrate that rapid evolution of body size in predictable directions can follow within 150 years of the introduction of species to new receiving communities.     

Space invaders? A search for patterns underlying the coexistence of alien black rats and Galápagos rice rats

The introduction and spread of the black rat Rattus rattus is believed to have caused the worst decline of any vertebrate taxon in Galápagos. However, the “extinct” Santiago rice rat Nesoryzomys swarthi has recently been rediscovered in sympatry with R. rattus providing the first exception to this general pattern of displacement. We carried out an exploratory investigation of this novel system with the aim of identifying patterns that may facilitate the apparent coexistence of the two species. We carried out an extensive survey of Santiago Island to map the current distribution of the endemic rice rat and to explore broad scale distribution–habitat associations. We then used live-trapping, radio-tracking, and spool-and-line tracking to quantify abundance–habitat correlations and to test for evidence of interspecific spatial segregation, alteration of N. swarthi activity patterns (spatial and temporal), and microhabitat partitioning. We found that N. swarthi has disappeared from part of its historical range and appears to be restricted to a 14 km stretch of the north-central coast, characterised by high density of the cactus Opuntia galapageia. In contrast, the generalist R. rattus was found at all survey sites. We found no evidence of spatial segregation, and home range size, temporal activity and density of N. swarthi did not vary with local density of R. rattus. However, pre-dawn and post-dusk N. swarthi activity levels increased with R. rattus density perhaps reflecting an increase in foraging effort necessary to compensate for the costs of interspecific exploitation or interference competition. The distribution, microhabitat selection, and abundance–habitat relations of N. swarthi suggest that the endemic cactus O. galapageia may facilitate interspecific coexistence. Further research should include a comparison of inter-seasonal resource preference and foraging activity of the two species coupled with replicated field experiments to confirm and quantify competition and to elucidate the mechanism of competitive coexistence.     

Foraging for nutrients, responses to changes in light, and competition in tropical deciduous tree seedlings

We evaluated (1) the responses of two co-occurring tropical tree species, Heliocarpuspallidus and Caesalpiniaeriostachys, to changes in light, (2) the ability of these species to search for and exploit a fertilized soil patch, (3) the relationship between the capacity to forage for a fertilized patch and the capacity to respond to changes in light availability and (4) how the relationship between light and nutrient acquisition influenced the competitive interactions between these species. Plants of the two species were exposed to a factorial combination of high (H) and low (L) light intensity and fertilized (+Fp) and unfertilized (−Fp) nutrient patches for 50 days. Half of the plants from H were then transferred to L (HL treatment), and half of the plants from L were transferred to H (LH). The remaining plants were kept in their original light condition and grown for another 50 days. Plants were grown in these light and patch treatments alone (one plant per pot) and in interspecific competition (one plant per species resulting in two plants per pot). Both species exploited fertilized patches by increasing root biomass and length in the patch. This enhanced plant productivity and growth rate mainly under LH and HH conditions for Heliocarpus and the HH condition for Caesalpinia). When plants in the HH light environment were grown with an unfertilized patch, plant biomass and relative growth rates (RGRs) were even lower than␣under the LL light environment [(HH–Fp)<LL]. However, the combined activity of shoot and roots when above- and below-ground resources were temporally and spatially heterogeneous influenced plant productivity and growth rate. The benefit from light increase (LH) was reduced when grown with an unfertilized patch. Larger reductions in root biomass, length and density in the patch, and in plant biomass and RGR, were exhibited by Heliocarpus than by Caesalpinia. These results suggest a close relationship between root foraging and light capture, where the benefit of the exploitation of the patch will be reflected in whole-plant benefit, if enough light is captured above-ground. In addition, the results suggest a change in the expected plant responses to light due to heterogeneity in soil nutrients, even though the fertilized patch was only a small proportion of the total soil volume. Leaf characteristics such as specific leaf area responded only to light conditions and not to patchily distributed nutrients. Root characteristics responded more strongly to nutrient heterogeneity. Competition modified the pattern of foraging under both high- and low-light conditions in Heliocarpus by 50 days, and the ability to forage for a fertilized patch under LL after 100 days of growth for Caesalpinia. Even though plant growth and productivity are greatly reduced under low-light conditions (HL and LL), competition modifies the ability of species to forage for a rich patch (especially for the fast-growing species Heliocarpus). Received: 24 November 1997 / Accepted: 15 June 1998     

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.     

Habitat use by endemic and introduced rodents along a gradient of forest disturbance in Madagascar

We used logistic and Poisson regression models to determine factors of forest and landscape structure that influence the presence and abundance of rodent species in the rain forest of Ranomafana National Park in southeastern Madagascar. Rodents were collected using live-traps along a gradient of human disturbance. All five endemic rodent species (Nesomys rufus, N. audeberti, Eliurus tanala, E. minor and E. webbi) and the introduced rat Rattus rattus were captured in both secondary and primary forests, but the introduced Mus musculus was only trapped in secondary forest. The abundance of R. rattus increased with the level of habitat disturbance, and it was most common in the heavily logged secondary forest. Furthermore, the probability of the presence of R. rattus increased with decreasing distance from forest edge and decreasing canopy cover, while the probability of presence increased with increasing herbaceous cover, altitude and overstory tree height. The species was never observed farther than 500 m away from human habitation or camp-site. N. rufus prefered selectively-logged forest at altitudes above 900 m a.s.l. Its probability of presence increased with increasing canopy cover, herbaceous cover and distance from forest edge, and with decreasing density of fallen logs, overstory tree height and distance from human habitation. N. audeberti prefered heavily-logged areas, while E. tanala was the only species occurring along the entire range of forest disturbance. We suggest that in the Ranomafana National Park the spread of R. rattus is associated with deforestation.     

Gap formation and regeneration of tropical mangrove forests in Ranong,Thailand

Tropical mangrove forests are characterized by clear zonation along a tidal gradient, and it has been supposed that the zonation is primarily controlled by soil factors. However, effects of disturbance on mangrove forests are still not well understood and may play an important role on the vegetation patterns and forest dynamics in some forest formations. In this study, the pattern of disturbance regime and its effects on regeneration of tropical mangrove forests along a tidal gradient were investigated in Ranong, Thailand. We established one or two 0.5 ha plots in four vegetation zones, i.e. Sonneratia alba–Avicennia alba zone, Rhizophora apiculata zone, Ra – Bruguiera gymnorrhiza zone, Ceriops tagal–Xylocarpus spp. zone. Gap size (percentage gap area to total study area and individual gap size) was the largest in Sa–Aa zone which is located on the most seaward fringe, and it declined from seaward to inland. Canopy trees of S. alba and A. alba had stunted trunks and showed low tree density. On the contrary, canopy dominants in the other three inland zones, e.g. R. apiculata, B. gymnorrhiza, and Xylocarpus spp., had slender trunks and showed high tree density. Accordingly, differences in disturbance regime among the four zones were resulted from the forest structural features of each zone. Disturbance regime matched with regeneration strategies of canopy dominants. Seedlings and saplings of S. alba and A. alba, which need sunny condition for their growth, were abundant in gaps than in understorey. By contrast, R. apiculata, B. gymnorrhiza, and Xylocarpus spp., which can tolerate less light than S. alba and A. alba, had greater seedling and sapling density under closed canopy than gaps. Many large gaps may enhance the abundance of S. alba and A. alba in Sa–Aa zone, and a few small gaps may prevent the light demanding species to establish and grow in the other inland zones. Correspondence of disturbance regime and regeneration strategies (e.g. light requirement) of canopy dominants may contribute to the maintenance of the present species composition in each of the vegetation zones.     

Interactive effects of grazing and shrubs on the annual plant community in semi‐arid Mediterranean shrublands

Question: We studied the interactive effects of grazing and dwarf shrub cover on the structure of a highly diverse annual plant community. Location: Mediterranean, semi‐arid shrubland in the Northern Negev desert, Israel. Methods: Variation in the biomass and plant density of annual species in the shrub and open patches was monitored during four years, inside and outside exclosures protected from sheep grazing, in two contrasting topographic sites: north and south‐facing slopes that differed in their dominant dwarf shrubs species: Sarcopoterium spinosus and Corydothymus capitatus, respectively. Results: Above‐ground biomass, density and richness of annual species were lower under the canopy of both shrub species compared to the adjacent open patches in the absence of grazing. Grazing reduced the biomass of annuals in open patches of both topographic sites, but not in the shrub patches. On the north‐facing slope, grazing also reduced plant density and richness in the open patches, but increased plant density in the shrub patches. At the species level, various response patterns to the combined effects of grazing and patch type were exhibited by different annuals. Protection against the direct impacts of grazing by shrub cover as well as species‐specific interactions between shrubs and annuals were observed. A conceptual mechanistic model explaining these interactions is proposed. Conclusion: In semi‐arid Mediterranean shrublands grazing and dwarf shrub cover interact in shaping the structure of the annual plant community through (1) direct impacts of grazing restricted to the open patches, (2) species‐specific facilitation/ interference occurring in the shrub patches and (3) subsequent further processes occurring among the interconnected shrub and open patches mediated through variation in seed flows between patches.     

Patch occupancy in the endangered butterfly Lycaena helle in a fragmented landscape: effects of habitat quality, patch size and isolation

While there is agreement that both habitat quality and habitat network characteristics (such as patch size and isolation) contribute to the occupancy of patches by any given species, the relative importance of these factors is under debate. This issue is of fundamental ecological importance, and moreover of special concern for conservation biologists aiming at preserving endangered species. Against this background we investigated patch occupancy in the violet copper Lycaena helle, one of the rarest butterfly species in Central Europe, in the Westerwald area (Rhineland-Palatinate, Western Germany). Occupied (n = 102) differed from vacant (n = 128) patches in altitude, size, connectivity, availability of wind shelter, in the abundance of the larval host-plant, in the abundance of a grass species indicating favorable habitat conditions and in the abundance of nitrophilous plants. Overall, patch occupancy was primarily determined by patch size, connectivity and the abundance of the larval host plant, while all other parameters of habitat quality were of subordinate importance. Therefore, our findings suggest that even for extremely sedentary species such as L. helle habitat networks are decisive and—next to the preservation of habitat quality—need to be an integral part of any conservation management for this species.